abstract - Department of Botany - Smithsonian Institution

SMITHSONIAN INSTITUTION 

Contributions from the United States National Herbarium 

Volume 52: 1-415 

Monocotyledons and Gymnosperms of 

Puerto Rico and the Virgin Islands 

Editors 

Pedro Acevedo-Rodríguez 

and 

Mark T. Strong 

Department of Botany 

National Museum of Natural History 

Washington, DC 

2005

ABSTRACT 

Acevedo-Rodríguez, Pedro and Mark T. Strong. Monocots and Gymnosperms of Puerto Rico and the 

Virgin Islands. Contributions from the United States National Herbarium, volume 52: 415 pages 

(including 65 figures). The present treatment constitutes an updated revision for the monocotyledon and gymnosperm flora 

(excluding Orchidaceae and Poaceae) for the biogeographical region of Puerto Rico (including all islets and islands) and the Virgin 

Islands. With this contribution, we fill the last major gap in the flora of this region, since the dicotyledons have been previously 

revised. This volume recognizes 33 families, 118 genera, and 349 species of Monocots (excluding the Orchidaceae and Poaceae) and 

three families, three genera, and six species of gymnosperms. The Poaceae with an estimated 89 genera and 265 species, will be 

published in a separate volume at a later date. When Ackerman’s (1995) treatment of orchids (65 genera and 145 species) and the 

Poaceae are added to our account of monocots, the new total rises to 35 families, 272 genera and 759 species. The differences in 

number from Britton’s and Wilson’s (1926) treatment is attributed to changes in families, generic and species concepts, recent 

introductions, naturalization of introduced species and cultivars, exclusion of cultivated plants, misdeterminations, and discoveries 

of new taxa or new distributional records during the last seven decades. Seven new taxa are described: Agave minor Proctor, 

Bulbostylis capillaris subsp. insulana M.T. Strong, Cyperus pulguerensis M.T. Strong, Guzmania lingulata var. concolor Proctor 

& Cedeño-Mald., Pitcairnia angustifolia var. simplicior Proctor & Cedeño-Mald., Pitcairnia jareckii Proctor & Cedeño-Mald., and 

Zephyranthes proctorii Acev.-Rodr. & M.T. Strong; and two new combinations are made: Cyperus subtenuis (Kük.) M.T. Strong 

and Rhynchospora divaricata (Ham.) M.T. Strong. Twenty one of the accepted names (or their basionyms) in this work are 

lectotypified: Aechmea lingulata (L.) Baker, Aechmea nudicaulis (L.) Griseb., Alocasia plumbea Van Houtte, Crocosmia × 

crocosmiiflora (Lemoine ex Morren) N.E. Brown, Cyperus confertus Sw., Cyperus subtenuis (Kük.) M.T. Strong, Cyperus urbanii 

Boeck., Eleocharis rostellata (Torr.) Torr., Guzmania monostachia (L.) Mez, Heteranthera limosa (Sw.) Willd., Heteranthera 

reniformis Ruiz & Pav., Hohenbergia antillana Mez., Neomarica northiana (Schneev.) Sprague, Rajania cordata L., 

Rhynchospora ciliata (G. Mey.) Kük., Rhynchospora odorata C. Wright ex Griseb., Spathiphyllum wallisii Regel, Tillandsia 

fasciculata Sw., Tillandsia polystachya (L.)L., and Xanthosoma helleborifolium (Jacq.) Schott. Four names are neotypified: 

Caladium bicolor (Aiton) Vent., Eleutherine bulbosa (Mill.) Urb., Smilax coriacea Spreng., and Spirodela polyrhiza (L.) Schleid. 

One name is epitypified: Anthurium scandens (Aubl.) Engl. Fifteen synonyms of accepted taxa are lectotypified: Amaryllis 

equestris Aiton, Bulbostylis curassavica var. pallescens Kük. & Ekman ex Urb., Bulbostylis ekmanii Kük., Bulbostylis papillosa 

Kük., Crinum amabile Donn ex Ker-Gawl, Cyperus ehrenbergii Kunth, Cyperus filiformis var. densiceps Kük., Eleocharis 

sintenisii Boeck., Pistia occidentalis Blume, Pitcairnia ramosa J. Jacq., Psilocarya rufa Nees, Rhynchospora longispicata Boeck., 

Scleria microcarpa var. subeciliata C.B.Clarke, Scleria grisebachii C.B. Clarke, and Xanthosoma atrovirens K. Koch & Bouché. 

Thirty-two of the taxa treated are newly reported for the flora area: Aechmea fasciata (Lindl.) Baker, Agave vivipara L., Bulbostylis 

stenocarpa Kük., Calathea crotalifera S. Watson, Carex longii Mack., Commelina rufipes var. glabrata (D.R. Hunt) Faden & D.R. 

Hunt, Costus malortieanus H. Wendl., Costus scaber Ruiz & Pav., Cyperus floridanus Britton ex Small, Cyperus papyrus L., 

Cyperus squarrosus L., Cyperus tenuis Sw., Cyperus unifolius Boeck., Eleocharis debilis Kunth, Eleocharis minutiflora Boeck., 

Eleocharis oligantha C.B. Clarke, Fuirena robusta Kunth, Furcraea stratiotes J. B. Petersen, Halophila engelmannii Asch., 

Heliconia rostrata Ruiz & Pav., Hydrocleys nymphoides (Willd.) Buchenau, Kyllinga nemoralis (J.R. & G. Forst.) Dandy ex 

Hutchinson & Dalz., Maranta gibba Sm., Rhynchospora domingensis Urb., Rhynchospora fascicularis subsp. fascicularis var. 

distans (Michx.) Chapm., Rhynchospora radicans subsp. microcephala (Bertero ex Spreng.) W.W. Thomas, Ruppia didyma Sw., 

Scleria brittonii Core, Scleria havanensis Britton, Scleria verticillata Muhl. ex Willd., Spathiphyllum wallisii Regel., and Xyris 

curtissii Malme. 

KEY WORDS: Botany, Caribbean, Greater Antilles, Greater Puerto Rico, Gymnosperms, Monocots, Puerto Rico,Virgin Islands. 

RESUMEN 

Acevedo-Rodríguez, Pedro y Mark T. Strong. Monocots and Gymnosperms of Puerto Rico and the Virgin 

Islands. Contributions from the United States National Herbarium, volume 52: 415 páginas (incluyendo 

65 figuras). El presente tratado constituye una revisión actualizada para los monocotiledóneos y los gimnospermos (excluyendo 

Orchidaceae y Poaceae) de la región biogeográfica de Puerto Rico (incluyendo cayos e islotes) y las Islas Vírgenes. En este volumen, 

se reconocen 33 familias, 118 géneros y 349 especies de monocotiledóneos y tres familias, tres géneros y seis especies de 

gimnospermos. Las Poaceas con un estimado de 89 géneros y 265 especies será publicada en un próximo volumen de esta serie. 

Cuando el numero (65 géneros y 145 especies) de Orquídeas (Ackerman, 1995) y el de Poaceas es tomado en consideración, el 

numero total de monocotiledóneas asciende a 35 familias, 272 géneros y 759 especies. La diferencia en el numero de especies entre 

el presente tratado y el de Britton y Wilson (1926) es atribuido a los cambios de conceptos de familias, géneros y especies, recientes 

introducciones, naturalización de especies anteriormente introducidas, la exclusión de plantas cultivadas, identificaciones erróneas 

y el descubrimiento de nuevos taxones o de nuevos registros de distribución en las ultimas 7 décadas. Los siguientes siete nuevos 

taxones son aquí descritos: Agave minor Proctor, Bulbostylis capillaris subsp. insulana M.T. Strong, Cyperus pulguerensis M.T. 

Strong, Guzmania lingulata var. concolor Proctor & Cedeño-Mald., Pitcairnia angustifolia var. simplicior Proctor & Cedeño-

Mald., Pitcairnia jareckii Proctor & Cedeño-Mald. y Zephyranthes proctorii Acev.-Rodr. & M.T. Strong, y dos combinaciones 

nuevas efectuadas: Cyperus subtenuis (Kük.) M.T. Strong y Rhynchospora divaricata (Ham.) M.T. Strong. Los siguientes 21 

nombres ( o sus basónimos) aquí aceptados son lectotipificados: Aechmea lingulata (L.) Baker, Aechmea nudicaulis (L.) Griseb., 

Alocasia plumbea Van Houtte, Crocosmia × crocosmiiflora (Lemoine ex Morren) N.E. Brown, Cyperus confertus Sw., Cyperus 

subtenuis (Kük.) M.T. Strong, Cyperus urbanii Boeck., Eleocharis rostellata (Torr.) Torr., Guzmania monostachia (L.) Mez, 

Heteranthera limosa (Sw.) Willd., Heteranthera reniformis Ruiz & Pav., Hohenbergia antillana Mez., Neomarica northiana 

(Schneev.) Sprague, Rajania cordata L., Rhynchospora ciliata (G. Mey.) Kük., Rhynchospora odorata C. Wright ex Griseb., 

Spathiphyllum wallisii Regel, Tillandsia fasciculata Sw., Tillandsia polystachya (L.) L., y Xanthosoma helleborifolium (Jacq.) 

Schott. Cuatro nombres son neotipificados: Caladium bicolor (Aiton) Vent., Eleutherine bulbosa (Mill.) Urb., Smilax coriacea 

Spreng., y Spirodela polyrhiza (L.) Schleid. Un nombre es epitipificado: Anthurium scandens (Aubl.) Engl.. Los siguientes quince 

sinónimos son lectotificados: Amaryllis equestris Aiton, Bulbostylis curassavica var. pallescens Kük. & Ekman ex Urb., 

Bulbostylis ekmanii Kük., Bulbostylis papillosa Kük., Crinum amabile Donn ex Ker-Gawl, Cyperus ehrenbergii Kunth, Cyperus 

filiformis var. densiceps Kük.,Eleocharis sintenisii Boeck., Pistia occidentalis Blume, Pitcairnia ramosa J. Jacq., Psilocarya rufa 

Nees, Rhynchospora longispicata Boeck., Scleria microcarpa var. subeciliata C.B. Clarke, Scleria grisebachii C.B. Clarke y 

Xanthosoma atrovirens K. Koch & Bouché. Los siguientes 32 taxones son nuevos registros para el área cubierta en esta flora: 

Aechmea fasciata (Lindl.) Baker, Agave vivipara L., Bulbostylis stenocarpa Kük., Calathea crotalifera S. Watson, Carex longii 

Mack., Commelina rufipes var. glabrata (D.R. Hunt) Faden & D.R. Hunt, Costus malortieanus H. Wendl., Costus scaber Ruiz & 

Pav., Cyperus floridanus Britton ex Small, Cyperus papyrus L., Cyperus squarrosus L., Cyperus tenuis Sw., Cyperus unifolius 

Boeck., Eleocharis debilis Kunth, Eleocharis minutiflora Boeck., Eleocharis oligantha C.B. Clarke, Fuirena robusta Kunth, 

Furcraea stratiotes J. Boye-Petersen, Halophila engelmannii Asch., Heliconia rostrata Ruiz & Pav., Hydrocleys nymphoides 

(Willd.) Buchenau, Kyllinga nemoralis (J.R. & G. Forst.) Dandy ex Hutchinson & Dalz., Maranta gibba Sm., Rhynchospora 

domingensis Urb., Rhynchospora fascicularis subsp. fascicularis var. distans (Michx.) Chapm., Rhynchospora radicans subsp. 

microcephala (Bertero ex Spreng.) W.W. Thomas, Ruppia didyma Sw., Scleria brittonii Core, Scleria havanensis Britton, Scleria 

verticillata Muhl. ex Willd., Spathiphyllum wallisii Regel. y Xyris curtissii Malme. 

PALABRAS CLAVES: Antillas Mayores, Banco de Puerto Rico, Botánica, Caribe, Gimnospermos, Islas Vírgenes, 

Monocotiledóneos y Puerto Rico. 

DATE OF PUBLICATION: 30 November 2005 

Cover Design by Alice Tangerini based on illustrations by Bobbi Angell. Front cover: Crinum 

zeylanicum and Hymenocallis latifolia. Back cover: Scleria canescens, S. scindens, and S. secans. 

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

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

7012, USA. 

POSTMASTER: Send address changes to Contributions from the U.S. National Herbarium, 

Department of Botany, National Museum of Natural History, MRC-166, P.O. Box 37012, 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. 

The periodical, Contributions from the United States National Herbarium, was first published in 

1890 by The United States Department of Agriculture. From 1 July 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 (2000) as a venue for publishing longer taxonomic 

papers, checklists, floras, and monographs, produced by the staff and associates at the U.S. National 

Herbarium. It is externally peer reviewed, and published at irregular intervals. Subscription and 

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

Natural History, MRC-166, Smithsonian Institution, Washington, DC, 20013-7012, USA e-mail: 

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previous volumes, please contact the editor, Paul M. Peterson (peterson@si.edu).

FAMILY TREATMENTS 

CONTRIBUTORS 

Acevedo-Rodríguez, P. Smithsonian Institution, Department of Botany, MRC-166, PO Box 

37012, Washington, DC 20013-7012 

Cedeño-Maldonado, J.A. United States Army Corps of Engineers, Antilles Regulatory 

Section, 400 Fernández Juncos Ave., San Juan, Puerto Rico 00901-3299 

Lewis, P. University of the West Indies, Department of Life Sciences, Herbarium 

Mona, Kingston 7, Jamaica. 

Nicolson, D. H. Smithsonian Institution, Department of Botany, MRC-166, POBox 37012, 

Washington, DC 20013-7012 

Proctor, G.R. University of the West Indies, Department of Life Sciences, Herbarium 

Mona, Kingston 7, Jamaica. 

Strong, M.T. Smithsonian Institution, Department of Botany, MRC-166, PO Box 37012, 

Washington, DC 20013-7012 

BOTANICAL ILLUSTRATIONS 

Angell, B. PO Box 158, Marlboro, VT 05344. Figs. 1-10, 13-20, 23, 25, 27, 29-32, 34, 36, 

43, 45, 36, 48, 50, 54, 56. 

Litwak, T. 10652 Wayridge Dr., Montgomery Village, MD 20886. Figs. 11, 12, 21, 22, 24, 

28, 33, 35, 37, 44, 47, 51-53, 55, 57. 

Tangerini, A. Smithsonian Institution, Department of Botany, MRC-166, PO Box 37012, 

Washington, DC 20013-7012. Figs. 26, 49.

CONTENTS 

FAMILY SEQUENCE ................................................................................................. 6 

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

Brief History of the Project ................................................................................ 9 

Methodology ..................................................................................................... 9 

Acknowledgements .......................................................................................... 9 

FLORISTIC TREATMENT ....................................................................................... 11 

GYMNOSPERMS .......................................................................................... 11 

MONOCOTYLEDONS .................................................................................. 18 

BIBLIOGRAPHY .................................................................................................... 387 

INDEX TO SCIENTIFIC NAMES ........................................................................... 390 

INDEX TO COMMON NAMES ............................................................................. 405

6 

1. Podocarpaceae 

2. Pinaceae 

3. Zamiaceae 

1. Araceae 

2. Lemnaceae 

3. Limnocharitaceae 

4. Alismataceae 

5. Hydrocharitaceae 

6. Cymodoceaceae 

7. Ruppiaceae 

8. Potamogetonaceae 

9. Smilacaceae 

10. Dioscoreaceae 

11. Burmanniaceae 

12. Hypoxydaceae 

13. Iridaceae 

14. Amaryllidaceae 

15. Agavaceae 

16. Asparagaceae 

17. Dracaenaceae 

Monocots and Gymnosperms of Puerto Rico and Virgin Islands 

FAMILY SEQUENCE 

GYMNOSPERMS 

MONOCOTYLEDONS 

18. Asphodelaceae 

19. Arecaceae (Palmae) 

20. Commelinaceae 

21. Haemodoraceae 

22. Pontederiaceae 

23. Musaceae 

24. Heliconiaceae 

25. Zingiberaceae 

26. Costaceae 

27. Cannaceae 

28. Maranthaceae 

29. Bromeliaceae 

30. Xyridaceae 

31. Juncaceae 

32. Cyperaceae 

33. Thyphaceae

Monocots and Gymnosperms of Puerto Rico and Virgin Islands 7 

Monocotyledons and Gymnosperms of Puerto Rico 

and the Virgin Islands 

Editors: Pedro Acevedo-Rodríguez¹ and Mark T. Strong¹ 

The Botany of Porto Rico and the Virgin 

Islands by Britton and P. Wilson (1923-26) treated 

27 families, 220 genera and 503 species of 

monocots plus two families, two genera and four 

species of gymnosperms. Modern revisions (this 

volume, Ackerman, 1995, and Peterson et al. in 

prep.) of the biogeographical Puerto Rican flora 

recognize 35 families, 272 genera and 759 species 

of monocots, and three families, three genera, and 

six species of gymnosperms. The differences in 

number is attributed to changes in families, 

generic and species concepts, recent introductions, 

naturalization of introduced species and cultivars, 

exclusion of cultivated plants, misdeterminations, 

and discoveries of new taxa or new distributional 

records during the last four decades. 

Botanical research in Puerto Rico and the 

Virgin Islands has lagged behind that carried out 

on other islands of the Caribbean and in the New 

World. In 1753 when Linnaeus published his 

Species Plantarum describing approximately 

9,000 plant species of the world, only two species 

were reported for Puerto Rico and one for the 

island of St. Croix. These numbers contrast with 

those from Jamaica with 135 species, the Lesser 

Antilles (including Curaçao and Bonaire) with 75, 

Hispaniola with 20, and Cuba with seven species 

reported. It wasn’t until the latter part of the 18 th 

century that botanical explorations started in 

earnest in Puerto Rico and the Virgin Islands. The 

first significant botanical expedition carried out on 

Puerto Rican territory was by F. Bredermeyer 

(1758-1839), part of an Austrian expedition lead 

by F. J. Maerter (1753-1827), whose collections 

INTRODUCTION 

are deposited in the Museum of Natural History of 

Vienna (W). In the following years many more 

botanists visited Puerto Rico or the Virgin Islands; 

their collections form the basis for the description 

of new genera or species as part of an ongoing 

effort to inventory the plants of the world. 

The first attempt to write a floristic treatment 

for any of the islands belonging to our region was 

started at the beginning of the 19 th century. From 

1828-31, Schlechtendal (1794-1866) published 

the first florula for the island of St. Thomas, 

treating 360 species. This effort was followed 

with a paper by Krebs in 1847, who contributed 

additional species for the flora of St. Thomas. In 

1879, von Eggers (1844-1903) published a florula 

of St. Croix, listing 1,028 species for an island that 

is only slightly larger than St. Thomas. A few 

years later, the first partial florula for Puerto Rico 

was published by D. Bello y Espinosa (1817-1884) 

from 1880 to 1883, listing 964 species of plants, 

some of which were new to science. His work was 

essentially an annotated checklist, based mostly on 

Bello’s herbarium specimens that were later 

destroyed by insects. Bello had collaborated with 

L. Krug (1833-1898), an amateur naturalist and 

German consul stationed in Mayagüez, but the 

checklist was published only with Bello’s 

authorship. Krug produced three unpublished 

manuscripts (destroyed by a fire in the library of 

the Botanical Museum in Berlin): a treatise on 

vernacular names of Puerto Rican plants, a catalog 

of West Indian plants, and most importantly, a 

volume illustrating some common Puerto Rican 

plants. 

1 Smithsonian Institution, Department of Botany, MRC-166, PO Box 37012, Washington, DC, 20013-7012 

acevedop@si.edu 

strongm@si.edu

8 


The first serious attempt to write a fullfledged 

flora of Puerto Rico came from A. Stahl 

(1842-1917), who from 1883-88 published six 

fascicles of various groups of flowering plants. His 

work was never finished as his treatment of the 

monocotyledons and approximately 720 

watercolors of Puerto Rican plants were never 

published. Although Stahl based his descriptions 

chiefly on live plants, he also collected 

approximately 1,330 dried specimens (with 

several duplicates) dating from 1882 to 1889. His 

collections were not cataloged in chronological 

sequence and it is therefore difficult to interpret his 

itinerary or whether he preserved specimens while 

writing his flora. The first set of his collections 

went to Berlin (Krug and Urban herbarium) and 

were destroyed during World War II. Duplicates 

of his collections are known to exist at various 

institutions as follows: B(6), BR(1), C(9), G(1), 

GH(1), GOET(6), L(7), M(1), NY(1), S(130), 

TO(2), and US(48). Stahl’s collections are 

important because they constitute early records for 

many native and introduced plants in Puerto Rico, 

some of which have never been recollected. His 

collections were the basis for numerous studies, 

some of them resulting in taxa new to science. 

Stahl described several new species himself (most 

of which are considered synonyms by modern 

botanists), but further study is necessary to assess 

whether his collections or his illustrations 

(watercolors) constitutes a type. 

Around the time when Stahl’s flora was being 

published, Paul Sintenis (1847-1907), a German 

botanist, arrived in Puerto Rico and collected on 

the island from 1884 to 1887. Sintenis was the 

most prolific collector in early Puerto Rican 

botany, with approximately 8,000 collections. His 

collections are very important because they 

contained large sets of duplicates that were 

distributed to numerous herbaria of the world, thus 

facilitating the study of Puerto Rican plants by a 

wide range of specialists. Many of his collections 

were designated as types of taxa new to science. 

His collections are equally important as early 

records of introduced plants and as a representation 

of the floristic content of an area from that time. It 

is very likely that Sintenis and Stahl collected 

together, as can be deduced from their collections 

which often represent the same species collected at 

the same site and on the same date. Sintenis’ first 

set of Puerto Rican plants went to Berlin (Krug and 

Urban Herbarium) but were largely destroyed 

during World War II. Surviving collections are 

found in numerous herbaria. 

By the close of the 19 th century and beginning 

of the 20 th century, even more botanists were 

actively studying and describing the flora of the 

West Indies. The most memorable and relevant 

ones to the flora of Puerto Rico and the Virgin 

Islands were I. Urban (1848-1931) and N. L. 

Britton (1859-1934). Urban published hundreds 

of scientific papers, many of which dealt with 

West Indian Botany. His Symbolae Antillanae, 

published in nine volumes from 1898 to 1928, still 

constitutes a fundamental and necessary work for 

the study of West Indian plants. Volume 4 (1903- 

11) of this series is dedicated to the flora of Puerto 

Rico, where numerous novelties were described 

either by him or by his numerous contributors. 

This work, although an excellent source of 

botanical bibliography and rigorous nomenclatural 

treatment, did not contain keys that would have 

facilitated its use by the non-expert or local 

botanist. 

From 1923-1926, Britton, then director of the 

New York Botanical Garden, published along with 

P. Wilson (1879-1944) the first full-fledged flora 

for the region of Puerto Rico and the Virgin 

Islands. They treated the flowering plants while 

W. R. Maxon (1877-1948), in volume 6(3) 1926, 

treated the Pteridophytes. Their work, although 

outdated, still constitutes one of the basic 

references for the study of Puerto Rican (and 

Virgin Islands) botany. 

During the decade of the 1960’s, A. Liogier 

started his work on the revision of the Puerto Rican 

flora, reporting numerous novelties as well as new 

records for the island. From 1985 to 1997 he 

published a new dicotyledon flora for Puerto Rico 

in five volumes, but did not cover the Virgin 

Islands in his treatments. In 2000, along with L. F. 

Martorell, Liogier published a revised edition to 

his synopsis for the Flora of Puerto Rico and 

adjacent islands, a checklist that lists all the 

vascular plants that occur on the islands under the 

jurisdiction of Puerto Rico. Their work includes a 

list of the monocots as well, but in the present work 

we have made substantial revisions to their 

nomenclature and species concepts. 

The present treatment constitutes an updated 

revision for the monocot (except for the Poaceae 

and the Orchidaceae) and gymnosperm flora for 

the biogeographical region of Puerto Rico 

(including all islets and islands) and the Virgin


Islands. With this contribution, we fill one of the 

last major gaps in the flora of this region, since 

other groups have been recently revised. The 

dicotyledon flora was treated by H. Alain Liogier 

& Martorell (1982, 2000) and by H. Alain Liogier 

(1985, 1988, 1994, 1995, 1997), the Pteridophytes 

by George R. Proctor (1989), and the orchids by 

James D. Ackerman (1992, 1995). The last 

remaining untreated group is the Poaceae, which 

will be treated by P.M. Peterson and collaborators 

in a separate work to be published in a forthcoming 

volume of this series. We follow the approach of 

Britton and P. Wilson (1923-26) and others (e.g., 

Proctor 1989, Ackerman 1995) in treating these 

islands as a biogeographical region instead of a 

flora dictated by political boundaries. 

BRIEF HISTORY OF THE PROJECT 

This project was started about two decades 

ago by Dr. George R. Proctor while working on his 

treatment for the Ferns of Puerto Rico and the 

Virgin Islands (1989). His position as a plant 

taxonomist at the Department of Natural 

Resources, San Juan, Puerto Rico (DRN) provided 

him with the opportunity to explore the island and 

to document its flora by collecting and preserving 

thousands of specimens. The first set of his Puerto 

Rican collections form the core of the SJ 

herbarium, while duplicates of his collections are 

distributed in several herbaria such as IJ, NY, and 

US. During his stay in Puerto Rico, Dr. Proctor 

collected numerous monocotyledons with the idea 

of producing a treatment for this important group 

of plants. Many of the new records reported and 

new taxa described in this volume are based on his 

collections. In early 1998, after retiring from 

DRN, Dr. Proctor came to the Smithsonian 

Institution as a Visiting Scientist, sponsored by an 

Andrew W. Mellon Foundation Fellowship to 

work on a treatment for the monocots of Puerto 

Rico and the Virgin Islands. Dr. Proctor spent a 

year and a half at our Institution with additional 

support from the U.S. Department of Interior, Fish 

& Wildlife Service. By the end of his visit, he 

produced the basis for the current publication. 

Several other workers contributed treatments to 

this volume, including students and various staff 

of the Department of Botany, Smithsonian. The 

editors of this volume checked the accuracy of 

treatments, sometimes enriching descriptions, 

providing typification information, solving 

nomenclatural problems, and citing the numerous 

collections deposited at SJ, NY, UPR, UPRRP, 

and US. The Gymnosperms were later added in 

order to fill a major gap in the knowledge of the 

flora area. After approximately five years of 

continuous work, we bring this volume to fruition. 

METHODOLOGY 

All taxa fully treated in this work are based on 

a specimen in an herbarium. These are indicated 

by bold face type. Synonyms are indicated by 

italisizing the names of taxa that have been used in 

major treatments for the islands: Urban’s Flora 

Portoricensis (1903, 1905, 1910, 1911) and 

Britton and P. Wilson’s Scientific Survey of 

Puerto Rico and the Virgin Islands (1923, 1924). 

Names misapplied to a species by an author are 

indicated in the sense (sensu) of the author 

followed by the true author of the name and date of 

publication. Names for species that could not be 

verified with a specimen are listed at the end of the 

corresponding genus under the heading: “Excluded 

species.” Major herbaria known to have extensive 

holdings of plants from Puerto Rico and the Virgin 

Islands were consulted to document the occurrence 

and distribution of the taxa treated here. The 

distribution (political divisions) given for each 

species is based on specimen records from the 

following herbaria: GH, GMUF, MAPR, MO, 

NA, NY, SJ, UPR, UPRRP, and US. The selected 

specimens cited for each species are provided as a 

guide to the variability within each species and in 

most cases do not include all of the material 

examined. Herbarium abbreviations within the 

text follow Index Herbariorum (Holmgren et al. 

1990). Abbreviations of author names follows 

Brummitt & Powell (1992). Names of taxa that 

appear in brackets in the keys indicate they are 

cultivated but not naturalized or persistent in 

natural areas. 

ACKNOWLEDGEMENTS 

We thank P. Acevedo Torres for locating 

representative material of Musaceae and Araceae 

cultivars; Franklin Axelrod (UPRRP) for careful 

review of the manuscript especially of name

10 


citations, answering numerous inquiries, and 

facilitating loan of specimens; Ronald Brashear 

and Kirsten van der Veen of the NMAH Dibner 

Library for facilitating the use of the rare books 

collection; James Ackerman (UPRRP) and Bruce 

Hansen (USF) for reviewing and improving the 

manuscript; Richard Carter (VSC) for helping to 

identify a Cyperus collection; Dwight Colón (SJ) 

for searching and providing information on some 

of the specimens housed at SJ; Mia Ehn for data on 

Cyperaceae type collections at C; Amy Farstad 

and John Kartesz (NCU) for information on 

Cyperaceae records for Puerto Rico; Socorro 

González-Elizondo (CIIDIR) for discussions on 

Eleocharis; Gerda Huisman (Universiteitsbibliotheek, 

Groningen) for valuable help in 

locating and providing images of several 

illustrations of Plumier’s Codex Boerhaavianus; 

Charles Jarvis (BM) for answering inquiries 

regarding Linnean names typification; Robert 

Kral (BRIT) for insightful discussions on 

Bulbostylis taxonomy; Alexander Krings (NCSU) 

for translating relevant German text into English; 

Paul and Hiltjie Maas (U) for information on the 

lectotypification of Canna indica; Rachel Newton 

for data on Cyperaceae type photographs at F; Dan 

Nicolson (US) and K. Gandhi (GH) for help in 

solving numerous questions on nomenclature and 

literature citation; Leslie Overstreet and Daria 

Wingreen of the NMNH Cullman Library for 

facilitating the use of numerous rare books; 

Lauren Raz (NY) for discussions on Dioscorea 

taxonomy and Rajania typification; Amneris 

Siaca for transcribing Proctor’s manuscript into 

electronic format; Gerald Smith (High Pt 

University, NC), for discussions on Hymenocallis 

taxonomy; Dennis Stevenson (NY) and Vivian 

Negrón-Ortiz (MU) for discussions regarding 

taxonomy of Zamia; Jeanine Vélez (MAPR) for 

answering inquiries, facilitating loans of material, 

and pulling specimens at NY; John Weirsema 

(USDA) for information regarding conservation 

of economically important monocots; J. Carlos 

Trejo (UPR) for answering numerous inquiries 

concerning collections at UPR; Taina Litwak for 

preparing many of the line drawings; Bobbi Angell 

for authorizing reprinting many of her illustrations 

here published; and Alice Tangerini for preparing 

several line drawings, support on image handling, 

and for designing the cover. We would like to 

thank the following herbaria for making their 

collections accessible either through loans or by 

imaging: BM, C, CGE, GH, JBSD, MAPR, MO, 

NCU, NY, P, SJ, UPR, and UPRRP. We greatfully 

acknowledge financial assistance from the 

Andrew W. Mellon Foundation for providing 

fellowships to George R. Proctor, José A. Cedeño- 

Maldonado, and Patrick Lewis for work at the 

Smithsonian Institution (NMNH); the U. S. Fish 

and Wildlife Service to George R. Proctor for 

additional time required for the completion of this 

project; the International Institute of Tropical 

Forestry for support of field work and 

inventorying of the Puerto Rican plant collections 

housed at US, and the USDA-Forest Service for 

helping with publication costs.


FLORISTIC TREATMENT 

Key to the taxa 

1. Plants woody, usually resinous; perianth absent; stigmas absent; ovules exposed during pollination; 

seeds naked, borne on a fleshy receptacle, or on the surface of cone scales or bracts .... Gymnosperms 

1. Plants herbaceous or woody; perianth usually present; stigmas present; ovules enclosed in ovaries 

during pollination; seeds enclosed in a fruit (capsule, follicle, achene, caryopsis, drupe, or berry) 

.............................................................................................................................................. 2 

2. Leaves usually reticulate-veined, not sheathing basally; vascular bundles usually in rings or 

concentric cylinders; flower perianth parts or segments generally 4 or 5 (rarely 3) or multiples 

thereof; cotyledons 2 ............................................................................. (Dicotyledons) 

2. Leaves usually parallel veined or with mid rib only, often sheathing basally; vascular bundles 

discrete, scattered; flower perianth parts or segments often in whorls of 3, or multiples thereof; 

cotyledons 1 ..................................................................................... Monocotyledons 

GYMNOSPERMS 

Key to the families 

1. Female cones fleshy with 1 or 2-several scales, drupe-like or nut-like .............................................. 2 

2. Female cone lacking a fleshy receptacle at base; fertile cone scales 2-several, often with dorsal 

projections; seeds several per cone .......................................................................... [Cupressaceae] 

2. Female cone with a fleshy receptacle at base; fertile cone scale 1, lacking a dorsal projection; seed 

1 per cone ..............................................................................................................1. Podocarpaceae 

1. Female cones woody with numerous scales, not drupe-like or nut-like ............................................. 3 

3. Leaves simple, needle-like, in fascicles of 2-5 .......................................................... 2. Pinaceae 

3. Leaves pinnate or if simple, not needlelike nor in fascicles ......................................................... 4 

4. Leaves pinnate, the pinnae with a mid rib only, lacking lateral longitudinal veins 

........................................................................................................................... [Cycadaceae] 

4. Leaves simple or pinnate, the leaf blades or pinnae lacking a mid rib, but with many 

longitudinal veins ............................................................................................................... 5 

5. Trees; leaves simple; cones ovoid or subglobose, of spirally-arranged flattened scales 

............................................................................................................ [Araucariaceae] 

5. Fern-like with a subterranean short trunk or caudex; leaves pinnate; cones cylindrical, of 

valvately arranged sporophylls ........................................................ 3. Zamiaceae

12 


Family 1. PODOCARPACEAE Podocarp Family 

Podocarpaceae Endl., Syn. Conif.: 203. 1847, nom. conserv. 

by M. T. Strong & P. Acevedo-Rodríguez 

Evergreen trees or shrubs, slightly resinous, dioecious or rarely monoecious. Trunks generally erect, 

simple, columnar, or sometimes branched from near the base. Leaves simple, usually spirally-arranged, 

sometimes subopposite or opposite and decussate, highly variable in shape, linear, lanceolate, oblong, 

needle-like, or scale-like, generally with a single midvein but sometimes parallel-veined, soft and flexible, 

occasionally leathery. Inflorescence composed of separate, unisexual scaly ament- or catkin-like cones 

(male) and fleshy drupe-like or nut-like cones (female). Male cones cylindrical, solitary or clustered, 

terminal or axillary, sessile or pedunculate, with spirally-arranged scales, each bearing 2 microsporangia 

(pollen sacs); pollen grains bearing 2 (-3) air-bladders, sometimes termed “wings”. Female cones usually 

solitary, terminal or sometimes axillary, with a fleshy receptacle at base; fertile cone scales 1-several, each 

or only one of each bearing one ovule. Mature seeds solitary with a fleshy outer layer or sometimes 

perched on, or partly embedded in a swollen fleshy or leathery receptacle which is often brightly colored. 

A family of 17 genera and approximately 170 species, distributed in tropical and antarctic regions 

worldwide. 

TYPE: Podocarpus L’Hér. ex Pers. 

Reference: Laubenfels, D. J. de. 1986. Coniferales. Pp. 337-453. In: Van Steenis, C. G. G. J. & W. J. 

J. O. De Wilde, (eds.), Fl. Malesiana vol. 10, Noordhoff International Publishing, Leyden, The 

Netherlands. 

1. PODOCARPUS 

Podocarpus L’Hérit. ex Pers., Syn. Pl. 2: 580. 1807, nom. conserv. 

Large evergreen trees or rarely shrubs, usually dioecious. Bark soft, fibrous, yellowish to reddish 

brown, weathering to gray, usually fissured, often peeling in vertical strips. Leaves spirally-arranged, 

bifacially flattened, with a single mid rib. Male cones axillary or sometimes terminal, pedunculate. Female 

cones axillary, on short naked peduncles, ellipsoid-globose, drupe-like with a fleshy or leathery receptacle 

at base, often apiculate at apex. Seeds green, often glaucous when immature. A genus of approximately 

100 species distributed primarily in the southern hemisphere worldwide in tropical or subtropical montane 

forests and antarctic forest zones. 

TYPE: Podocarpus elongatus (W. T. Aiton) L’Hérit. ex Pers. ( ≡ Taxus elongata W. T. Aiton), type 

conserv. 

1. Podocarpus coriaceus Rich. & A. Rich., 

Comm. Bot. Conif. Cycad. 14, pl. 1, fig. 3. 

1826; Nageia coriacea (Rich. & A. Rich.) 

Kuntze, Revis. Gen. Pl. 2: 800. 1891. Type: 

Lesser Antilles, Montserrat. Collector 

unknown. (holotype: probably at P). 

Fig. 58. A 

Shrub or tree to 20 m tall, divaricately 

branching, with narrow to spreading crown. Bark 

smoothish and scaly on new growth, becoming 

fissured and eventually shaggy and peeling off in 

brown or gray strips, the inner bark pink. Buds (at 

branch tips) with long-acuminate scales, these 

sometimes becoming foliaceous. Leaves crowded, 

spreading, linear-lanceolate, straight to subfalcate, 

4-13 (-17) × 0.8-1.6 (-2) cm, stiff and leathery, 

semi-glossy, dark green adaxially, green to 

yellow-green abaxially, long-acuminate or acute 

with acuminate tip, gradually narrowed to base, 

short-petiolate, the midvein prominent abaxially, 

indistinct adaxially, lateral veins not evident. Male 

cones solitary, sessile, cylindrical, (14-) 20-40 × 4- 

6 mm, surrounded at base by several broad obtuse 

scales, yellowish green; microsporophylls sparingly 

lacerulate, the apex obtusely rounded, often 

abruptly narrowed to an acute apiculum. Female 

cones solitary, axillary, peduncles 3-10 mm long,


receptacle red to dark red, fleshy, 2-lobed, 6-10 

mm long; scales 2-3, each bearing a solitary ovule. 

Seeds broadly ovoid or ellipsoid, 7-10 × 5.5-7 mm, 

obtusely or bluntly crested at apex, slightly ridged 

dorsally, light brown to brown. 

General distribution: Puerto Rico, Lesser 

Antilles, Trinidad, and Tobago. 

Distribution in Puerto Rico: In montane 

forests and wooded slopes, 500-1065 m. Recorded 

from Maricao, Río Grande, and San Germán. 

Common names: Puerto Rico: Caoba del país, 

Caobilla; Podocarp. 

Selected specimens examined: PUERTO RICO: 

Maricao: Sargent 509 (US); Sintenis 291(US); Río 

de Maricao, Britton et al. 2462 (US); Monte 

Alegrillo, Britton et al. 2598 (US); Bosque Insular 

de Maricao, Las Tetas de Cerro Gordo, Webster et 

al. 8868 (US); Insular Forest (Monte del Estado), 1 

mi. E of stone observation tower, Stimson 1270 

(US); Maricao State Forest: Bo. Cain Alto, 

Proctor 47174 (US); Bosque Insular de Maricao, 

near Buena Vista, Little 13343 (US); Bosque 

Family 2. PINACEAE Pine Family 

Pinaceae Lindl., Intr. Nat. Syst. Bot., ed. 2: 313. 1836, nom. conserv. 


Insular de Maricao, trail from Vivero de Peces 

southward, Little 13363 (US); Maricao State 

Forest, W side of Rd. 120, km 14.1, at lookout 

tower, Proctor & McKenzie 43831 (US). Río 

Grande: Trail to El Yunque, Luquillo Mts, Little 

21609 (US); Rt. 191, from Luquillo Experimental 

Forest, El Yunque peak, Pfeifer & class 2779, 

2780 (US); Sierra de Luquillo, El Yunque, Hartley 

13347 (US); El Yunque, Sargent 13343 (US) 

Sierra de Luquillo, El Yunque, Little 13558 (US); 

Caribbean National Forest, El Yunque, Axelrod & 

Lindgren 6438 (US). San Germán: Bo. Minillas, 

Maricao Forest Reserve, ca. 2 km down disused 

Rt. 362 (S of Rt. 120), Axelrod & Potter 9589 

(US). 

Cultivated Species 

Podocarpus macrophyllus var. maki Endl., a 

native of China, has been planted as an ornamental 

in Puerto Rico, but is not recorded as being 

naturalized there. 

Evergreen or deciduous trees or sometimes shrubs, resinous and aromatic, monoecious; roots fibrous 

to woody. Bark furrowed, plated, scaly, or sometimes smooth; twigs terete, sometimes bearing persistent 

primary scale-like cataphylls. Leaves (needles) acicular, linear, or oblong, alternate and spirally arranged 

or fascicled, sessile to short-petiolate, shed singly or in fascicles; resin canals present. Inflorescence 

composed of separate, unisexual scaly ament- or catkin-like cones (male) and woody cones (female). Male 

cones solitary or clustered, axillary, ovoid to ellipsoid or cylindric, maturing and shed annually; 

sporophylls overlapping, bearing 2 abaxial pollen sacs; pollen spherical, 2-winged, less commonly with 

wings reduced to frill, or not winged. Female cones solitary, grouped, or whorled, erect or pendent, sessile 

or short-stalked, ovoid to ovoid-ellipsoid or subglobose, maturing and shed in 1-3 seasons or longpersistent; 

scales spirally arranged, leathery or fleshy when young, relatively thin to strongly thickened 

and woody at maturity, with 2 inverted, adaxial ovules. Seeds winged or unwinged, two to each scale; aril 

lacking; cotyledons 2-12 (-18). A family of 10 genera and ca. 200 species confined almost exclusively to 

the Northern Hemisphere. 

TYPE: Pinus L. 

Reference: Thieret, J.W. 1993. Fam. 3. Pinaceae. Pp. 352-398. In: Flora of North America Editorial 

Committee (eds.), Fl. North Amer., vol. 2. 

Pinus L., Sp. Pl. 1000. 1753. 

1. PINUS 

Evergreen trees or sometimes shrubs. Bark variously furrowed or plated and scaly, exfoliating.

14 


Branches usually in pseudowhorls; shoots dimorphic, long or short; the short ones borne in close spirals 

from axils of scaly bracts and bearing fascicles of leaves. Leaves solitary and spirally arranged or in 

fascicles of (1-) 2-5 (-6); blades linear-lanceolate, linear, or scale-like, ± 2- to 3-angled and rounded on 

abaxial surface, sessile, sheathed at base by persistent or deciduous, scarious basal sheaths; resin canals 

2 or more. Male cones in dense, ament-like, mostly ovoid to cylindric-conic spikes, each in the axil of a 

scale leaf, often numerous and closely clustered around base of current year’s growth. Female terminal or 

subterminal, sessile or stalked, ovoid to oblong-ellipsoid, maturing in 2 (-3) years; cone scales persistent, 

woody or pliable, apical portion of each scale thickened, with a scar, or extended into a hook, spur, claw, 

or prickle. Seeds winged or wingless. Approximately 100 species in north temperate and tropical upland 

regions of North America, Mexico, West Indies, Central America, Europe, Asia, northern Africa, 

Philippines, and Sumatra. 

LECTOTYPE: Pinus sylvestris L., designated by Britton, N. Amer. Trees 5. 1908. 

Reference: Farjon, A. & B. T. Styles. 1997. Pinus (Pinaceae). Fl. Neotrop. Monogr. 75: 1-291. 

1. Pinus caribaea Morelet, Rev. Hort. Côte d’Or 

1: 107. 1851. Neotype: Cuba. Isla de la 

Juventud [Isla de Pinos]. Lückhoff 11608 

(PRF), designated by Lückhoff, Ann. Univ. 

Stellenbosch, Reeks, A, Wis-Naturk. 39(1): 

1-153. 1964. 

Fig. 58. B 

Medium to tall tree, 20-35 (-45) m tall, 50-100 

cm in diam. Trunk erect, terete, slender, clear 

(unbranched) for 2 /3 to ¾ of tree height; bark rough 

and scaly, breaking into irregular square plates 

divided by shallow or deep fissures proximally, 

grey-brown; crown broadly conical, open or 

irregular. Leaves in fascicles of (2-) 3 (-4), very 

rarely 5, straight and rigid, slightly twisted, (12-) 

15-26 (-30) cm × (1-) 1.3-1.8 mm, margins 

serrulate, narrowed to an acute-pungent apex; 

fascicle sheaths 15-20 mm long, papery, with 

erose-ciliate margins. Male cones clustered at ends 

of new shoots, cylindrical, often curved at 

maturity, 20-30 × 5-7 mm, pink or yellow, 

maturing yellowish or reddish brown. Female 

cones subterminal, mostly in pairs or whorls of 3- 

5 (-8), narrowly conic when closed, ovoid to 

ovoid-conical when open, (4-) 5-12 (-13) × (3-) 4- 

Family 3. ZAMIACEAE Zamia Family 

Zamiaceae Horan., Prim. Lin. Syst. Nat. 45. 1834. 

by P. Acevedo-Rodríguez & M. T. Strong 

6 (-7) cm; peduncles curved, 2-2.5 cm long, 

deciduous; cone scales ca. 120-200, thin, oblong, 

nearly flat, straight or recurved; umbo dorsal, flat 

or slightly raised, 3-4 mm wide, up to 3 mm long, 

with a minute persistent prickle. Seeds obliquely 

obovoid, obtuse or angular, slightly flattened, 5-7 

× 2.5-3.5 mm, light grey-brown with dark spots or 

dark brown to blackish, the wings obliquely ovate 

or oblong, 10-20 × 3.8-8 mm, thin, nearly 

transparent, grey or light brown, very finely and 

closely blackish striate. 

General distribution: Naturally occurring in 

the Bahamas, Turks-Caicos Islands, western 

Cuba, Mexico, and Central America. 

Distribution in Puerto Rico: Introduced, 

planted, and naturalized in areas of the central and 

the Luquillo mountains. 

Common names: Puerto Rico: Pino antillano, 

Pino hondureño, Caribbean pine tree. 


Pinus occidentalis Sw., a native of Hispaniola, 

has been planted as an ornamental in Puerto Rico, 

but is not recorded as being naturalized there. 

Palm-like plants; stems subterranean to tall and erect, usually unbranched, cylindrical, with persistent


leaf bases (in Australian genera). Leaves simple-pinnate, spirally arranged, interspersed with cataphylls; 

leaflets sometimes dichotomously divided. Longitudinal ptyxis erect to inflexed or sometimes reflexed, 

horizontal ptyxis conduplicate. Leaflets with several sub parallel dichotomously branching longitudinal 

veins, lacking a mid rib; stomata on both surfaces or undersurface only; individual ptyxis flat. Trichomes 

colored or transparent, branched or unbranched, short-curved or idioblastic. Male and female sporophylls 

spirally aggregated into determinate axillary cones. Female sporophylls simple, appearing peltate, with a 

barren stipe and an expanded and thickened lamina with 2 (rarely 3 or more) sessile, orthotropous ovules 

inserted on the inner (axis facing) surface and directed inwards (“inverted”). Seeds radiospermic. Two 

subfamilies with nine genera and ca. 150 species in tropical and warm temperate regions of Africa, 

Australia and North and South America. 

TYPE. Zamia L. 

References: Eckenwalder, J. E., 1980. Taxonomy of the West Indian Cycads. J. Arnold Arbor. 61: 

701-722. Jones, D. L. 1993. Cycads of the world. Smithsonian Institution Press, Washington DC, 312 pp. 

Negrón Ortiz, V. & G. J. Breckon. 1989. A note on the dispersal of Zamia (Zamiaceae) in Puerto Rico. 

Caribbean J. Sci. 25(1-2): 86-87. Newell, S. J., 1983. Reproduction in a natural population of cycads 

(Zamia pumila L.) in Puerto Rico. Bull. Torrey Bot. Club 110 (4): 464-473. Newell, S. J. 1986. Variation 

in leaf morphology among three populations of Zamia L. in Puerto Rico. Taxon 35(2): 234-242. Newell, 

S. J., 1989. Variation in leaflet morphology among populations of Caribbean Cycads (Zamia). Amer. J. 

Bot. 76 (10): 1518-1523. The Cycad Pages. 2004. [http://plantnet. rbgsyd.gov.au/PlantNet/cycad/ 

index.html]. Stevenson, D. W., 1987. Again the West Indian Zamias. Fairchild Trop. Gard. Bull. 42(3): 

23-27. Stevenson, D. W. (ed.), 1990. The biology, structure, and systematics of the Cycadales. Mem. New 

York Bot. Gard. 57: 1-210. 

Zamia L., Sp. Pl. ed. 2, 1659. 1763. 

1. ZAMIA 

Terrestrial or very seldom epiphytic, fern-like or palm-like, dioecious woody plants; stems massive, 

unbranched or branched, tuber-like and wholly or partly subterranean or cylindrical and wholly emergent. 

Leaves spirally arranged, few to many in a terminal crown, pinnate; leaflets coriaceous, opposite or nearly 

so, with numerous longitudinal veins but lacking a mid rib, entire, toothed toward the apex, or sometimes 

spinulose, articulate at base; petioles elongated, swollen at base, usually with spiny prickles; “stipules” 

paired, inconspicuous. Cones solitary to numerous, with sporophylls in vertical rows, these hexagonal at 

apex; staminate cones cylindrical, erect or decumbent, with wedge-shaped, short-stalked sporophylls; 

pistillate cones ovoid or barrel-shaped, erect, with peltate sporophylls that are faceted or flattened at apex. 

Seeds ovoid or subglobose with orange, red, yellow or brown sarcotesta. A Neotropical genus with 50-60 

species (some of which are still undescribed) occurring in a wide range of habitats, many of them 

cultivated in the tropics as house and garden plants. 

There has been much disagreement with respect to the number of Zamia species recognized in Puerto 

Rico and the West Indies. In a detailed study of leaf morphology, based on herbarium specimens, 

Eckenwalder (1980) found a continuous gradation of foliar characters throughout the West Indies, 

concluding that Zamia was represented in the West Indies by two subspecies of Z. pumila. Accordingly, 

all native populations of Zamia in Puerto Rico are Z. pumila ssp. pumila. A contrasting view is presented 

by Stevenson (1987) where 4 species are recognized in Puerto Rico based on foliar and strobili characters 

derived from herbarium specimens and live plants. For the most part (except for the recognition of Z. 

integrifolia), the present authors follow Stevenson’s opinion realizing that further work is needed (DNA 

sequencing) in order to confirm or reject his assessment. 

TYPE: Zamia pumila L.

16 


Key to the species of Zamia 

(modified after Stevenson, 1987) 

1. Leaflet narrowly lineate, 5-7 mm wide, with entire or seemingly entire margins, the apex long acute; 

seeds red to blood-red ……….........................................................................…….. 2. Z. portoricensis 

1. Leaflet lineate-oblong, oblong or narrowly obovate, 10 or more mm wide, with serrate or callous 

margins, the apex acute, obtuse, erose or truncate; seeds orange or orange-red. ................................ 2 

2. Leaflets oblong, 1-2 cm wide …………………........................................……… 3. Z. pumila 

2. Leaflet oblanceolate to obovate, 1.5-3.5 (-5) cm wide ……….....….…. 1. Z. amblyphyllidia 

1. Zamia amblyphyllidia D.W. Stev., Fairchild 

Trop. Gard. Bull. 42(3): 26. 1987. Type: 

Puerto Rico; Utuado (Río Abajo). Acevedo- 

Rdgz. 576 (holotype: NY, isotypes: FTG, 

NY). 

Zamia latifoliolata sensu Britton & P. Wilson in 

part, non Prenleloup, 1872. 

Fig. 58. C 

Palm-like, acaulescent shrub 0.7-1.2 m tall; 

stem subterranean, tuberous, 3-10 (-25) cm in 

diameter, often dichotomously branched, smooth, 

devoid of old leaf bases. Young leaves rusty 

pubescent. Leaves arched-ascending, several to 

numerous, forming a crown; leaflets (2)5-15 pairs, 

the distal ones opposite, but the basal ones 

alternate, narrowly oblanceolate to obovate, 12-25 

× 1.5-3.5 cm, base decurrent onto the rachis, the 

apex obtuse, rounded or truncate, often erose or 

notched apically, the margins remotely and 

minutely serrate in upper half; petioles smooth. 

Cataphylls ca. 1.5 cm long, subulate. Staminate 

cones pedunculate, resembling a corn-cob, 6-20 × 

1-3 cm, densely reddish brown pubescent. Ovulate 

cones 1(3), pedunculate, cylindrical with long, 

gradually tapering blunt sterile apex, 6-15 × 4-6.5 

cm, densely reddish brown or dark brown 

pubescent. Seeds ovoid-angled, 1-2 cm long, with 

a orange-red fleshy coat. 

General distribution: Known to occur in 

Jamaica, western Cuba, and Puerto Rico. 

Distribution in Puerto Rico: Widely distributed 

along the northern karst region. Rare to locally 

common. Recorded for Arecibo, Bayamón, 

Ciales, Manatí, Toa Alta, Toa Baja, and Utuado. 

Common name: Puerto Rico: Marunguey. 


Arecibo: Cambalache State Forest Reserve, 

Strong et al. 463 (US). Bayamón: Bo. Hato Tejas, 

Axelrod 7616 (UPRRP). Ciales: ca. 80 m E of 

coordinates 18 23.593’N, 66 29.29’W, Acevedo- 

Rdgz. et al. 13494 (US). Toa Alta: Along Rt. 677, 

2 mi. S of Rt. 2, Miller & Taylor 5917 (UPRRP). 

Toa Baja: Finca Nevares al lado de Carrt. # 2, 

Acevedo-Rdgz. & Chinea 2144 (NY, UPR, US). 

2. Zamia portoricensis Urb., Symb. Antill. 1: 

291. 1899. Type: Puerto Rico; Guánica. 

Sintenis 3782 (holotype: B, isotype: NYfragment). 

Fig. 58. D 

Fern-like shrub, 40-90 cm tall; stem 

subterranean, tuberous, 6-15 cm in diameter, 

dichotomously branched, wrinkled. Leaves 2-10, 

ascending-arched; leaflets 5-30 pairs, opposite or 

subopposite, linear, 8-25 × 0.5-0.8 cm, often 

canaliculate above, acute at apex, entire; petioles 

smooth. Staminate cones 1-10, pedunculate, 

resembling a corn-cob, 3-15 × 0.8-2 cm, densely 

pubescent. Ovulate cones 1 (-3), short-pedunculate, 

cylindrical to slightly ovoid with blunt or slightly 

acute apex , 6-12 × 4-5 cm, dark reddish brown 

sometimes becoming gray when mature, densely 

pubescent. Seeds ovoid, 1-2 cm long, with red or 

blood-red fleshy coat. 

General distribution: Endemic to Puerto 

Rico. 

Distribution in Puerto Rico: Known only 

from SW Puerto Rico. Recorded for Guánica, 

Sabana Grande, and Yauco. 

Common name: Puerto Rico: Marunguey. 


Guánica: Guánica Forest Reserve, Axelrod & 

Escobar 2879 (UPRRP). Sabana Grande: Susúa 

Forest Reserve, Axelrod et al. 3821 (UPRRP). 

Yauco: Susúa Forest Reserve, Axelrod & Chavez 

4044 (UPRRP).


3. Zamia pumila L., Sp. Pl., ed. 2, 1659. 1763. 

Lectotype: Hispaniola. J. Commelin, Horti 

Med. Amstelod. 1: t. 58, 1697, designated by 

Eckenwalder, J. Arnold Arbor. 61: 715. 1980. 

Zamia debilis L. f. in Aiton, Hort. Kew. 3: 478. 

1789. Type: Hispaniola. J. Commelin, Horti 

Med. Amstelod. 1: t. 58, 1697. 

Zamia integrifolia L. f. in Aiton, Hort. Kew. 3: 

477. 1789. Holotype: United States; Florida, 

(a plant cultivated at Kew Gardens), Aiton s.n. 

(K). 

Zamia media Jacq., Pl. Hort. Schoenbr. 3: 77, t. 

397, 398. 1798. Lectotype: West Indies. Jacq., 

Hort. Schoenb. 3: t. 398. 1798, designated by 

Eckenwalder, J. Arnold Arbor. 61: 715. 1980. 

Zamia floridana A. DC., Prodr. 16(2): 544.1868; 

Zamia angustifolia var. floridana (A. DC.) 

Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 

4(4): 315. 1876. Type: United States; Florida. 

Hulse s.n. (holotype: G-DC). 

Zamia latifoliolata Prenl., Bull. Soc. Vaud. Sci. 

Nat. 11: 278. 1872. Type: Dominican 

Republic. Prenleloup s.n. (holotype: LAU). 

Zamia allison-armourii Millsp., Publ. Field 

Columbian Mus., Bot. Ser. 2: 23. 1900. Type: 

Dominican Republic; Santo Domingo. 

Millspaugh 817 (holotype: F-60817, isotype: 

NY-fragment). 

Zamia erosa O.F. Cook & G.N. Collins, Contr. U. 

S. Natl. Herb. 8: 267. 1903. Type: Puerto 

Rico; Vega Baja. Cook s.n. (lectotype: NY, 

isolectotype: NY), designated by Eckenwalder, 

J. Arnold Arbor. 61: 716. 1980. 

Fern-like shrub to 1 m tall; stem subterranean, 

tuberous, 3-10(25) cm in diameter, often 

dichotomously branched, wrinkled. Leaves 2-15; 

leaflets 5-30 pairs, opposite or subopposite, 

oblong, 8-25 × 0.5-2 cm, acute or somewhat 

CULTIVATED FAMILIES 

rounded at apex, often deeply cleft or erose, with 

10-15 distinct teeth in upper fourth; petioles 

smooth. Cataphylls subulate, to 3.5 cm long. 

Staminate cones 1-10 (30), pedunculate, resembling 

a corn-cob, 3-15 × 0.8-2 cm, dark reddish brown. 

Ovulate cones 1-5, pedunculate, cylindrical to 

slightly ovoid with blunt or slightly acute apex , 6- 

12 × 4-6 cm, dark reddish brown sometimes 

becoming gray when mature, densely pubescent. 

Seeds ovoid-angled, 1-2 cm long, with an orangered 

fleshy coat. 

General distribution: Distributed in S.E. 

United States (Georgia & Florida), Bahamas, 

central Cuba, Hispaniola, and southern Puerto 

Rico. 

Distribution in Puerto Rico: Known from the 

municipalities of Aguadilla, Bayamón, Coamo, 

Dorado, Guayanilla, Isabela, Loíza, Manatí, 

Ponce, Salinas, Santa Isabel, Toa Alta, Vega Baja, 

and Yauco. 

Common names: Puerto Rico: Marunguey, 

Palmita de jardin. 

Note: Zamia integrifolia is here lumped under 

Z. pumila because the putative distinguishing 

character, i.e., the entire leaflet margins, is not 

present in its lectotype. 


Aguadilla: Rt. 107, km 2.2 (near Rt. 458), Axelrod 

& Ackerman 261 (UPRRP). Dorado: Rt. 677, ca. 2 

km S of Rt. 2, ca. 1 km N of Hacienda Dorado, 

Taylor & Molano 8629, 8630 (UPRRP). 

Guayanilla: El convento, Quebrada los Cerros, 

limestone hills, Galarza et al. 108 (UPRRP). 

Isabela: Bo. Jauca 2, Proctor & Colón 50478 (SJ- 

2). Loíza: Calle Villanueva of Pueblo Indio, 

Taylor et al. 9469 (UPRRP). Manatí: Bo. Tierras 

Nuevas Saliente, Proctor & Figueroa 50647 (SJ). 

Salinas: Campamento Santiago, Bo. Río Jueyes, 

Proctor 50391 (SJ). 

A number of species of the gymnosperm families Araucariaceae and Cupressaceae are planted as 

ornamentals in Puerto Rico and the Virgin Islands. In the Araucariaceae, these include: Agathis robusta 

(F.J. Müll.) F.M. Bailey (native to Australia), Araucaria angustifolia (Bertol.) Kuntze (native to Brazil), 

A. bidwillii Hook. (native to Australia), and A. heterophylla (Salisb.) Franco (native to the Norfolk 

Islands). In the Cupressaceae, these include: Chamaecyparis funebris (Endl.) Franco (native to China), 

Cupressus arizonica Greene (native to Arizona, U.S.A.), C. lusitanica Mill. (native to Mexico and Central 

America), C. macrocarpa Hartw. (native to California, U.S.A.), C. sempervirens L. (native to southern 

Europe), and Thuja orientalis L. (native to China and Korea).

18 


MONOCOTYLEDONS 

Key to the families 

1. Plants of aquatic or marine environments ........................................................................................ 2 

2. Plants submersed, floating, or emersed in fresh to brackish waters ............................................ 3 

3. Plants plate-like, tiny, 1-2.5 mm long .................................................................... Lemnaceae 

3. Plants not plate-like, leaves well-developed with blades well over 1 cm long ...................... 4 

4. Inflorescence sessile or essentially so, 1-flowered ........................................................... 5 

5. Plant a floating rosette of leaves, acaulescent; inflorescence included within a spathe 

............................................................................................................. Araceae (Pistia) 

5. Plant submersed and rooting with elongate often branching stems; inflorescence axillary, 

not included within a spathe ................................. Hydrocharitaceae (Najas) 

4. Inflorescence pedunculate, 1- to many-flowered, not included within a spathe, or may be 

included within a spathe-like bract when immature ....................................................... 6 

6. Perianth present, white, blue, lilac, or rarely yellow ................................................... 7 

7. Leaves submersed, strap-like or ribbon-like ................................ Hydrocharitaceae 

7. Leaves emersed or floating on surface (at least some) with petioles and dialated 

linear, lanceolate, ovate, elliptic, orbicular, or sagittate blades ............................ 8 

8. Leaf blades floating on surface or ascending slightly above it, orbicular or broadly 

elliptic ................................................................................................. 9 

9. Flowers unisexual, inconspicuous; ovary inferior; fruit a many-seeded berry 

.................................................................... Hydrocharitaceae (Limnobium) 

9. Flowers bisexual, showy; ovary superior; fruit of distinct follicles 

.................................................................... Limnocharitaceae (Hydrocleys) 

8. Leaf blades emersed, generally erect, linear, lanceolate, ovate, elliptic, or sagittate 

.......................................................................................................... 10 

10. Fruit a many-seeded capsule ............................................. Pontederiaceae 

10. Fruit an achene ..................................................................... Alismataceae 

6. Perianth absent .......................................................................................................... 11 

11. Inflorescence a spike of 2-many flowers; stipes of fruits (druplets) not elongating 

................................................................................................... Potamogetonaceae 

11. Inflorescence a cluster of 2 or rarely more flowers; stipes of fruits (druplets) 

elongating ............................................................................................. Ruppiaceae 

2. Plants submersed in marine or brackish environment ............................................................. 12 

12. Leaves strap-like, 4-12 mm wide ............................................................. Hydrocharitaceae 

12. Leaves linear or cylindrical, to 2 mm wide ...................................................................... 13 

13. Leaves linear, with a conspicuous midvein, narrowly acuminate at apex .. Ruppiaceae 

13. Leaves cylindrical, without midvein, truncate to concave at the expanded apex (in ours) 

.................................................................................................. Cymodoceaceae 

1. Plants terrestrial, epiphytic, or of marshy environments ............................................................... 14 

14. Plants woody, with a single trunk and an unbranched crown of large (> 75 cm long) leaves 

........................................................................................................................................ Arecaceae 

14. Plants herbaceous or woody; stem short and covered by a basal rosette of leaves, or the stem 

elongate and often branched, with leaves scattered along the stem ........................................ 15 

15. Plant with a definite woody stem ..................................................................................... 16 

16. Trunks with prop roots at base; flowers in heads or spadices ............... [Pandanaceae] 

16. Trunks without prop roots at base; flowers in branching inflorescences ................... 17 

17. Inflorescence an axillary raceme or panicle; fruit a fleshy berry or sometimes hard 

and woody, usually red or orange; seeds globose or elongate 

........................................................................................................ Dracaenaceae


17. Inflorescence a large terminal panicle; fruit a fleshy or chartaceous capsule; seeds compressed 

........................................................................................ Agavaceae (Yucca) 

15. Plant herbaceous, or if woody, the stem very short and leaves in a rosette ................................. 18 

18. Plants climbing (vines) ......................................................................................................... 19 

19. Plants with bisexual flowers .......................................................................................... 20 

20. Leaves reduced and scale-like; branches transformed into verticillate or fasciculate 

needle-shaped cladodes ...................................................................... Asparagaceae 

20. Leaves well-developed; branches not forming cladodes ................. Araceae (in part) 

19. Plants with unisexual flowers ........................................................................................ 21 

21. Plant monoecious, climbing aided by adventitious roots; leaves with a stout midvein 

and secondary pinnate veins; inflorescence a spadix, subtended by a spathe 

......................................................................................................... Araceae (in part) 

21. Plants dioecious, twining, or climbing by tendrils; leaves with 3-9 arcuate mid veins; 

inflorescence spike-like, racemose, paniculate or umbellate, not subtended by a bract 

............................................................................................................... 22 

22. Plants with stipular tendrils; inflorescence umbellate; fruit a berry 

..........................................................................................................Smilacaceae 

22. Plants twining, without tendrils; inflorescence a spike, raceme or panicle; fruit a 

chartaceous samara or a 3-winged capsule. .............................Dioscoreaceae 

18. Plants erect, prostrate or decumbent, not climbing ............................................................... 23 

23. Inflorescence a spadix subtended by a spathaceous bract. ................................... Araceae 

23. Inflorescence a spike, a cyme, a raceme or a panicle not subtended by a spathaceous bract 

............................................................................................................................. 24 

24. Flowers inconspicuous, perianth scale-like or reduced to bristles or scales .......... 25 

25. Flowers subtended by 3-6, whorled, scale-like perianth segments (tepals); fruit a 

3- to many-seeded capsule .............................................................. Juncaceae 

25. Flowers subtended by a single or pair of scale-like bracts; fruit a one-seeded 

achene-like follicle, achene, or utricle ............................................................ 26 

26. Flowers unisexual, densely arranged in separate, large cylindrical spikes on 

the same stalk; fruit a one-seeded achene-like follicle, subtended by whorls 

of fine perigonal hairs ..................................................... Typhaceae 

26. Flowers bisexual or sometimes unisexual, arranged in small, compressed or 

terete, spikes or spikelets in branching or head-like inflorescences; fruit an 

achene or utricle, naked or subtended by hypogynous barbed bristles .... 27 

27. Flowers spirally imbricate or less often 2-ranked, subtended by a single 

scale; leaf-sheath usually closed; stems triangular or cylindrical; fruit 

an achene with or without bristles .......... Cyperaceae 

27. Flowers distichously arranged, subtended by a pair of scales; leaf-sheath 

usually open; stems cylindrical; fruit an utricle without bristles 

............................................................. Poaceae (forthcoming volume) 

24. Flowers showy, either the sepals or the petals large, not scale-like nor reduced to 

bristles or scales ..................................................................................................... 28 

28. Ovary superior ................................................................................................. 29 

29. Leaves in a rosette or rosulate at base ...................................................... 30 

30. Leaves chartaceous to rigid-coriaceous .......................... Bromeliaceae 

30. Leaves fleshy, thickened .................................................................... 31 

31. Leaves 30-60 cm long, with a gelatinous core, not fibrous; scapes to 

1 m tall .............................................................. Asphodelaceae 

31. Leaves 90-275 cm long, with a fibrous core; scapes 5-7 m tall. 

........................................................ Agavaceae (Agave, Furcraea) 

29. Leaves alternate, opposite or spirally arranged ........................................ 32

20 


32. Leaves reduced and scale-like; branches transformed into verticillate or fasciculate 

needle-shaped cladodes .............................................................................. Asparagaceae 

32. Leaves well-developed; branches not forming cladodes ......................................... 33 

33. Flowers subtended by indurate scales, imbricately arranged in a solitary, ovoid or 

ellipsoid, cone-like spike at the apex of a naked culm ............................. Xyridaceae 

33. Flowers not subtended by scales, arranged in cymes, racemes or panicles ....... 34 

34. Inflorescence cymose, usually enclosed by a folded, boat-shaped, spathelike 

bract; flower with distinct calyx and corolla, free or basally connate 

..................................................................................... Commelinaceae 

34. Inflorescence racemose or paniculate, not enclosed by a boat-shaped bract; 

flower perianth tubular or funnel-shaped ................................... Dracaenaceae 

28. Ovary inferior ............................................................................................................................... 35 

35. Leaf veining pinnate, secondary veins arising along the primary mid rib from base to apex, 

these parallel ........................................................................................................................ 36 

36. Fertile stamens 5(6) ....................................................................................................... 37 

37. Leaves and bracts spirally arranged; flowers unisexual; inflorescences with chartaceous, 

wine-colored bracts ............................................................... Musaceae 

37. Leaves and bracts distichously arranged; flowers bisexual; inflorescences with fleshy, 

showy, colorful bracts .............................................................. Heliconiaceae 

36. Fertile stamen 1 ............................................................................................................. 38 

38. Leaves eligulate; ovary or fruit warty ...................................................... Cannaceae 

38. Leaves with a pair of stipule-like ligules; ovary or fruit smooth ....................... 39 

39. Leaves with a swollen pulvinus or joint at the junction of the petiole and blade 

.........................................................................................................Marantaceae 

39. Leaves lacking a swollen pulvinus or joint at the junction of the petiole and blade 

................................................................................................................ 40 

40. Leaves distichously arranged; sheaths open; ovary with 2 apical nectary glands 

.................................................................................... Zingiberaceae 

40. Leaves spirally arranged; sheaths closed; ovary without apical glands 

..................................................................................................... Costaceae 

35. Leaf veining parallel, secondary veins arising from the base of the primary mid rib .......... 41 

41. Flowers zygomorphic; anthers usually 1, sessile near the apex of a column; plants commonly 

epiphytic, less often lithophytic or terrestrial .................. Orchidaceae (see Ackerman 1995) 

41. Flowers actinomorphic or nearly so; stamens 3 or 6, with long, distinct or basally connate 

filaments; plants terrestrial .............................................................................. 42 

42. Perianth conspicuously pubescent .................................................... Haemodoraceae 

42. Perianth not conspicuously pubescent ..................................................................... 43 

43. Stamens 3 ........................................................................................................ 44 

44. Leaves reduced, scale-like; plant saprophytic (not green and photosynthetic) 

.................................................................................................. Burmanniaceae 

44. Leaves well-developed; plant not saprophytic .............................. Iridaceae 

43. Stamens 6 ........................................................................................................ 45 

45. Leaves swollen at base into a bulb; perianth segments > 6.5 cm long; fruit a 

capsule; seeds glabrous ....................................................... Amaryllidaceae 

45. Leaves not swollen at base; perianth segments to 1.5 cm long; fruit a berry; 

seeds pilose ........................................................................... Hypoxidaceae


Family 1. ARACEAE Philodendron Family 

Araceae Juss., Gen. Pl. 23. 1789, nom. conserv. 

by P. Acevedo-Rodríguez & D. H. Nicolson 

Erect perennial herbs, often with creeping or tuberous rhizomes, or climbing robust herbs with 

adventitious roots. Leaves alternate, heterophyllous in numerous genera, often all basal, stout, simple, 

compound, dissected or lobed; petioles forming a sheath at base. Flowers minute, bisexual or unisexual 

(monoecious), in a terminal, densely-flowered spadix with a large, subtending bract (spathe) of variable 

size and shape; perianth of 4 or 6 distinct or connate tepals in two cycles, or these reduced or wanting; 

stamens (1-) 4 or 6 (8), the filaments mostly short, distinct or connate, the anthers opening by terminal 

pores or longitudinal slits; ovary superior, often unilocular with basal to lateral parietal placentation, or 

less often with several locules of (2) 3 (-15) united carpels with axile placentation, the ovules few or 

solitary or less often numerous, the style terminal and short or the stigma sessile. Fruit a berry or 

developing with the spadix and maturing as a multiple fruit. A family of about 110 genera and ca. 2, 450 

species, most with tropical and subtropical distribution. 

TYPE: Arum L. 

References: Engl., A. & K. Krause, 1920. Araceae-Colocasioideae. In: A. Engler (ed.) Das 

Pflanzenreich IV. 23E (Heft 71). Leipzig. Govaerts, R. & D. Frodin. 2002. World Checklist and 

Bibliography of Araceae. Royal Botanic Garden, Kew. Howard, R. A. 1979. Nomenclatural notes on the 

Araceae of the Lesser Antilles. J. Arnold Arbor. 60: 272-289. Mayo, S. J., J. Bogner, and P. C. Boyce, 

1997. The Genera of Araceae. Royal Botanic Garden, Kew. Nicolson, D. H. 1987. Araceae. Pp. 17-101. 

In: Dassanayake, M. D. & F. R. Fosberg (eds.). A Revised handbook to the Flora of Ceylon. Vol. 6. 

Smithsonian Institution, Washington, DC. 

Key to the genera 

1. Plants floating, aquatic; leaves subsessile, in a dense rosette ...............................................11 Pistia 

1. Plants terrestrial, epiphytic or of marshy environments but not floating; leaves petiolate, alternate or 

in a loose rosette .......................................................................................................................... 2 

2. Plants erect, not climbing ............................................................................................................ 3 

3. Stems aerial, erect, non-rhizomatous (non-starchy) .................................................................... 4 

4. Leaf blades lanceolate or triangular-lanceolate, cordate at base; petioles to 1 m long ............. 

......................................................................................................10 Philodendron giganteum 

4. Leaf blades elliptic, ovate or nearly lanceolate, the base acute, obtuse, truncate or seldom subcordate; 

petioles 30-40 cm long ..................................................................................... 5 

5. Spathe ovate, not differentiated into a tube and blade, free from the spadix; stamens solitary, 

free .................................................................................................. [Aglaonema] 

5. Spathe convolute, surrounding the spadix, slightly constricted at the middle and basally 

adnate to the pistillate portion of the spadix; stamens 4-5, connate into a synandrium ..... 

......................................................................................................................... 5 Dieffenbachia 

3. Stems subterranean and modified into tubers or rhizomes, or aerial but massive, and starchy 6 

6. Leaves solitary or less often 2 or 3; blades trisect; petioles with banded variegation .............. 

............................................................................................................................... 6 Dracontium 

6. Leaves in a loose rosette; blades entire; petioles not band-variegated ................................... 7 

7. Flowers bisexual and with obvious perianth ..................................................................... 8 

8. Tertiary veins reticulate; spathe green, reflexed (red, orange, white, erect or spreading in 

cultivated species) ..................................................................................... 2 Anthurium 

8. Tertiary veins parallel-pinnate; spathe white, erect ............................. 12 Spathiphyllum 

7. Flowers unisexual, lacking perianth .................................................................................. 9 

9. Leaves peltate at base ................................................................................................. 10

22 


10. Leaf blades usually variegated with small white, pink, or reddish, spots; spadix 

lacking a distal appendage ......................................................................... 3 Caladium 

10. Leaf blades green; spadix usually with a distal sterile appendage .......... 4 Colocasia 

9. Leaves sagittate or cordate at base ............................................................................ 11 

11. Spadix with a sterile distal appendage ................................................... 1 Alocasia 

11. Spadix with distal fertile portion (without a sterile appendage) ......................... 12 

12. Spathe strongly constricted, the lower part forming a tube with convolute 

margins, the upper part a concave, evanescent blade ............. 14 Xanthosoma 

12. Spathe not constricted, sub-cylindrical or obconic, persistent 

................................................................................................. [Zantedeschia] 

2. Plants climbing herbs or vines aided by adventitious roots ....................................................... 13 

13. Flowers bisexual ................................................................................................................. 14 

14. Perianth present; spathe chartaceous and reflexed, not covering the spadix; stems to 5 

mm in diam.; leaves 7-13 cm long, simple, green, not-perforated ............... 2 Anthurium 

14. Perianth absent; spathe fleshy, erect, covering (surrounding) the spadix; stems to 2 cm in 

diam.; leaves 20-50 cm long, pinnately compound or simple, simple leaves variegated 

or perforated ............................................................................................................... 15 

15. Leaves pinnately divided, or if simple green or variegated (green and yellow) and 

non-perforated; ovary unilocular; seeds reniform .............................. 7 Epipremnum 

15. Leaves simple, perforated; ovary bilocular; seeds ovoid or cylindrical ...... 18 Monstera 

13. Flowers unisexual ............................................................................................................... 16 

16. Leaves pedately dissected; sap milky ........................................................ 13 Syngonium 

16. Leaves entire, cordate, elliptic or pinnately compound; sap clear .................... 

.......................................................................................................10 Philodendron (in part) 

1. ALOCASIA 

Alocasia (Schott) G. Don in Sweet, Hort. Brit., ed. 3, 631. 1839, nom. conserv. 

Erect, medium-sized to rarely arborescent herbs, terrestrial; stems thick, with clear or milky sap, often 

subterranean, stoloniferous or bulbiferous, aerial portion erect. Leaves alternate, few to many in a terminal 

crown, simple, sinuate to deeply pinnatifid, sagittate, or less often hastate or cordate, sometimes peltate at 

base, long-petiolate, sometimes subtended by a cataphyll. Inflorescence 2-many per node. Spathe usually 

herbaceous, partially enclosing the spadix, strongly constricted, forming a tube with convolute margins, 

shorter than the erect, boat-shaped blade; spadix sessile or short-stipitate, shorter than the spathe, with a 

sterile distal portion (appendage), a pistillate conical-cylindrical basal zone, separated from the staminate 

zone by a narrower zone of sterile flowers, and a staminate cylindrical or conical zone. Flowers unisexual, 

lacking perianth; stamens 3-12(36), connate into a synandrium; ovary 1-locular or partially 3- or 4-locular 

at apex, with 6-10 basal ovules per locule, the style short, the stigma depressed-capitate, 3- or 4-lobed. 

Fruit fleshy, usually bright red, with 1-5 seeds. A genus of 60-70 species native to the Old World tropics 

and subtropics, with 3 species cultivated and naturalized in the Neotropics. 

TYPE: Alocasia cucullata (Lour.) G. Don (≡ Arum cucullatum Lour.), typ. conserv. 

References: Bunting, G. S. & D. H. Nicolson. 1963. The Alocasia plumbea confusion. Baileya 11: 

142-146. Hay, A., 1998. The genus Alocasia (Araceae-Colocasieae) in West Malaysia and Sulawesi. 

Gard. Bull. Straits Settlem. 50: 221-334. 

Key to the species of Alocasia 

1. Leaf blades green, coriaceous; petioles green; plant not flowering or fruiting in Puerto Rico 

...................................................................................................................................…. 1. A. macrorrhizos


1. Leaf blades purplish tinged at least abaxially, chartaceous; petioles purplish variegated to completely 

burgundy or dark purple; inflorescence paired; spathe blade reflexed at anthesis, chartaceous, 

adaxially pinkish ….....................................................................................................… 2. A. plumbea 

1. Alocasia macrorrhizos (L.) G. Don in Sweet, 

Hort. Brit. ed. 3, 631. 1839 [as 

“macrorrhizon”]; Arum macrorrhizon L., Sp. 

Pl. 965. 1753. Lectotype: Sri Lanka. 

Hermann, Parad. Bat. t. 73. 1698, designated 

by Furtado, Gard. Bull. Straits Settlem. 11: 

252. 1941 (see Nicolson, Rev. Fl. Ceylon 6: 

58. 1987, for discussion). 

Fig. 58. E 

Glabrous, terrestrial herb, 1-1.5 m tall, 

producing watery sap; acaulescent or developing 

an elongated caudex with age; acaulescent plants 

with a short, conical corm. Leaves in a rosette, 

ascending; blades flattened, ascending, with basal 

sinus projecting downward, 25-50 (-100) × 20-36 

(-100) cm, green (although white-variegated in 

some cultivars), slightly lustrous, lance-ovate, 

coriaceous, wavy or slightly plicate along 

secondary veins, the apex acute or obtuse and 

apiculate, the base hastate, the sinuses nonoverlapping, 

to 30 cm long, the margins wavy, 

with a submarginal vein within 2 mm from 

margin; midvein and secondaries stout, prominent 

on both surfaces, lower surface with dark spots on 

secondary vein angles; petioles 60-100 cm long, 

invaginate on lower ½, upper margins of 

invagination revolute. Not known to flower or 

fruit in Puerto Rico. 

General distribution: Native to India through 

SE Asia and Malesia. Introduced and escaped 

throughout the tropics as an ornamental and as a 

source of food. 

Distribution in Puerto Rico: Common herb of 

moist secondary forests, abandoned farms, along 

stream and river banks. Recorded for Arecibo, 

Bayamón, Gurabo, Loíza, Naguabo, and San 

Sebastián. 

Common name: Puerto Rico: Yautía 

cimarrona. 


Arecibo: Río Abajo State Forest, Acevedo-Rdgz. 

10755 (UPRRP, US). Bayamón: Bo. Santa Olaya, 

Quebrada Collores, Proctor 43188 (SJ). Gurabo: 

Bo. Mamey: Road 189, km 10.4, Proctor 45122 

(SJ). Loiza: Vacia Talega, disturbed area of sand, 

Axelrod & Thomas 12688 (UPRRP). Naguabo: 

Bo. Río Blanco: near Road 191 ca. 200 m from jct 

w/Road 31, Proctor 45127 (SJ). San Sebastián: 

Sargent 280 (US). 

2. Alocasia plumbea Van Houtte, Fl. Serres 

Jard. Eur. 21: 93. 1875. Lectotype: Java (?). 

Van Houtte, Fl. Serres Jard. Eur. 21: t. 

2206. 1875, here designated. 

Fig. 58. F 

Glabrous, acaulescent, terrestrial herb, ca. 1 m 

tall, with a short starchy corm or rhizome at base. 

Leaves in a basal rosette; blades more or less 

flattened, ascending or less often pointing 

downward, 40-60 × 20-35 cm, slightly lustrous, 

adaxially green, abaxially purplish or pinkish 

tinged, elliptic-lanceolate, chartaceous, wavy or 

slightly plicate along secondary veins, with a 

submarginal vein within 3 mm from margin, the 

apex obtuse or acute, apiculate, the base sagittate, 

symmetrical, the sinuses nearly touching each 

other, ca. 20 cm long; midvein and secondaries 

stout, whitish, prominent on lower surface, lower 

surface with dark spots on secondary vein angles; 

petioles to 1 m long, green, variegated with purple 

or completely purple, ascending or slightly arched, 

invaginate on lower ½. Inflorescences axillary, 

two per leaf, ascending; peduncles to 25-30 cm 

long, cylindrical, green, tinged with purple or 

pinkish; spathe chartaceous, the tube green, ca. 4 

cm long, the blade oblong-elliptic, reflexed at 

anthesis and soon withering, 20-21 cm long, 

abaxially green with purplish tinge, pinkish 

adaxially; spadix cream, the distal sterile zone 12- 

14 cm long. 

General distribution: Native to SE Asia, 

introduced in Puerto Rico as an ornamental, now 

naturalized. 

Distribution in Puerto Rico: Occasional, 

cultivated and escaped or naturalized in moist 

disturbed habitats, such as roadsides. Recorded for 

Adjuntas, Arecibo, Luquillo, and Toa Baja. 

Common name: Puerto Rico: Yautía 

ornamental. 

Note: Although recent workers (Hay, 1998;

24 


Govaerts & Frodin, 2002) suggest Alocasia 

plumbea to be a cultivar of the variable species A. 

macrorrhizos, we are treating it as a distinctive 

species because the populations of both taxa as 

found in Puerto Rico and other parts of the 

Neotropics are quite distinctive from each other 

(refer to key above). 

2. ANTHURIUM 

Anthurium Schott, Wiener Z. Kunst 1829: 828. 1829. 


Adjuntas: Bo. Limaní, Acevedo-Rdgz. et al. 13377 

(MAPR, US). Luquillo: Luquillo Mountains, 

Caribbean National Forest, Acevedo-Rdgz. 10808 

(UPRRP, US). Toa Baja: Bo. Media Luna, 1.7 km 

due E of Campanilla, Proctor & González 50496 

(SJ-2). 

Erect herbs or vines climbing by adventitious roots, terrestrial or epiphytic; stems elongate, fleshy or 

the plant acaulescent. Leaves alternate, simple, or palmately lobed, long-petiolate, enveloped by a 

cataphyll in early stages. Inflorescence solitary. Spathe usually herbaceous, not enclosing the spadix, 

reflexed, usually long-lived, green, whitish to brightly colored; spadix sessile or short-stipitate, cylindrical 

or conical, many-flowered, flowering from base to apex. Flowers bisexual, sessile; perianth segments 4; 

stamens 4; ovary 2-locular, with 1 or 2 pendulous ovules per locule, the style short or wanting, the stigma 

disk-like to 4-lobed. Fruit a 2-locular, fleshy, bright red, white or lavender berry, often pendulous from a 

threadlike structure when ready for dispersal; seeds oblong. The largest genus in the family with ca. 800 

species native to the Neotropics. 

LECTOTYPE: Anthurium acaule (Jacq.) Schott (≡ Pothos acaulis Jacq.), designated by Britton & P. 

Wilson, Bot. Porto Rico 5: 128. 1923. 

References: Croat, T. B. 1991. A revision of Anthurium section Pachyneurium (Araceae). Ann. 

Missouri Bot. Gard. 78: 539-855. Mayo, S. J. 1982. Anthurium acaule (Jacq.) Schott (Araceae) and the 

West Indian ‘bird’s nest’ Anthuriums. Kew Bull. 36: 691-719. Sheffer, R. C. et al. 1980. Taxonomy of 

Anthurium scandens (Araceae). Aroideana 3: 86-93. 

Key to the species of Anthurium 

1. Plant an epiphytic vine with slender, scandent stems, 1-2 m long; leaves alternate, to 13 cm long 

.........................................................................................................................................4. A. scandens 

1. Plant terrestrial or epiphytic but not climbing, acaulescent or stems 10-50 cm long; leaves in a rosette, 

20-80 cm long ……………........................................................................................................…….. 2 

2. Leaves attenuate at base; petioles much shorter than the blade …………........……. 2. A. crenatum 

2. Leaves cordate or truncate at base; petioles as long or nearly as long as the blade ……..……… 3 

3. Leaves ovate to triangular-ovate, the blades directed downward from the point of attachment 

at base ………................................................................................................ 1. A. cordatum 

3. Leaves lanceolate or elliptic, the blades ascending …...........................................…………. 4 

4. Blades 30-70 cm long; petioles 30-50 cm long; inflorescence sterile, the peduncle 60-103 

cm long; spathe 10-15 cm long, attenuate at apex, spadix 15-40 cm long 

…….....................................................................................................….. 5. A. × selloum 

4. Blades 20-40 cm long; petioles 16-35 cm long; inflorescence fertile, the peduncle 7-28 

cm long; spathe 2.5-4.5 cm long, caudate at apex; spadix 2.2-4.6 (11) cm long 

………...........................................................................................…. 3. A. dominicense 

1. Anthurium cordatum (L.) Schott, Wiener Z. 

Kunst 1829: 828. 1829; Pothos cordatus [ as 

“cordata”] L., Sp. Pl. ed. 2, 1373. 1763. 

Lectotype: Martinique (?), Plumier (Burman 

ed.), Pl. Amer., t. 38. 1756, designated by R.A. 

Howard, Fl. Lesser Antill. 3: 376. 1979. 

Anthurium cordifolium sensu Eggers, Fl. St. Croix 

98. 1879; Millspaugh, Publ. Field Columbian 

Mus., Bot. Ser. l: 477. 1902, and Little et al., 

Fl. Virgin Gorda 16. 1976, non Kunth, 1841. 

Fig. 2. A-G.


Glabrous, erect, terrestrial herb 0.5-1.5 m tall; 

stem erect or decumbent, cylindrical, 10-50 (75) 

cm long and ca. 5 cm diam., with numerous, large 

leaf scars; cataphylls 10-20 cm long, weathering 

into persistent fibers. Leaves spirally arranged in a 

loose rosette; blades directed downward, 30-52 × 

25-30 cm, broadly ovate to triangular-ovate, 

coriaceous, strongly reticulate-veined, the apex 

obtuse, the base cordate-sagittate with overlapping 

sinuses, the margins entire, slightly wavy; petioles 

erect, cylindrical, 20-60 cm long. Inflorescence 

ascending; peduncles to 50 cm long; spathe 

reflexed, green, leafy, oblong to ligulate, to 10 cm 

long; spadix ascending, pendulous when fruits are 

mature, 10-30 cm long, dark brown, widened at 

base. Berry bright red, obovoid, 5-8 mm long, 

produced on proximal portion of spadix, 

embedded in the axis when immature, pendulous 

from a thread-like structure when ripe. 

General distribution: Jamaica, Virgin Islands 

and Lesser Antilles. 

Distribution in the Virgin Islands: Locally 

common understory herb of shaded moist forests. 

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

Virgin Gorda. 

Common name: St. John: Maroon jancole. 

Selected specimens examined: ST. CROIX: Mt. 

Eagle, Thompson 457 (US). N slope Bodkin Hill, 

Fosberg & Ogden 55352 (US). E slopes of Mt. 

Eagle; ca. 1.7 km S of Canebay, Croat 61021 (US). 

Maroon Ridge area NE of Springgarden, Proctor 

44985 (SJ, US). ST. JOHN: Lamesur, N.L. Britton & 

Shafer 516 (US). Cinnamon Bay on road to 

Herman Farm, Mori & Woodbury 16998 (NY-2). 

TORTOLA: Sage Mt. Natl. Park, Proctor 50569, 

50570 (SJ). Pickate Ghut, W of Mt. Sage, Proctor 

& Barwick 41983, 41984 (SJ). High Bush, N.L. 

Britton & Shafer 813 (US).VIRGIN GORDA: Gorda 

Peak Natl. Park, Proctor & Barwick 41959 (SJ). 

2. Anthurium crenatum (L.) Kunth, Enum. Pl. 3: 

75. 1841; Pothos crenatus [ as “crenata’] L., 

Sp. Pl. ed. 2, 1373. 1763. Lectotype: St. 

Thomas, U. S. Virgin Is., Plumier (Burman 

ed.), Pl. Amer. t. 39. 1756, designated by 

Croat, Ann. Missouri Bot. Gard. 78: 635. 

1991. 

Anthurium acaule sensu many West Indian 

authors, non (Jacq.) Schott, 1832. 

Anthurium huegellii sensu Eggers, Fl. St. Croix 

98. 1879, Bello, Anales. Soc. Esp. Esp. Hist. 

Nat. 12: 115. 1883, and Millspaugh, Publ. 

Field Columbian Mus., Bot. Ser. l: 477. 1902, 

non Schott, 1855. 

Anthurium lanceolatum Kunth ex Bello, Anales 

Soc. Esp. Hist. Nat. 12: 115. 1883, nom. nud. 

Anthurium acaule var. portoricense Kuntze, 

Revis. Gen. Pl. 2: 738. 1891. Type: Puerto 

Rico. Kuntze, s. n. (holotype: NY-2 sheets!; 

isotypes: K, MO). 

Anthurium acaule var. brevipes Engl., Bot. Jahrb. 

Syst. 25: 362. 1898. Type: Puerto Rico. 

Sintenis 1582 (isotype: K). 

Fig. 58. G; 59. A 

Glabrous, erect terrestrial or epiphytic herb 

0.8-1.5 m tall; stem prostrate, ca. 10 cm long, with 

numerous adventitious roots, and numerous 

persistent cataphylls. Leaves ascending, spirally 

arranged in a loose rosette; blades 40-80 × 15-25 

cm, elliptic, oblong-elliptic or oblanceolate, 

coriaceous, strongly reticulate-veined, very often 

weathering into a network of veins, the apex 

obtuse, with a short apiculum, the base obtuse or 

rounded, tapering from below the middle of the 

blade, the margins entire to slightly crenate; 

petioles 5-20 cm long, flattened along the ventral 

surface, swollen at point of attachment with blade. 

Inflorescence ascending; peduncles to 80 cm long; 

spathe to 12 cm long, reflexed, green, leafy, 

lanceolate, deciduous; spadix ascending, pendulous 

when fruits are mature, 10-30 cm long, dark 

brown, widened at base. Berry bright red, obovate, 

5-6 mm long, produced on proximal portion of 

spadix, embedded in the axis when immature, 

pendulous from a thread-like structure when ready 

for dispersal. 

General distribution: Hispaniola, Puerto 

Rico, and the Virgin Islands. 

Distribution in Puerto Rico and the Virgin 

Islands: Common understory herb of shaded moist 

forests, and scrub forests. Recorded for Arecibo, 

Barranquitas, Bayamón, Cabo Rojo, Camuy, 

Cayey, Culebra, Juana Díaz, Luquillo, Maricao, 

Mayagüez, Peñuelas, Río Grande, Salinas, San 

Juan, Toa Baja, Yabucoa, Yauco, and Vieques; St. 

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

Common names: Puerto Rico: Flor de 

Culebra, Hoja de costado, Moco de pavo; St. John: 

Scrub-bush. 


Arecibo: Cambalache State Forest: Bo. 

Garrochales, Proctor 42413 (SJ); Río Abajo State

26 


Forest: Bo. Río Arriba, Proctor et al. 49978 (SJ). 

Barranquitas: Bo. Barrancas, upper NE slope & 

summit of Monte La Torrecilla, Proctor & 

Thomas 44444 (SJ). Bayamón: Stevenson 384 

(US); Bo. Hato Tejas, Webster & Miller 8607 

(US). Cabo Rojo: Sierra Bermeja: Bo. Llanos 

Costa, upper slopes & summit of Cerro Mariquita, 

Proctor & McKenzie 44031 (SJ). Camuy: 

Montañas Guarionex, Bo. Puertos, top of Mogote 

0.45 km due SSE of intersection of Roads 119 & 

456, Proctor et al. 48920 (SJ). Culebra: Bo. 

Flamenco, on peak 0.6 km due S of Playa Resaca, 

Proctor 39985 (SJ). Culebra: Britton & Wheeler 

114 (US). Juana Díaz: Bo. Guayabal, upper S side 

& summit of Cerro de las Cuevas, Proctor & Pinto 

40788 (SJ). Maricao: Bo. Maricao Afuera. Río 

Maricao margins, Cedeño, & Jiménez 550 (SJ). 

Mayagüez: Vicinity of Mayagüez, NE of Ponce, 

Heller 6283 (US). Peñuelas: Bo. Rucio, upper 

slopes of Cerrote de Peñuelas, Proctor & Haneke 

43267 (SJ, US). Río Grande: Bo. Zarzal: 0.9 km 

due NNW of Cerro Bravo, Proctor 42672 (SJ). 

Salinas: Bo. Lapa, summit of E peak, Las Tetas de 

Cayey, Proctor 44406 (SJ). San Juan: Stone 

Quarry, 2 mi. E of Santurce, Heller & Heller 1267 

(US). Toa Baja: Bo. Candelaria, wooded mogotes 

0.4-0.7 km WSW of Pico Nevárez, Proctor & 

Carrasquillo 45306 (SJ). Vieques: Isabel Segunda 

to Campo Cielo, Shafer 2377 (US). Yabucoa: Bo. 

Calabazas, hwy 3, app. Km 101.5, Proctor & 

Rivera 46942 (SJ). Yauco: Bo. Sierra Alta, summit 

area of Pico Rodadero, Proctor & Díaz 44738 (SJ). 

ST. JOHN: Bethania to Rosenberg, N.L. Britton & 

Shafer 225 (US). Bordeaux, N.L. Britton & Shafer 

545 (US). Cinnamon Bay on road to Herman 

Farm, Mori & Woodbury 16999 (NY). ST. 

THOMAS: Eggers 191 (US). Pearl to Bonne 

Resolution, Britton & Marble 1320 (US). 

Raunkiaer s.n. (US). TORTOLA: Pickate Ghut, W of 

Mt. Sage, Proctor & Barwick 41989 (SJ), N.L. 

Britton & Shafer 811 (US). 

3. Anthurium dominicense Schott, Oesterr. Bot. 

Z. 8: 350. 1858. Type: Dominica. Imray s. n. 

(K), according to R.A. Howard, Fl. Lesser 

Antill. 3: 377. 1979. 

Anthurium dominicense var. sintenisii Engl., Bot. 

Jahrb. Syst. 25: 418. 1898. Type: Puerto Rico. 

Sintenis 4206 (syntype: F), Sintenis 1581 

(syntypes: NY!, US!). 

Glabrous, erect terrestrial or epiphytic herb, 

50-75 cm tall; stem erect, 3-5(10) cm long, with 

persistent cataphylls, these subulate, 5-9 cm long, 

weathering into persistent fibers. Leaves ascending, 

spirally arranged in a loose rosette; blades 15- 

(35)50 × 3-14(18.5) cm, lanceolate or oblonglanceolate, 

coriaceous, strongly reticulate-veined, 

the apex acute, with a short apiculum, the base 

subcordate, truncate or less often rounded, the 

margins entire to slightly crenate; petioles 20-41 

cm long, furrowed along the ventral surface, 

darker at point of attachment with blade. 

Inflorescence ascending; peduncles 7-28 cm long; 

spathe 2.5-4.5 cm long, reflexed, green, leafy, 

lanceolate, caudate at apex; spadix ascending, 2.2- 

4.6(11) cm long, green, subcylindrical or widened 

at base. Berry bright red, ellipsoid to subglobose, 

5-6 mm long, produced on spadix for most of its 

length. 

General distribution: Puerto Rico and the 

Lesser Antilles. 

Distribution in Puerto Rico: Occasional 

understory herb or epiphyte of moist forests. 

Adjuntas, Canóvanas, Ciales, Luquillo, Maricao, 

Naguabo, Patillas, Peñuelas, Ponce, and Río 

Grande. 

Common name: Puerto Rico: Lengua de vaca. 


Adjuntas: Alto de la Bandera, N.L. Britton & 

Shafer 2013 (US). Canóvanas: Sierra de Luquillo, 

Caribbean Natl. Forest. W end of El Toro trail, first 

1.5 km from road 186, Proctor & Taylor 46300 

(SJ). Ciales: Toro Negro State Forest, upper SE 

slopes & summit of Cerro Rosa, Proctor et al. 

44594 (SJ). Luquillo: Caribbean Natl. Forest, 

Proctor 44751(SJ, US). Maricao: Maricao State 

Forest, NE ridge of Monte Allegrillo, Proctor & 

Padrón 41665 (SJ). Naguabo: Loma la Mina, SE 

side to E Peak, Shafer 3281 (US). Patillas: Carite 

Forest Reserve, Proctor 48487 (SJ). Peñuelas: Bo. 

Rucio, upper slopes of Cerrote de Peñuelas, 

Proctor & Haneke 43268 (SJ). Ponce: Toro Negro 

State Forest, Proctor 40079 (US, SJ). Río Grande: 

Caribbean Natl. Forest, summit of Mt. Britton, 


4. Anthurium scandens (Aubl.) Engl. in Martius, 

Fl. Bras. 3(2): 78. 1878; Dracontium 

scandens Aubl., Hist. Pl. Guiane 836. 1775. 

Epitype: Haiti; vic. of Port de Paix, La Coup 

road, 22 Jan 1929, Leonard 12281 (US), here 

designated. 

Fig. 1. A-F


Fig. 1. A-F. Anthurium scandens. A. Fertile branch. B. Inflorescence. C. Flower, and stamen with accompanying tepal. D. 

Stamen, frontal and side views. E. Fertilized ovary. F. Infructescence. From Acevedo-Rdgz., P. 2003, Bejucos y plantas trepadoras 

de Puerto Rico e Islas Vírgenes, Smithsonian Institution. 

Glabrous, epiphytic, herbaceous vine, climbing 

by means of adventitious roots; stem cylindrical, 

to 5 mm in diam., 1-2 m long, internodes with 

persistent, straw-colored, fibrous leaf sheaths 

(cataphylls), weathering into persistent fibers. 

Leaves alternate; blades simple, 7-13 × 2.5-5 cm, 

lanceolate-elliptic or elliptic, chartaceous, the 

apex acuminate, the base obtuse or attenuate, the 

margins entire; upper surface glabrous, foveate; 

lower surface glabrous, with numerous dark 

punctations; petioles 0.7-4.5 cm long. Inflorescence 

axillary, ascending; spathe 1-1.2 cm long, 

reflexed, green with purplish venation, lanceolate; 

spadix ascending, 1.5-2 cm long. Berry white, 

depressed-globose, ca. 5 mm in diam.; seeds 

ellipsoid, ca. 1 mm long. 

General distribution: Widely distributed 

throughout the Neotropics. 

Distribution in Puerto Rico: Common 

epiphytic vine of moist to wet forest. Adjuntas, 

Aibonito, Arecibo, Bayamón, Camuy, Cayey, 

Ciales, Guayama, Luquillo, Maricao, Naguabo, 

Ponce, Río Grande, Salinas, San Sebastián, 

Utuado, and Yauco. 

Common name: Puerto Rico: Guinda. 

Note: In his original publication of Dracontium 

scandens, Aublet cited Plumier’s Descr. Pl. Amér. 

fig. 74 as the type for this name. However, fig. 74 

is a species of Pothomorphe (Piperaceae). This 

apparent mistake was corrected by Sheffer et al.

28 


(1980) when they indicated that the type was 

actually Plumier’s fig. 78. As they indicated, 

Plumier’s fig. 78 does not match what has been 

traditionally recognized as Anthurium scandens, 

the inflorescence as depicted and described by 

Plumier is way too long. According to Dr. Tom 

Croat (MO), the referred plate, does not represent 

any West Indian species even remotely. He thinks 

that more likely it represents a mixed collection or 

even something entirely made up. The glanddotted 

leaves are fine but the inflorescence must be 

from a different (birds nest) Anthurium. In view of 

the ambiguity of the current type of Anthurium 

scandens we have decided to epiypify this name 

with a specimen from the type locality (near Port 

de Paix) cited by Plumier, which clearly represents 

this well-known and common species. 


Adjuntas: Monte Guilarte State Forest, Bo. 

Garzas, Mountain slopes c. 1.3-1.5 km due SE of 

Monte Guilarte Peak, Proctor & Alemañy 45835 

(SJ). Aibonito: Stevenson 5044 (US). Arecibo: Río 

Abajo Forest Reserve, Acevedo-Rdgz. 9395 (UPR, 

UPRRP, US). Bayamón: Bo. Guaraguao Arriba: 

upper slope & summit of Cerro La Peña, Proctor 

41706 (SJ). Camuy: Montañas Guarionex, Bo. 

Puertos, Top of mogote 0.45 km due SSE of 

intersection, Roads 119 & 456, Proctor et al. 

48919 (SJ). Cayey: Bo. Carmen, along crest (S 

side) of Sierra Jajome, 2-2.8 km due NE-ENE. of 

village of Carmen, Proctor 46929 (SJ, US); 

Cayey, Underwood & Griggs 415 (US). Maricao: 

Monte Montoso, Britton & Cowell 4138 (US). 

Ciales: along trail Camino de la Ceiba towards 

Quebrada del Pozo Azul, Acevedo-Rdgz. 11880 

(UPRRP, US). Naguabo: Río Icaco and adjacent 

hills, Shafer 3562 (US). Ponce: on the Adjuntas 

Road, 11 mi. from Ponce, Heller 6357 (US). Río 

Grande: along Road 186 to El Verde area of 

Caribb. Natl. Forest, Acevedo-Rdgz. & Siaca 9339 

(UPR, UPRRP, US). Salinas: Bo. Lapa, summit of 

western peak, Las Tetas de Cayey, Proctor 41931 

(SJ). Río Grande: Sierra de Luquillo, Sintenis 1505 

(US). San Sebastián: Bo. Guajataca, along W side 

of Río Guajataca in gorge S of Lago de Guajataca, 

Proctor et al. 48118 (SJ). Utuado: Britton & 

Cowell 1031 (US). Yauco: Bo. Sierra Alta, summit 

area of Pico Rodadero, Proctor & Díaz 44737 (SJ). 

5. Anthurium × selloum K. Koch in Klotzsch, 

App. Gen. Sp. Nov. 1855: 8. 1855-56. 

Neotype: St. John, U. S. Virgin Is. Krebs, s. n. 

(C), designated by Croat [as lectotype ¹], Ann. 

Missouri Bot. Gard. 78: 727. 1991. 

Fig. 59. B 

Glabrous, erect, terrestrial or seldom epiphytic 

herb, 0.5-1 m tall; stem erect, short, with numerous 

adventitious roots; cataphylls to 18 cm long, 

persistent, weathering to show numerous fibrous 

remnants. Leaves numerous, ascending, spirally 

arranged in a loose rosette; blades 30-70 × 13-26 

cm, lanceolate, oblong-lanceolate or cordatelanceolate, 

coriaceous, strongly reticulate-veined, 

the apex obtuse to short-acuminate, the base 

cordate to nearly truncate, the margins entire, 

slightly wavy; petioles 30-50 cm long, cylindrical, 

canaliculate along the ventral surface, swollen at 

point of attachment with blade. Inflorescence 

ascending; peduncles 60-103 cm long, cylindrical; 

spathe to 15 cm long, spreading to reflexed, green, 

leafy, oblong-lanceolate; spadix ascending, 15-40 

cm long, dark brown, cylindrical, narrowed 

toward the apex. Sterile, not producing berries. 

General distribution: Endemic to the 

mountains of St. John and Tortola; locally 

common understory herb of shaded moist forests. 

Grown as a garden plant on Tortola. 

Note: Anthurium × selloum seems to be a 

sterile hybrid between A. cordatum and A. 

crenatum. It has never been described, collected, 

or observed in fruit; the inflorescences develop 

normally but die before setting fruits. Ovules and 

pollen grains are produced, but the pollen is not 

available since the anthers do not dehisce. 

Although A. × selloum is capable of vegetative 

reproduction, most individuals seem to be the 

result of crossing from the two putative parent 

species. The morphology of Anthurium × selloum 

is intermediate between that of the parent species, 

and the taxon is normally found where the parent 

species occur together. 

¹ Koch’s description of A. selloum was based 

on plants cultivated at B, and did not include the 

specimen collected by Krebs on St. John. 

Therefore, the Krebs specimen is not original 

material and cannot be regarded as a lectotype but 

instead as a neotype. 

Selected specimens examined: ST. JOHN: 

Bordeaux, N.L. Britton & Shafer 560 (US); Cruz 

Bay Quarter, Acevedo-Rdgz. & Siaca 3810 (MO, 

NY, US). TORTOLA: Pickate Ghut, W of Mt. Sage, 

Proctor & Barwick 41984 (SJ).



Anthurium andreanum Linden and A. 

longilaminatum Engl., both native to South 

3. CALADIUM 

Caladium Vent., Mag. Encycl. 4(16): 463. 1801. 

America, were reported by Martorell et al. (1981) 

as cultivated in Puerto Rico. 

Acaulescent, tuberous herbs. Leaves simple or rarely trisect, several in a loose rosette, often peltate 

and variegated, cordate or sagittate, secondary veins forming an irregular collecting vein between the 

primary lateral veins; petiole long, with distinct sheath. Inflorescence pedunculate, 1 per node. Spathe 

erect, constricted, the basal area forming a convolute, green tube that surrounds the spadix, the upper area 

opened into a whitish and withering blade; spadix cylindrical, erect, stipitate or sessile, pistillate below 

and staminate to the apex with a constricted transition zone of sterile flowers. Flowers unisexual, the 

perianth wanting; stamens 3-5, united into truncate synandria with marginal thecae; ovary incompletely 

2- or 3-locular, with many parietal ovules, the stigma sessile. Fruit a fleshy, 1- to many-seeded, whitish 

berry; seeds ovoid to ellipsoid, with many longitudinal grooves. A genus of about 12 species native to the 

Neotropics, but one species widely cultivated and escaping in the tropics. 

LECTOTYPE: Caladium bicolor (Aiton) Vent. (≡ Arum bicolor), typ. conserv. prop. 

1. Caladium bicolor (Aiton) Vent., Mag. Encycl. 

4(16): 464. [22 Dec 1800-21 Jan]1801; Arum 

bicolor Aiton, Hort. Kew. 3: 316. 1789. 

Cyrtospadix bicolor (Aiton) Britton & P. 


Neotype: Madeira Is., Bot. Mag. 21: fig. 820. 

1805, here designated ². 

Xanthosoma sylvestre Bello, Anales Soc. Esp. 

Hist. Nat. 12: 114. 1883. Type: Puerto Rico. 

Bello, s. n. (destroyed). 

Glabrous, erect acaulescent herb with an 

obconical, fleshy corm at base. Leaves 1-2; blades 

pointing downward, ca. 30 × 20 cm, lanceolate to 

hastate, chartaceous, usually with small, irregular 

whitish or pinkish spots, or variegated along 

secondary veins, less often completely green, 

glaucous beneath, the apex acute or shortly 

acuminate, the base peltate, cordate, the margins 

more or less wavy; petioles erect, 35-55 cm long, 

sheathing, white at the very base, usually with 

purple stripes. Inflorescences axillary, ascending, 

solitary; peduncles as long as or little shorter than 

the petioles, cylindrical, green, usually with purple 

stripes; spathe chartaceous, glaucous, to 14 cm 

long, the blade twice as long as the tube, withering, 

elliptic, apiculate at apex; spadix shorter than the 

spathe, the staminate zone twice as long as the 

pistillate. 

General distribution: Native to South 

America, but widely cultivated as a foliage 

ornamental, naturalized in tropical and subtropical 

areas. 


Islands: Cultivated as ornamental and naturalized 

in open disturbed, moist areas, common along 

rivers and swampy areas. Aguas Buenas, 

Aibonito, Arecibo Bayamón, Ciales, Mayagüez, 

Quebradillas, and San Juan. 

Common names: Puerto Rico: Paleta de 

pintor, Malanga cimarrona, Yautía cimarrona. 

Note: ² The plant used by Aiton to describe 

this species was presumably from the Madeira 

Islands and perhaps cultivated at K. Searches in K 

or BM for a specimen that could be regarded as the 

type for this name have failed, suggesting that the 

plant was never vouchered with a herbarium 

specimen. As a result, we are designating a 

neotype for this commonly cultivated species. 


Aguas Buenas: Sintenis 2515 (US). Arecibo: 

Bosque de Río Abajo, carretera 621, Acevedo- 

Rdgz. & Laboy 174 (SJ). Bayamón: Stevenson 386 

(US). Mayagüez: vicinity of Cerro Las Mesas, 

Cowell 552 (US). Quebradillas: Bo. Cacao, along 

E side of Río Guajataca at bottom of canyon, 0.5- 

1 km SE to SSE of mouth of Quebrada, Proctor 

50208 (SJ, US). San Juan: Santurce, Goll et al. 78 

(US).

30 


4. COLOCASIA 

Colocasia Schott in Schott & Endlicher, Melet. Bot. 18. 1832, nom. conserv. 

Small to large herbs, the stems subterranean and tuberous, or epigeal and massive. Leaves simple, 

several in a rosette in acaulescent species or terminal in arborescent species, often peltate, secondary veins 

forming a collective vein between the primary lateral veins; petiole long, with distinct sheath. 

Inflorescence pedunculate, 1 to many per node. Spathe erect, constricted, the basal area forming a 

convolute, accrescent tube that surrounds the spadix base, the blade much longer than the tube, white or 

yellow and withering; spadix erect, sessile, shorter than the spathe, with a sterile distal portion 

(appendage), and a basal pistillate zone, separated from the staminate zone by a narrower zone of sterile 

flowers. Flowers unisexual, lacking perianth; stamens 3-6, connate into a truncate synandrium; ovary 1locular, 

with numerous parietal ovules, the style short, the stigma discoid-capitate or 2-lobed. Berry 

oblong, green or white, many-seeded; seeds ovoid to ellipsoid, with numerous grooves. A genus of 8 

species native to tropical Asia, with 1 species cultivated and naturalized throughout the tropics. 

TYPE: Colocasia antiquorum Schott (≡ Arum colocasia L.), typ. conserv. 

1. Colocasia esculenta (L.) Schott in Schott & 

Endicher, Melet. Bot. 18. 1832; Arum 

esculentum L., Sp. Pl. 965. 1753. Lectotype: 

Jamaica. Sloane, Voy. Jamaica 1: t. 106, f. 1. 

1707, designated by Nicolson in A. C. Smith, 

Fl. Vit. Nov. 1: 456. 1979. 

Arum colocasia L., Sp. Pl. 965. 1753; Colocasia 

antiquorum Schott in Schott & Endicher, 

Melet. Bot. 18. 1832; Caladium colocasia 

(L.) W. Wight, Contr. U. S. Natl. Herb. 9: 206. 

1905. Lectotype: “Habitat in Cretae, Cypri, 

Syriae, Aegypti aquosis.” (LINN-1079.4), 

designated by Mayo in Bosser et al., Fl. 

Mascareignes 192: 18.1984. 

Fig. 59. C 

Glabrous, acaulescent herb, with a massive, 

fleshy corm at base, with lateral, thick, edible 

runners. Leaves in a loose rosette; blades pointing 

downward, 23-55 × 12-38 cm, cordate or 

lanceolate, sub-coriaceous, green above, glaucous 

below, the apex obtuse, acute or shortly 

acuminate, the base peltate-cordate, the margins 

more or less wavy, with a submarginal collecting 

vein; petioles erect, to 85 cm long, inserted 3-7 cm 

from base of blade, invaginate on lower 1 /3. 

Inflorescences axillary, ascending, solitary; 

peduncles nearly as long as the petiole, cylindrical; 

spathe fleshy, to 35 cm long, the tube green, the 

blade lanceolate, not much wider than the tube, 

yellow to orange, flexing open near the base, then 

deflexing and dropped; spadix yellow, much 

shorter than the spathe, the sterile flower zone and 

the distal appendage shorter than the fertile zones. 

General distribution: Perhaps native to SE 

Asia, now pantropically cultivated and escaped. 


Islands: Commonly cultivated for its edible 

tubers, sometimes persistent or naturalized along 

river banks and moist forest understory. Arecibo, 

Caguas, Carolina, Jayuya, Loíza, Salinas, San 

Juan, and Toa Baja. 

Common names: Puerto Rico: Malanga, 

Yautía malanga. 



10783 (UPRRP, US). Caguas: Bo. Bairoa, along or 

near Río Bairoa just NE of hwy 30 bridge, Proctor 

& Colón 50045 (SJ). Carolina: 204 St. Country 

Club, Ortega 121 (UPRRP). Jayuya: Cerro Punta, 

open eroded slope, Acevedo-Rdgz. et al. 14450 

(UPRRP, US). Loíza: Bo. Medianía Alta, along 

sandy dike, NW bank of Río Herrera at S end of 

road 964, Proctor et al. 49607 (SJ, US). Salinas: 

Bo. Lapa, vicinity of Las Tetas de Cayey, Proctor 

& Thomas 44612 (SJ). San Juan: Río Piedras, UPR 

Botanic Garden, muddy field on edge of secondary 

forest, Acevedo-Rdgz. 14454 (UPRRP, US). Toa 

Baja: Bo. Candelaria, wooded mogotes 0.6-0.7 km 

WSW of Pico Nevárez, Proctor & Thomas 45619 

(SJ).


5. DIEFFENBACHIA 

Dieffenbachia Schott, Wiener Z. Kunst 1829: 803. 1829. 

Erect to decumbent herbs, usually of wet or swampy areas; stems thickened, elongate, usually fragile, 

producing a whitish, poisonous, caustic sap. Leaves simple, usually spotted or variegated, mostly borne 

distally on stems, long-petiolate. Inflorescence 2-several per node; cataphylls short and inconspicuous. 

Spathe greenish, persistent, partially enclosing the spadix, more or less constricted at the middle, the basal 

part elongated, forming a tube with convolute margins, the upper part a short, erect or recurved blade; 

spadix cylindrical, shorter than the spathe, the pistillate zone (basal) basally to entirely adnate to the 

spathe, laxly-flowered, separated from the staminate zone by a sub-naked axis with scattered sterile 

staminate flowers, the staminate zone distal, sub-cylindrical, longer than the free pistillate portion of the 

spadix. Flowers unisexual, the perianth wanting, pistillate flowers with 4 staminodes; stamens 4-5; ovary 

1- to 3-locular, with a solitary ovule per locule, the stigma sessile, nearly globose to 3-lobed. Fruit a fleshy 

berry; seeds ovoid to globose. A genus of about 135 species native to the Neotropics. 

TYPE: Dieffenbachia seguine (Jacq.) Schott (≡ Arum seguine Jacq.) 

Reference: Croat, T.B. 2004. Revision of Dieffenbachia (Araceae) of Mexico, Central America, and 

the West Indies. Ann. Missouri Bot. Gard. 91: 668-772. 

1. Dieffenbachia seguine (Jacq.) Schott, Wiener 

Z. Kunst 1829: 803. 1829; Arum seguine 

Jacq., Enum. Syst. Pl. 31. 1760; Dieffenbachia 

plumieri Schott, Oesterr. Bot. Wochenbl. 2: 

69. 1852, nom. illeg. Lectotype: West Indies. 

Plumier, Descr. Pl. Amér., t. 61, 1693, 

designated by R.A. Howard, Fl. Lesser Antill. 

3: 389. 1979. 

Caladium maculatum Lodd., Bot. Cab. 7: 608. 

1822; Dieffenbachia maculata (Lodd.) G. 

Don in Sweet, Hort. Brit., ed. 3, 632. 1839. 

Type: South America. Lodd., Bot. Cab. 7: t. 

608. 1822. 

Figs. 2. H, I; 59. D 

Erect to decumbent herb to 2 m tall; stem 

cylindrical, ca. 3 cm diam., green with leaf scars, 

producing a milky or watery, caustic sap when cut. 

Leaves borne distally on stems; blades usually 

spotted or variegated, 20-50 × 12-24 cm, elliptic or 

oblong-ovate, chartaceous, the apex acute or 

acuminate, the base rounded or cordate, usually 

asymmetric, the margins entire; petioles 30-40 cm 

long, sheathing. Inflorescences axillary, ascending, 

in groups of 2 or 3; peduncles 4-15 cm long, 

cylindrical; spathe coriaceous, 18-30 cm long, 

green, erect, opening throughout its length, keeled 

at apex; spadix shorter than the spathe, with green 

axis; pistillate zone with widely spaced flowers 

with yellow stigmata and 4 white staminodes; 

staminate zone densely clustered, cream. Berry 

orange-red. 

General distribution: Throughout tropical 

America. 


Islands: An occasional herb of swamps, moist 

forests, wet seepage, sunny to partly shaded areas. 

Arecibo, Bayamón, Cayey, Ciales, Fajardo, 

Maricao, Naguabo, Río Grande, San Juan, Utuado, 

and Vega Alta; St. John, St. Thomas, and Tortola. 

Common names: Puerto Rico: Rábano, 

Rábano cimarrón. Virgin Islands: Dumb cane. 

Notes: The sap of this plant is irritating to the 

skin, and if ingested is inflammatory to the tongue 

and digestive tract, hence the common name, 

dumb cane. 


Arecibo: Río Abajo State Forest, wetland just S of 

Road 621, 0.8 km due W of Campamento Crozier, 

Proctor & Chabert 44721 (SJ). Bayamón: Bo. 

Guaraguao Arriba, along Road 879 ca. 1 km due 

WSW of Cerro La Peña, Proctor & Thomas 44558 

(SJ). Cayey: Bo. Lapa, NW side, summit ridge of 

Cerro Avispa, Proctor 42761 (SJ). Ciales: along 

trail Camino de la Ceiba towards Quebrada del 

Pozo Azul, Acevedo-Rdgz. & Vicens 11876 

(UPRRP, US). Fajardo: Colonia San Miguel, N.L. 

Britton & Shafer1625 (US). Maricao: Sargent 643 

(US). Mayagüez: vic. of Cerro Las Mesas, Cowell 

550 (US). Naguabo: Río Icaco & adjacent hills, 

Shafer 3484 (US). Río Grande: Caribbean Natl. 

Forest, El Verde Field Station, Thompson & 

Thompson 3238 (US). San Juan: Río Piedras, 

Stevenson 2160 (US). Utuado: Bo. Don Alonso,

32 


Fig. 2. A-G. Anthurium cordatum. A. Habit. B. Leaf. C. Infructescence. D. Flower, top view. E. Stamen with subtending perianth 

segment. F. Stamen, frontal and lateral views. G. Ovary and l.s. ovary. H-I. Dieffenbachia seguine. H. Fertile branch. I. 

Inflorescence and subtending leaf. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78.


Acevedo-Rdgz. 13418 (FTG, NY, US).Vega Alta: 

Bo. Sabana; NE of Regadera, Proctor & Thomas 

44571 (SJ, US). ST. JOHN: Maho Bay Gut, 

Dracontium L., Sp. Pl. 967. 1753. 

6. DRACONTIUM 

Erect, small to large herbs, acaulescent with a depressed-subglobose tuber. Leaves solitary; blade 

trisected; petiole stout, long, often verrucose and with banded variegation, sheathed only at base. 

Inflorescence 1(2) per node; cataphylls conspicuous. Spathe erect, boat-shaped, brown-purple outside, 

reddish purple outside, marcescent or persistent, partially enclosing the spadix, not constricted, basally 

convolute; spadix cylindrical or ellipsoid, sessile or stipitate, much shorter than the spathe, flowers fertile 

throughout, except for a few sterile apical ones. Flowers bisexual, with 4-8 sepals; stamens 4-5, free; ovary 

ovoid incompletely 1-to 6-locular, with a solitary axial or basal ovule per locule, the style elongated, the 

stigma sub-globose to 3-lobed. Berry obpyramidal greenish brown; seeds rounded to reniform. A genus 

of about 23 species native to Central America, South America, Lesser Antilles and Puerto Rico. 

LECTOTYPE: Dracontium polyphyllum L., designated by Britton & P. Wilson, Bot. Porto Rico 5: 130. 

1923. 

References: Zhu, G. H. & M. H. Grayum. 1995. Two hitherto confused species, Dracontium 

polyphyllum and D. asperum (Araceae), and typification of their names. Taxon 44: 519-523. Zhu, G. 1995. 

Systematics of Dracontium (Araceae). Ph. D. dissertation, University of St. Louis, Missouri. Zhu, G. & 

T. B. Croat. 2004. Revision of Dracontium (Araceae). Ann. Missouri Bot. Gard. 91: 593-667. 

1. Dracontium asperum K. Koch, Wochenschr. 

Gärtnerei Pflanzenk. 33: 259. 1859. Neotype: 

Surinam. Irwin 55045 (NY; COL, F, GH, K, 

US!, VEN), designated by Zhu & Grayum, 

Taxon 44: 522. 1995. 

Dracontium polyphyllum sensu authors, non 

Linnaeus, 1753. 

Fig. 59. E, F 

Acaulescent herb with a hemispherical tuber 

and numerous lateral smaller, elongated tubercles. 

Leaves solitary or less often two or three; blades 

ascending, then the lobes spreading horizontally, 

trisected, green, chartaceous, 70-100 × 70-100 cm, 

the middle segments usually twice triparted from 

the middle, the laterals usually 2-parted below the 

middle, lower surface with prominent primary and 

secondary veins, the segments acuminate at apex, 

with entire to wavy margins; rachis nearly terete; 

petioles 1-2 m long, nearly terete, with banded 

variegation and rough excrescences, especially in 

lower half. Inflorescences solitary, ascending; 

peduncles 15-45 cm long above ground, 

cylindrical, ca. 1 cm in diam. at base, with same 

banded pattern as petiole; cataphyll much shorter 

than the peduncle; spathe 14-20 cm long, reflexed 

Woodbury 863/7293 (VINPS). TORTOLA: Road 

Town to High Bush, N.L. Britton & Shafer 762 

(US). 

at apex, with overlapping margins on lower 

portion completely covering the spadix, inner 

surface purple to purplish red, with a whitish area 

at the base; spadix cream, ca. 7 cm long, nearly 

elliptical. Berries depressed-globose to discoid, 8- 

12 mm in diam., 1-to 3-seeded, purplish brown or 

bluish black at maturity; seeds triangular to 

rounded. 

General distribution: Puerto Rico, Lesser 

Antilles, Trinidad and Tobago, Guyana, Surinam, 

and Brazil. 

Distribution in Puerto Rico: A rare herb of 

wet seepage, shaded areas or shaded river banks. 

Arecibo, Quebradillas, Río Grande, San Sebastián. 

Common name: Puerto Rico: Guapa. 


Arecibo: Río Abajo State Forest, Proctor et al. 

46104 (SJ); Sector Arrozales; Bo. Sabana Hoyos, 

understory of secondary forest, Acevedo-Rdgz. et 

al. 13402 (US). Quebradillas: Bo. San Antonio, 

Proctor et al. 46460, 48047 (SJ). Río Grande: 

Sierra de Luquillo, Proctor & Rivera 47976 (SJ). 

San Sebastián: Bo. Aibonito, Area Recreativa de 

Guajataca, Proctor 48044 (SJ). Bo. San Antonio, 

0.9 km due SW of jct. of Rd. 437 with Rd. 113, 

Proctor 48047 (US).

34 


7. EPIPREMNUM 

Epipremnum Schott, Bonplandia 5: 45. 1857. 

Robust, root-climbing vines. Leaves alternate, simple or pinnately dissected; petioles distally 

pulvinate, with a winged sheath that usually withers into numerous, persistent fibers. Inflorescences 1(2) 

per node. Spathe fleshy, boat-shaped, not constricted, usually deciduous or withering after anthesis, 

covering the spadix; spadix sub-cylindrical, thickened, erect, sessile or stipitate, shorter than the spathe, 

flowers fertile throughout. Flowers bisexual, without perianth; stamens 4, free; ovary prismatic, truncate 

at apex, unilocular, with 2 (-8) basal or parietal ovules. Berries yellowish, 1- to 8-seeded; seeds reniform, 

smooth. Genus of 20 species native to tropical SE Asia, Australia, and western Pacific, with a few species 

cultivated and naturalized throughout the tropics. 

TYPE: Epipremnum mirabile Schott (≡ E. pinnatum (L.) Engl.). 

Key to the taxa of Epipremnum 

1. Leaves green, not variegated; pinnately dissected, or the juveniles simple; stems and petioles green, 

usually flowering and fruiting ……....................................................................…… 1. E. pinnatum 

1. Leaves variegated (green-yellow), simple or less often pinnately dissected; stems and petioles yelloworange; 

seldom flowering …………......................................................….. 2. E. pinnatum ‘Aureum’ 

1. Epipremnum pinnatum (L.) Engl., Pflanzenr. 

IV. 23B (Heft 37): 60. 1908; Pothos pinnatus 

L., Sp. Pl. ed. 2, 1374. 1763. Holotype: 

Ambon (Molucca Is.). Rumphius, Herb. 

Amboin. 5: 489, t. 183, f. 2. 1747. 

Philodendron nechodomae Britton in Britton & P. 

Wilson, Bot. Porto Rico 6: 335. 1926. [as 

“nechodomi”] Type: Puerto Rico. Britton & 

Boynton 8261 (holotype: NY!) 

Fig. 3. A-C 

Herbaceous root-climber, 6-8 m long. Stems 

flexible, cylindrical, green, to 3 cm in diam., 

producing scanty watery sap; bark papery. Leaves 

alternate, glabrous, green, not variegated, 

chartaceous, dimorphic; juvenile leaves simple, 

ascendant; adult leaves hanging, pinnately 

dissected, to 50 × 30 cm, the apex acuminate, the 

base asymmetric, sub-cordiform; lower surface 

with prominent venation, and a line of domatia 

along both sides of the midvein, and sometimes 

with pin-pricks; petioles to 35 cm long, marginate 

throughout. Inflorescence axillary, solitary, erect; 

spathe sub-cylindrical, to 18 cm long, fleshy, 

yellowish, opening along one side, acuminate at 

apex; spadix sessile, nearly cylindrical, 12-15 cm 

long. Berries not known. 

General distribution: Native to E India to SE 

Asia into Oceania. Occasionally cultivated 

throughout the tropics and becoming naturalized. 


Islands: Cultivated and perhaps naturalized in 

Puerto Rico, Dorado, Río Grande, and Utuado; 

naturalized on St. Thomas. 


Mayagüez: Federal Experimental Extension, 

Acevedo-Rdgz. 10838 (US). San Juan: Pennock’s 

Nursery, Hato Rey, cultivated, Howard & Nevling 

16921 (US). 

2. Epipremnum pinnatum ‘Aureum’ Nicolson, 

Allertonia 1: 347. 1978. 

Pothos aureus [as “aurea”] André, Ill. Hort. 27: 69 

t. 381. 1880; Epipremnum aureum (André) G. 

S. Bunting, Ann. Missouri Bot. Gard. 50: 28. 

1963. Type: Solomon Is. André, Ill. Hort. 27: 

69, t. 381. 1880. 

Fig. 3. D, E 

This cultivar is similar to the wild species 

except that it is usually more robust, reaching more 

than 10 m long, the stems and petioles are yelloworange, 

and the leaves are variegated with yellow. 

Juvenile leaves are 7-10 cm long and entire while 

the adult ones reach 35 cm in length, and are 

irregularly pinnatifid. Seldom collected in flower, 

fruits not known. 

General distribution: Originally reported 

from Solomon Islands but probably from


Fig. 3. A-C. Epipremnum pinnatum. A. Sterile branch. B. Leaf. C. Inflorescence. D-E. Epipremnum pinnatum ‘Aureum’. D. 

Sterile branch. E. Leaf. From Acevedo-Rdgz., P. 2003, Bejucos y plantas trepadoras de Puerto Rico e Islas Vírgenes, Smithsonian 

Institution.

36 


cultivation; cultivated and naturalized throughout 

the tropics. 

Distribution in Puerto Rico: Naturalized, 

common in moist secondary forests. Arecibo, 

Dorado, Jayuya, Río Grande, and Vega Baja. 

Common name: Puerto Rico: Bejuco de agua. 


Birdsey s.n. (US). Dorado: Bo. Higuillar, moist 

8. MONSTERA 

Monstera Adans., Fam. Pl. 2: 470. 1763, nom. conserv. 

secondary forest 0.8 km due NW of intersection, 

Roads 683 & 697, Proctor 48592 (SJ). Río 

Grande: along Road 191 to El Yunque Forest 

Reserve, Acevedo-Rdgz. & Siaca 9334 (UPR). 

Utuado; along Rt. 146, ca. 5 km from Dos Bocas 

Lake, Acevedo-Rdgz. et al. 7141 (HAC, UPRRP, 

US). ST. THOMAS: Peterborg, Acevedo-Rdgz. 

11260 (UPRRP, US). 

Robust or slender root climbers; stems smooth or verrucose. Leaves alternate; blades simple, entire 

or laciniate, usually perforated; petioles elongated and winged. Inflorescences 1-several per node. Spathe 

thickened, ovate, oblong-ovate, boat-shaped, not constricted, convolute at base, white to rose within, 

deciduous; spadix sessile, cylindrical, thickened, erect, little shorter than the spathe; lower flowers usually 

sterile. Flowers bisexual, lacking perianth; stamens 4, free; ovary prismatic, truncate at apex, bilocular, 

with 2 basal ovules per locule, the stigma punctate or linear. Berries green, white, yellow or orange; seeds 

1-3, obovoid or ellipsoid. Genus of approximately 40 species native to continental tropical America and 

the Lesser Antilles. 

TYPE: Monstera adansonii Schott (≡ Dracontium pertusum L.), typ. conserv. 

Reference: Madison, M. 1977. A revision of Monstera (Araceae). Contr. Gray Herb. 207: 3-100. 

1. Monstera adansonii Schott, Wiener Z. Kunst 

1830: 1028. 1830; Dracontium pertusum L., 

Sp. Pl. 967. 1753, non Monstera pertusa 

(Roxb.) Schott, 1830. Type: Martinique. 

Plumier, Descr. Pl. Amér., t. 56, 57. 1693. 

Fig. 4. A-C 

Glabrous, herbaceous, root-climber. Stems 

flexible, cylindrical, green, 0.5-2.5 cm in diam., 

producing scanty watery sap. Leaves alternate, 

simple, dimorphic; blade asymmetrically ellipticlanceolate, 

chartaceous, basal leaves not perforated, 

distal leaves perforated, 26-47 × 13-22.5 cm, the 

apex short- or long-acuminate, the base asymmetric, 

one side cuneate, the other obtuse; upper surface 

dull, with wide, flattened midvein; lower surface 

light green, with prominent midvein and darker 

lateral secondary veins; petioles winged almost to 

the apex, shorter than the blade. Inflorescence 

axillary, solitary, erect; spathe boat-shaped, to 20 

9. PHILODENDRON 

Philodendron Schott, Wiener Z. Kunst 1829: 780. 1829, nom. conserv. 

cm long, erect, adaxially cream or yellowish; 

spadix yellowish, ellipsoid, 8-13 cm long. Fruiting 

spadix yellow to orange. 

General distribution: Native to continental 

tropical America. 

Distribution in Puerto Rico: Exotic, cultivated 

and sometimes naturalized in moist forest 

understory. Río Grande. 


Río Grande: Bo. Jiménez, Finca San Pedrito, 

Proctor 50421 (SJ); Road 191 to El Yunque Forest 

Reserve, Acevedo-Rdgz. & Siaca 9313 (UPR, 

US). 


Monstera deliciosa Liebm., a more robust 

scandent or decumbent shrub, is sometimes 

cultivated in Puerto Rican gardens, but it is not 

known to grow in the wild. 

Stout or slender vines, climbing by means of adventitious roots, or less often erect herbs; stems 

elongate, usually producing a watery, caustic sap. Leaves simple, lobed or variously divided or pinnatifid,


Fig. 4. A-C. Monstera adansonii. A. Fertile branch. B. Leaf. C. Spathe and spadix. From Mori, S. et al. 1997. Vascular plants of 

central French Guiana. Mem. NYBG Vol. 76(1).

38 


long-petiolate. Inflorescence 1-11 per node. Spathe erect, convolute, surrounding the spadix, usually 

thickened, and constricted between the tube and the blade, not adnate to the spadix; spadix cylindrical, 

erect, usually nearly sessile, pistillate zone basal, separated from the staminate zone by a zone of sterile 

staminate flowers. Flowers unisexual, the perianth wanting; stamens 2-6; ovary 2- to many-locular, with 

1 to many ovules per locule, the stigma sessile, entire or lobed. Fruit a fleshy, 1- to many-seeded berry; 

seeds ovoid to ellipsoid. A genus of about 275 species native to the Neotropics. 

TYPE: Philodendron grandifolium (Jacq.) Schott (≡ Arum grandifolium Jacq.) 

Reference: Arias Granda, I. 1998. El género Philodendron Schott (Araceae) en Cuba. Feddes Repert. 

109: 33-39. 

Key to the species of Philodendron 

1. Petiole winged or margined nearly to the apical portion ……...............................…. 4. P. lingulatum 

1. Petioles not winged or winged only on the lower half ………................................................……… 2 

2. Leaves pinnatilobed at margins ………….........................................................……. [P. radiatum] 

2. Leaves with entire margins ………….......................................................................................…. 3 

3. Leaf blades 25-60 cm long; petioles 40-100 cm long; cataphylls persistent ……...…………. 4 

4. Stems 8-10 cm in diam.; spathe shortly stipitate, ca. 45 cm long, thick, abaxially green, 

adaxially white ……..........................................................................…….. 2. P. giganteum 

4. Stems 2-2.5 cm in diam.; spathe long-stipitate, ca. 18 cm long, coriaceous, abaxially whitish, 

adaxially burgundy ………………...............................................… 5. P. ornatum 

3. Leaf blades 13.5-35 cm long; petioles 10-35 cm long; cataphylls deciduous ……..………… 5 

5. Leaves oblong or lanceolate ……………......................................….. 1. P. consanguineum 

5. Leaves cordiform …………..................................................................…. 3. P. hederaceum 

1. Philodendron consanguineum Schott, Syn. 

Aroid. 88. 1856. Type: “S. Domingue”. 

Plumier, Descr. Pl. Amér. t. 51a, 52. 1693. 

Philodendron angustatum Schott, Syn. Aroid. 

103. 1856. Type: Haiti; Isle de la Tortue. 

Plumier, Descr. Pl. Amér. t. 51c & 54. 1693. 

Philodendron krebsii Schott, Bonplandia 7: 164. 

1859. Type: St. Thomas, U. S. Virgin Is. Krebs 

s. n. (holotype: C). 

Philodendron wrightii Griseb., Cat. Pl. Cub. 219. 

1866. Type: Cuba. Wright 3211 (holotype: 

GOET; isotypes: GH, MO, NY, US!). 

Philodendron marginatum Urb., Symb. Antill. 4: 

135. 1903. Type: Puerto Rico. Sintenis 4496b 

(syntype: B, destroyed); Sintenis 6927 

(syntype: B, destroyed). 

Philodendron fuertesii K. Krause in Urban, Symb. 

Antill. 7: 172. 1912. Type: Dominican 

Republic. Fuertes 1184 (holotype: B, 

destroyed). 

Philodendron urbanianum K. Krause in Urban, 

Symb. Antill. 7: 172. 1912. Type: Cuba. 

Baker & van Hermann 4263 (syntype: B, 

destroyed); Eggers 5143 (syntype: B ?). 

Fig. 5. A, B 

Herbaceous, glabrous root-climber, 10-30 m 

long. Stems flexible, cylindrical, reaching 2 cm in 

diam., producing scanty, watery, caustic sap; bark 

smooth, light brown or grayish, papery; nodes with 

an annular leaf scar. Leaves alternate, simple; 

blades 13.5-35 × 7.5-17.5 cm, oblong or less often 

lanceolate, coriaceous, lustrous, the apex acute or 

acuminate, the base cordiform, midvein stout, 

prominent on both surfaces; abaxial surface 

yellowish green with dark venation; petioles 

cylindrical, as long as the blade, winged along the 

lower ½. Inflorescence axillary, solitary; spathe 

green, erect, sub-cylindrical, with overlapping 

margins, 9-13 cm long; spadix sessile. 

General distribution: Cuba, Hispaniola, 

Puerto Rico and the Virgin Islands. 


Islands: In moist and wet forests. Adjuntas, 

Arecibo, Barranquitas, Bayamón, Cabo Rojo, 

Canóvanas, Dorado, Fajardo, Guayama, Guaynabo, 

Isabela, Lajas, Maricao, Ponce, Quebradillas, Río 

Grande, Sabana Grande, Salinas, Toa Baja, Vega 

Baja, and Yauco; St. Thomas. 

Common names: Puerto Rico: Bejuco de 

calabazón, Rasca garganta.


Fig. 5. A-B. Philodendron consanguineum. A. Branch. B. Inflorescence. C-D. Philodendron hederaceum. C. Fertile branch. D. 

Inflorescence, longitudinal section, with detail of the gynoecium and stamens. From Acevedo-Rdgz., P. 2003, Bejucos y plantas 

trepadoras de Puerto Rico e Islas Vírgenes, Smithsonian Institution.

40 



Adjuntas: Sintenis 4347 (US). Arecibo: Bo. 

Arrozal, NE of Road 637, km 6.4, Proctor & Díaz 

44774 (SJ). Barranquitas: Bo. Barrancas, upper 

NE slope & summit of Monte La Torrecilla, 

Proctor & Thomas 44454 (SJ). Bayamón: Bo. 

Guaraguao Arriba, upper slopes & summit of 

Cerro La Peña, Proctor & Foy 42323 (SJ). Cabo 

Rojo: Between Cabo Rojo and San Germán, N. L. 

Britton et al. 4313 (US). Canóvanas: Sierra de 

Luquillo, Caribbean Natl. Forest, Proctor & 

Taylor 46292 (SJ). Dorado: Bo. Higuillar, grounds 

of Dorado Beach Hotel, Proctor 45104 (SJ). 

Fajardo: Colima San Miguel, N.L. Britton & 

Shafer 1656 (US). Guayama: Bo. Carmen, along 

crest (S side) of Sierra de Jajome 2-2.8 km due NE- 

ENE of village of Carmen, Proctor & Rivera 

46941 (SJ, US). Guaynabo: Bo. Pueblo Nuevo, E 

end of Montes de Caneja, Proctor & Rivera 51113 

(SJ). Isabela: Bo. Coto, hwy 2, km 104.5, Proctor 

& Vives 45119 (SJ). Lajas: Limestone cliffs, 

Sargent 342 (US). Maricao: Bo. Indiera Fría, 

along track near Río Lajas below El Salto de Curet, 

Proctor & Padrón 45676 (SJ). Ponce: Road from 

Ponce to Adjuntas, Underwood & Griggs 753 

(US). Quebradillas: Bo. San Antonio, distrito 

Montedero, Proctor 46440 (SJ). Río Grande: 

Sierra de Luquillo, Proctor 50419 (SJ). Salinas: 

Bo. Lapa, near summit of E peak, Las Tetas de 

Cayey, Proctor & Thompson 44935 (SJ). Toa 

Baja: Bo. Candelaria, wooded mogotes 0.4-0.7 km 

WSW of Pico Nevárez, Proctor & Carrasquillo 

45305 (SJ). Vega Baja: Goll 1032 (US). Yauco: 

Bo. Sierra Alta, summit area of Pico Rodadero, 

Proctor & Díaz 44739 (SJ). 

2. Philodendron giganteum Schott, Syn. Aroid. 

89. 1856. Type: Unknown; representative 

material for this species and a good candidate 

for a lectotype of its name is Schott, Icones 

Aroideae, t. 2685 (NYBG neg. 4123). 

Fig. 6. A-D 

Erect, terrestrial or epiphytic plant, to 2 m tall, 

or a root-climber to 8 m long; stem cylindrical, 8- 

10 cm diam., producing scanty watery sap; 

cataphylls to 60 cm long, initially entire, 

weathering into persistent fibers. Leaf blades 

horizontal or directed downward, 25-60 × 17-50 

cm, lanceolate or triangular-lanceolate, nearly 

coriaceous, slightly paler below, the apex obtuse 

to acute or acuminate, the base cordate, with 

sinuses not overlapping, the margins sinuate; 

petioles erect, to 1 m long, nearly cylindrical. 

Inflorescence axillary, solitary; peduncles 6-9 cm 

long, stout; spathe 14-21 cm long, constricted at 

the middle, with convolute margins on lower ½, to 

form an abaxially reddish tinged tube, the blade 

adaxially cream or white, abaxially yellowish 

green, with two brown spots where the margins 

overlap; spadix sessile, stout, as long as the spathe, 

with staminate portion whitish and pistillate 

portion yellowish green. Berry yellow to orange. 


Rico, Virgin Islands, Lesser Antilles, Trinidad and 

Venezuela. 


Islands: Uncommon terrestrial or epiphytic herb 

of moist forests. Aguas Buenas, Arecibo, Cabo 

Rojo, Fajardo, Guaynabo, Las Piedras, Luquillo, 

Río Grande, Toa Baja, Vega Alta, and Yabucoa; 

St. John, St. Thomas, and Tortola. 


Aguas Buenas: Bo. Sonadora, S side of summit 

ridge, Cerro Marquesa, Proctor 43062 (SJ). 

Arecibo: Río Abajo State Forest, Proctor 41185 

(SJ). Cabo Rojo: between Cabo Rojo and San 

Germán, Britton et al. 4312 (US). Fajardo: 

Hillside forest, N.L. Britton & Shafer 1685 (US). 

Guaynabo: Bo. Sonadora, upper N slopes of Cerro 

Marquesa, Proctor 43085 (SJ). Las Piedras: Bo. 

Montones, Road 917, 0.2 km due NE of Escuela 

José D. Zayas, Proctor 41568 (US). Luquillo: El 

Yunque forest reserve, Visley, Acevedo-Rdgz. 

10806 (US). Toa Baja: Bo. Candelaria, 0.4-0.7 km 

WSW of Pico Nevárez, Proctor & Carrasquillo 

45304 (SJ).Vega Alta: Bo. Sabana, limestone hills 

NE of Regadera, Proctor 48503 (SJ). Yabucoa: 

Bo. Calabazas, hwy 3, appr. Km 101.5, Proctor & 

Rivera 46944 (SJ). ST. JOHN: Coral Bay Quarter; 

Bordeaux Mountain, Acevedo-Rdgz. 4705 (JBSD, 

MO, NY, UPR, US). TORTOLA: High Bush, N.L. 

Britton & Shafer 828 (US); Upper slopes of Mt. 

Sage, Proctor & Barwick 41978 (SJ). 

3. Philodendron hederaceum (Jacq.) Schott, 

Wiener Z. Kunst 1829: 780. 1829; Arum 

hederaceum Jacq., Enum. Syst. Pl. 31. 1760. 

Lectotype: Colombia; Cartagena. Jacquin, 

Select. Stirp. Amer. Hist., t. 152, 1763, 

designated by G. S. Bunting, Baileya 11: 62. 

1963.


Fig. 6. A-D. Philodendron giganteum. A. Non-climbing habit. B. Climbing habit. C. Leaf. D. Inflorescence, whole and longitudinal 

section. From Acevedo-Rdgz., P. 2003, Bejucos y plantas trepadoras de Puerto Rico e Islas Vírgenes, Smithsonian Institution.

42 


Philodendron scandens K. Koch & Sello in 

Klotzsch, App. Gen. Sp. Nov. 1853: 4. 1854. 

Type: A cultivated plant grown in Berlin (see 

G. S. Bunting, Gentes Herb. 10: 160. 1968, for 

discussion). 

Philodendron micans Klotzsch ex K. Koch in 

Klotzsch, App. Gen. Sp. Nov. 1854: 7. 1854- 

55. Type: A plant from the Neotropics, 

brought into cultivation in Germany by 

Warszewicz. 

Philodendron isertianum Schott, Prodr. Syst. 

Aroid. 242. 1860. Philodendron scandens 

subsp. isertianum (Schott) G. S. Bunting, 

Gentes Herb. 10: 165. 1968. Type: Martinique: 

Isert s. n. (holotype: C). 

Philodendron oxycardium sensu Britton & P. 

Wilson, Bot. Porto Rico 5: 124. 1923, non 

Schott, 1856. 

Figs. 5. C, D; 60. E 

Vine to 10 m long, rooting at nodes; stem 

cylindrical, 1-2 cm diam., producing watery, 

caustic sap; cataphylls to 12 cm long, deciduous. 

Leaf blades 14-30 × 10-20 cm, ovate, coriaceous, 

the apex acuminate to cuspidate, the base cordate 

with non-overlapping sinuses, the margins slightly 

sinuate; petioles curved to straight, 10-15 cm long, 

nearly cylindrical, invaginate at base. Inflorescence 

axillary, solitary, ascending; peduncles 5-9 cm 

long, stout; spathe ca. 15 cm long, thickened, with 

convolute margins, nearly cylindrical, abaxially 

green to yellow at maturity, adaxially red to 

maroon; spadix nearly sessile, cylindrical, stout, 

whitish, slightly shorter than the spathe. 

General distribution: Throughout tropical 

America. 


Islands: Common in moist forests of low and 

middle elevations. Fajardo, Guaynabo, Maricao, 

Río Grande, and Toa Baja; St. John, St. Thomas, 

and Tortola. 


calabazón, Paisaje. 


Fajardo: Hillside forest, N.L. Britton & Shafer 

1691 (US). Guaynabo: Bo. Sonadora, upper N 

slopes of Cerro Marquesa, Proctor 43086 (SJ). 

Maricao: Indiera Fría, N.L. Britton et al. 4489, 

4544 (US). Maricao: Bo. Montoso, Río Prieto 

valley, Breckon & Cedeño 4264 (US). Río Grande: 

Sierra de Luquillo, Proctor 50420 (SJ). Toa Baja: 

Bo. Candelaria, E sector, Mogotes de Nevárez, 

Proctor 45457 (SJ). ST. JOHN: Coral Bay Quarter; 

east side of Bordeaux Mountain, Acevedo-Rdgz. 

2610 (US, VINPS). ST. THOMAS: Estate Caret Bay, 

NW side of Crown Mountain, Proctor & Pinto 

40341 (SJ). TORTOLA: Pickate Ghut, W of Mt. 

Sage, Proctor & Barwick 41982 (SJ). High Bush, 

N.L. Britton & Shafer 826 (US). 

4. Philodendron lingulatum (L.) K. Koch in 

Klotzsch, App. Gen. Sp. Nov. 1855: 2. 1855- 

56; Arum lingulatum L., Syst. Nat. ed. 10, 

1251. 1759. Lectotype: Martinique. Plumier 

(Burman ed.), Pl. Amer. t. 37. 1756, 

designated by R.A. Howard, Fl. Lesser Antill. 

3: 392. 1979. 

Fig. 7. A 

Herbaceous, glabrous root-climber, to 15 m 

long. Stems flexible, cylindrical, dark green, 

sulcate along one side, 05-2.5 cm in diam., 

producing scanty watery sap; bark papery, peeling 

off. Leaves distichous; blades simple, 13-42 × 7- 

28 cm, elliptic or ovate, subcoriaceous, semilustrous, 

the apex acute or abruptly acuminate, the 

base cordiform, sub-cordiform, cuneate or 

truncate, midvein wide, prominent on lower 

surface, secondary venation light green; petiole 

winged for most of its length, 12-48 cm long. 

Inflorescence axillary, solitary or in pairs. Spathe 

10-23.5 × 2.9-5.3 cm, erect, the tube cylindricalurceolate, 

abaxially dark green, adaxially greenish 

or whitish; spadix 8.7-21.2 cm long, whitish or 

cream, the pistillate portion 2.1-4.7 cm long, the 

sterile portion 0.6-1.3 cm long, and the staminate 

portion to 12 cm long. 


Rico, Virgin Islands, and the Lesser Antilles. 

Distribution in Puerto Rico and the 

Virgin Islands: In moist and rain forests along the 

Central mountain range and in karst limestone. 

Adjuntas, Arecibo, Ciales, Naguabo, and Río 

Grande; Tortola. 


calabazón, Calabazón cimarrón. 


Ciales: along trail Camino de la Ceiba towards 

Quebrada del Pozo Azul, Acevedo-Rdgz. & Vicens 

11875 (UPRRP, US). Naguabo: Bo. de Maizales, 

N.L. Britton & Shafer 2139 (US). Río Grande: El 

Yunque, Sargent 559 (US); Sierra de Luquillo,


Fig. 7. A. Philodendron lingulatum, branch. B-C. Philodendron ornatum. B. Leaf, with detail of tubercles. C. Cataphylls. From 

Acevedo-Rdgz., P. 2003, Bejucos y plantas trepadoras de Puerto Rico e Islas Vírgenes, Smithsonian Institution.

44 


Sintenis 1746 (US). TORTOLA: Pickate Ghut, W of 

Mt. Sage, Proctor & Barwick 41981 (SJ). 

5. Philodendron ornatum Schott, Oesterr. Bot. 

Wochenbl. 3: 378. 1853. Type: Unknown; 

representative material of this species and a 

good candidate for a lectotype of its name is 

Schott, Icones Aroideae, t. 2453 (NYBG neg. 

4116). 

Fig. 7. B, C 

Herbaceous, glabrous, root climber, 2-10 m 

long. Stems cylindrical, 2-2.5 cm in diam., 

smooth, producing scanty watery, caustic sap, the 

nodes with an annular scar. Cataphylls ca. 24 cm 

long, persistent. Leaves distichous; blades 32-60 × 

24-46 cm, cordiform, chartaceous, glabrous, the 

apex acuminate, the base cordiform, the margins 

undulate, slightly revolute, upper surface dull, 

dark green, with sunken venation, lower surface 

light green, with prominent, reddish venation; 

petioles ascending, 14-42 cm long, ventrally 

compressed, distally pulvinate, the pulvinus 

tuberculate. Inflorescence axillary, in groups of 3, 

ascending; peduncles robust, 11-13 cm long. 

Pistia L., Sp. Pl. 963. 1753. 

10. PISTIA 

A unispecific genus, characterized by the following species. 

TYPE: Pistia stratiotes L. 

1. Pistia stratiotes L., Sp. Pl. 963. 1753. 

Lectotype: India. Rheede, Hort. Malab. 11: t. 

32. 1692, designated by Suresh et al., Taxon 

32: 127. 1983. 

Pistia occidentalis Blume, Rumphia 1: 79. 1835. 

Lectotype: Madeira or Barbados. Sloane, 

Voy. Jamaica 1: t. 2, f. 2. 1707, here 

designated. 

Fig. 8. A-C 

Floating acaulescent, stoloniferous herb, 

forming large colonies. Leaves subsessile, in a 

rosette; blades 3-12 × 1.5-5 cm, cuneate, fleshy, 

with sunken parallel veins, impermeably pubescent, 

the apex rounded, truncate, usually notched, the 

base cuneate, the margins revolute. Inflorescence 

Spathe persistent, 16-18 cm long, fleshy, 

convolute, sub-infundibuliform, abaxially whitish 

with pinkish hue, adaxially burgundy, aristate at 

apex; spadix cylindrical-ellipsoid, robust, cream, 

almost as long as the spathe. 

General distribution: Venezuela, Trinidad, 

Tobago, Surinam and Brazil. Introduced elsewhere. 

Distribution in Puerto Rico: Naturalized 

along the lower part of the Guajataca Gorge. 


Guajataca: Guajataca River, Woodbury s.n.(SJ). 

Quebradillas: Bo. Cacao, along E side of Río 

Guajataca at bottom of canyon, Proctor 50206 

(SJ). San Sebastián: Bo. Cibao, Río Guajataca 

gorge, Proctor 47073 (SJ). 


Philodendron bipinnatifidum Schott ex Endl. 

with large deeply pinnatifid leaves, is sometimes 

cultivated as an ornamental in Puerto Rico (Hato 

Rey, Pennock’s Nursery, Howard & Nevling 

16928, US); Philodendron radiatum Schott is also 

cultivated in Puerto Rico and has been collected as 

a persistent plant in abandoned fields (Utuado: Bo. 

Don Alonso, Acevedo-Rdgz. 13411, MAPR, US). 

axillary, solitary, ascending; peduncles 5-7 mm 

long, slender; spathe 1.3-1.5 cm long, convolute 

and adnate to the spadix below, spreading above, 

whitish; spadix with a single pistillate flower at 

base, and with 2-8 staminate flowers above, 

shorter than the spathe. Flowers unisexual, the 

perianth wanting; stamens 2; ovary 1-locular, with 

numerous ovules, the style slender, the stigma 

penicillate. Fruit thin-walled, many-seeded. Seeds 

cylindrical, rugulose. 

General distribution: A variable species 

found throughout tropical and subtropical regions. 


Islands: In calm, temporary or permanent 

freshwater bodies such as rivers, lakes, ponds, 

cisterns or pools, sometimes cultivated. Arecibo,


Fig. 8. A-C. Pistia stratiotes. A. Habit. B. Inflorescence, l.s. inflorescence showing staminate and pistillate flowers, and l.s. pistil 

showing ovules. C. Seed. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78. 

Arroyo, Barceloneta, Cabo Rojo, Gurabo, Lajas, 

Loíza, Toa Alta, and Vega Alta, Vega Baja; St. 

John and St. Thomas. 

Common names: Puerto Rico: Lechuga de río, 

Lechuguilla de río. 


Arecibo: E.G. Britton 5094 (US). Arroyo: Bo. 

Palmas, recently-established, temporary “lake” 

along road to Punta Guilarte, Proctor et al. 46371 

(SJ). Barceloneta: Bo. Garrochales, area of Caño 

Matos, Proctor et al. 46144 (SJ). Cabo Rojo: 

Sintenis 831 (US). Gurabo: Bo. Celada, Road 941 

at W end of Río Gurabo bridge, Proctor 42248 

11. SPATHIPHYLLUM 

Spathiphyllum Schott in Schott & Endicher, Melet. Bot. 22. 1832. 

(SJ). Lajas: Bo. Sabana Yeguas, 0.3 km E of Road 

116 and 1.2 km NE of Finca Juanita, Proctor 

46387 (SJ). Loíza: Bo. Torrecilla Alta, 0.2-1.2 km 

NNE of NE corner of Santa Barbara urbanization, 

Proctor & Colón 50494 (SJ). Manatí: Sintenis 

6798 (US). Toa Alta: Bo. Contorno, Road 861, km 

1.1, beside Río de la Plata below hwy bridge, 

Proctor & Rivera 47441 (SJ). Vega Alta: between 

Dorado and Vega Baja, Strong et al. 405 (GMUF). 

Vega Baja: Bo. Cibuco, along Río Cibuco ca. 1 km 

due SE of Cerro Cibuco, Proctor et al. 45627 (SJ). 

ST. JOHN: Virgin Is. Natl. Park: just SE of Peter 

Peak, Proctor & Pinto 40428 (SJ). 

Terrestrial herbs; stems short, erect or creeping (seemingly acaulescent). Leaves in a loose rosette, 

simple, long-petiolate, the petiole geniculate at apex, with a long sheath. Inflorescence solitary, erect; 

peduncle shorter than the petiole; spathe herbaceous, not enclosing the spadix, ascending, marcescent, 

white, with distinct midvein and pinnate laterals; spadix sessile or short-stipitate, cylindrical, 

flowering from base to apex. Flowers bisexual, the perianth segments 4-6, free or connate into a short cup; 

stamens 4-6, free; ovary (2-) 3 (-4) -locular, with 2-8 axile ovules per locule, the style long, the stigma 2or 

3-lobed or sub-capitate or punctiform. Berry 1- to 8-seeded, green; seeds oblong, ellipsoid or ovoid. A 

genus of about 41 species of Central America, northern South America, and the eastern Malay 

archipelago.

46 


TYPE: Spathiphyllum lancifolium (Jacq.) Schott (≡ Dracontium lancifolium Jacq. [as lancaefolium]). 

Reference: Bunting, G. S. 1960. A revision of Spathiphyllum (Araceae). Mem. New York Bot. Gard. 

10: 1-53. 

1. Spathiphyllum wallisii Regel, Gartenflora 26: 

323. 1877. Lectotype: Venezuela. Regel, 

Gartenflora 26: t. 920. 1877, here designated. 

Glabrous, erect, terrestrial herb to 50 cm tall; 

rhizome very short. Leaves in a loose rosette; 

blades ascending, 12-22 × 2.3-4 cm, elliptic, 

chartaceous, midvein stout, prominent underneath, 

the secondaries slightly prominent, the apex long 

acuminate, the base attenuate, slightly 

asymmetrical, the margins entire, slightly wavy; 

petioles erect, nearly cylindrical, 13-25 cm long, 

narrowly winged throughout its extension. 

12. SYNGONIUM 

Syngonium Schott, Wiener Z. Kunst 1829: 780. 1829. 

Inflorescence ascending; peduncles to 45 cm long; 

spathe ascending, green, leafy, oblong-lanceolate, 

ca. 11 cm long, acuminate at apex; spadix 

ascending, long-stipitate, ca. 3 cm long. Berries 

green, conical, ca. 4 mm long, produced 

throughout the spadix. 

General distribution: Native of Venezuela 

and Colombia (?) introduced into the West Indies. 

Distribution in Puerto Rico: Cultivated, an 

escaped population found in Adjuntas: Cordillera 

Central, Bo. Portugués ca. 0.9 km due SW of Alto 

de la Bandera, Proctor 42130 (SJ). 

Epiphytes or hemi epiphytes, usually with long, root-climbing stems, producing milky sap. Leaves 

simple or variously divided, with 5-11 leaflets; petioles sheathed toward the base. Inflorescences 1-11 per 

axil; peduncles erect in flower, pendant in fruit; spathe fleshy, convolute, conspicuously constricted 

medially, the tube ellipsoid, the blade whitish to greenish, broadly spreading at anthesis; spadix much 

shorter than the spathe, erect, with pistillate flowers on basal portion. Flowers unisexual, the perianth 

wanting; stamens 3-4, united into a synandrium; ovary (1-) 2 (-3)-locular, with 1 (-2) ovules per locule, the 

stigma discoid or bilabiate. Fruit a 1-seeded berry, connate into an ovoid syncarp; seeds obovoid or ovoid. 

A genus of 33 species native to the Neotropics with most species in Costa Rica and Panama. 

TYPE: Syngonium auritum (L.) Schott (≡ Arum auritum L.) 

References: Birdsey, M. 1955. The morphology and taxonomy of the genus Syngonium (Araceae). 

Ph. D. dissertation, University of California, Berkeley. 382 pp. Croat, T. B. 1981. A revision of Syngonium 

(Araceae). Ann. Missouri Bot. Gard. 68: 565-651. 

Key to the species of Syngonium 

1. Leaves trisect, sub-5-pedatisect or rarely 5-pedatisect; leaflets 3(5), acute at apex ….... [S. auritum] 

1. Leaves pedately divided; leaflets 3-11, acuminate at apex ………….................... 1. S. podophyllum 

1. Syngonium podophyllum Schott, Bot. Zeitung 

(Berlin) 9: 85. 1851. Type: Unknown; 

representative material for this species and a 



4329). 

Figs. 9. A-C; 60. F 

Vine to 10 m long, rooting at nodes; stem 

cylindrical, glaucous, 1-2 cm diam., producing 

abundant milky sap. Leaves pedately divided; 

leaflets 3-11, united or free to base, coriaceous, the 

apex acuminate, the base variously auriculate, the 

margins sinuate, outermost leaflets smaller, the 

medial leaflets 16-38 × 6-17 cm, obovate, elliptic


Fig. 9. A-C. Syngonium podophyllum. A. Fertile branch. B. Leaf. C. Inflorescence with detail of synandrium. From Mori, S. et 

al. 1997. Vascular plants of central French Guiana. Mem. NYBG Vol. 76(1).

48 


or lanceolate; petioles 15-60 cm long, nearly 

cylindrical, sheathed of their length. Inflorescences 

4-11 per axil, ascending; peduncles 8-9 cm long, 

slender; spathe ca. 10 cm long, convolute into an 

ellipsoid tube at base, the blade cream-colored, 

concave, ephemeral; spadix whitish, sessile, 

cylindrical with a constriction between the 

pistillate and the staminate areas. Syncarp ovoid, 

red, reddish orange or yellow, 3-5.5 cm long. 

General distribution: Native to Mexico but 

grown as an ornamental and naturalized throughout 

the Caribbean. Also, now throughout tropical 

America and Florida (U.S.A.). 


Islands: Becoming common throughout moist 

disturbed habitats. Ceiba, Ciales, Luquillo, Río 

Grande, Utuado, Vega Baja, and Yabucoa; St. 


Common name: Puerto Rico: Malanga 

trepadora. 


Ceiba: Bo. Chupacallos, former quarry area along 

Quebrada Aguas Claras, 1.45 km due WSW of 

plaza in Ceiba town center, Proctor & Colón 

50190 (SJ). Ciales: Bo. Frontón, Road 140, km 49, 

13. XANTHOSOMA 

Xanthosoma Schott in Schott & Endicher, Melet. Bot. 19. 1832. 

Proctor et al. 51149 (SJ). Luquillo: Bo. Mameyes 

I, Fortuna area, E side of the Río Mameyes near its 

mouth, Proctor & Torres 49244 (SJ). Río Grande: 

Bo. Herreras: just N of Caño San Luis c. 0.2 km W 

of Road 187, Proctor & Morris 45994 (SJ). 

Utuado: Bo. Don Alonso, Acevedo-Rdgz. et al. 

7122 (UPRRP, US).Vega Baja: Bo. Cibuco, 

border of mangrove thicket 1.2 km due ENE of 

Cerro Cibuco, Proctor et al. 45631 (SJ). Yabucoa: 

Bo. Calabazas, hwy 3, ca. Km 101.5, Proctor & 

Rivera 46943 (SJ). ST. JOHN: Cruz Bay Quarter; 

Susannaberg, Acevedo-Rdgz. & Angell 4061 (MO, 

NY, UPR, US). 


Syngonium auritum (L.) Schott was reported 

by Liogier and Martorell (Fl. Puerto Rico, Syst. 

Synop. 2000) as occurring in shady forests at low 

elevations. However, we have not been able to 

locate a specimen to verify this record, nor have we 

collected it in naturalized conditions. The species 

has been collected however, persisting on an 

abandoned farm in Utuado: Bo. Don Alonso, 

Acevedo-Rdgz. 13410 (MAPR, US). 

Small to large herbs, the stems subterranean and tuberous with smaller tubers or stolons, or epigeal 

and massive; producing milky sap. Leaves simple or dissected, several in a rosette in acaulescent species 

or terminal in arborescent species, cordate or sagittate at base, secondary veins forming a poorly organized 

collective vein between the primary lateral veins; petiole long, with distinct, elongated sheath. 

Inflorescence pedunculate, 1 to many per node. Spathe erect, strongly constricted, the basal area forming 

a convolute, tube that surrounds the spadix, the marcescent and deciduous blade longer than the tube, boatshaped, 

oblong to oblong-lanceolate, gaping and erect or sometimes reflexed; spadix erect, shorter than 

the spathe, with a basal pistillate zone, separated from the distal staminate zone by a narrower zone of 

staminate sterile flowers. Flowers unisexual, lacking perianth; stamens 4-6, connate into a truncate 

synandrium; ovary ovoid, (1)2-4-locular, with numerous parietal to basally axial ovules, the style broader 

than the ovary, coherent or weakly connate with those of neighboring flowers, the stigma sub-capitate or 

2-lobed. Berry cylindrical, white or orange, many-seeded; seeds ovoid, costate. A genus of about 57 

species native to tropical America, including the West Indies. Several species cultivated throughout the 

tropics for their edible starchy tubers and leaves. 

LECTOTYPE: Xanthosoma sagittifolium (L.) Schott (≡ Arum sagittifolium L. [as sagittaefolium]), 

designated by Nicolson, Taxon 24: 345-347. 1975. 

Note: As commonly happens with cultivated species, the taxonomy of Xanthosoma is notoriously 

difficult. The collections available are inadequate as the plants do not preserve well and usually are not 

representative, even of the individual collected. We are basically following the treatments of previous 

workers in the region, most of which stem from the work of Engler & Krause in 1920. A revision of this 

important genus is badly needed.


Key to the species of Xanthosoma 

1. Leaves entire, sagittate or nearly lanceolate, without lateral lobes …….........................................… 2 

2. Petiole’s invagination with revolute margins; petioles and lower surface of leaf purplish tinged; 

basal lobes of the leaf subtriangular ………......................................…. 5. X. violaceum 

2. Petiole’s invagination with involute or straight margins; petioles and lower surface of leaf light 

green; basal lobes of the leaf sub-rhomboid, obtuse ……….............................................…… 3 

3. Leaf with 8-9 basicopic secondary veins on basal lobes, these forming a 75-80° with the 

primary vein; spathe with green tube and abaxially white blade …….....3. X. sagittifolium 

3. Leaf with ca. 12 basicopic secondary veins on basal lobes, these forming a 45-60° with the 

primary vein; spathe with maroon tube and abaxially rosy blade …........ 4. X. undipes 

1. Leaves pedately dissected or with two large lateral lobes ……….............................................…….. 4 

4. Leaves with two divergent basal lobes …………….......................................… 1. X. brasiliense 

4. Leaves pedately dissected into 5-12 segments ………….......................…. 2. X. helleborifolium 

1. Xanthosoma brasiliense (Desf.) Engl., 

Pflanzenr. IV. 23E (Heft 71): 58. 1920; 

Caladium brasiliense Desf., Tabl. Ecole Bot., 

ed. 3, 386. 1829. Type: Not known, a 

cultivated plant grown in Paris, (FI?). 

Acontias hastifolius Schott in Schott & Endlicher, 

Melet. Bot. 19. 1832. Type: Unknown; 

representative material for this species and a 


Schott, Icones Aroideae, t. 3450 (NYBG Neg. 

4295). 

Acontias cubensis sensu Bello, Anales Soc. Esp. 

Hist. Nat. 12: 115. 1883, non Schott, 1859. 

Xanthosoma hastatum Eggers, Fl. St. Croix 99. 

1879*. Type: St. Croix, U.S. Virgin Islands. 

Eggers ? (C?). 

Glabrous, erect, acaulescent herb, with 

slender base. Leaves 1-3; blades nearly horizontal, 

21-40 × 11-25 cm, hastate, chartaceous, upper 

surface green, lower surface pruinose, the apex 

obtuse or less often shortly acuminate, the basal 

lobes divergent, oblong-lanceolate, 8-25 cm long, 

the margins undulate; petioles erect, 22-45 cm 

long, green, terete except for the invaginate lower 

1 /3. Inflorescences axillary, ascending; peduncles 

to 32 cm long; spathe coriaceous, ca. 28 cm long, 

the tube 8-10 cm long, green, oblong-ovoid, the 

blade elliptic to lanceolate, erect, concave, 

abaxially greenish, with network of veins, 

acuminate at apex; spadix stipitate, shorter than 

the spathe, the pistillate zone ca. 3 cm long, conical 

or subconical, the sterile staminate zone ca. 4 cm 

long, the fertile staminate zone ca. 12 cm long. 

General distribution: Central and South 

America, Cuba, Puerto Rico, Virgin Islands and 

the Lesser Antilles. 

Distribution in Puerto Rico and the Virigin 

Islands: Marshy grounds, and disturbed, moist 

edges of forests. Arecibo, Coamo, Mayagüez, San 

Juan, and Toa Baja; reported by Eggers (1879) as 

naturalized on all islands of the Virgin Islands, not 

collected there in recent years. 

Note: * Questionably validly published, i.e., 

not labelled as “n. sp.” 



10685 (US). Coamo: Sintenis 3222 (US). 

Mayagüez: Cowell 643, 645 (US). San Juan: Río 

Piedras, Stevenson 2496 (US). Toa Baja: Bo. 

Candelaria, 0.4-0.7 km WSW of Pico Nevárez, 

Proctor 45337 (SJ-2); 45337a (US). 

2. Xanthosoma helleborifolium (Jacq.) Schott, 

Oesterr. Bot. Z. 15: 33. 1865; Arum 

helleborifolium Jacq., Collectanea 3: 217. 

1789; Acontias helleborifolius (Jacq.) Schott 

in Schott & Endicher, Melet. Bot. 19. 1832. 

Lectotype: Venezuela. Jacq., Ic. Pl. Rar. pl. 

613. 1792, here designated. 

Xanthosoma hastifolium K. Koch, App. Gen. Sp. 

Nov. 1854: 2. 1854-55. Type: Unknown, a 

cultivated plant grown in Berlin. 

Xanthosoma angustisectum Engl., Arac. Exsicc. et 

illustr. 88. 1884; Xanthosoma helleborifolium 

var. angustisectum (Engl.) Engl., Pflanzenr. 

IV. 23E (Heft 71): 61. 1920. Type: Puerto 

Rico. Grosourdy, s. n. (holotype: P, photo at 

US)

50 


Glabrous, erect, acaulescent herb, with 

swollen base. Leaves 1-2; blades 20-90 × 15-35 

cm, in outline, pedately dissected into 5-12 

segments, chartaceous, upper surface green, lower 

surface pruinose, the segments oblong to 

oblanceolate, acute or acuminate at apex, the 

margins undulate; petioles erect, 22-50 cm long, 

green with dark green banding, invaginate on 

lower ¼. Inflorescences axillary, ascending, 

solitary; peduncles 10-15 cm long; spathe 

chartaceous, 11-14 cm long, the tube 4-5.5 cm 

long, green, ellipsoid, the blade elliptic to 

lanceolate, erect, concave, yellowish green, 

acuminate at apex; spadix shortly stipitate, slightly 

shorter than the spathe, the pistillate zone 

cylindrical, the sterile staminate zone conical, the 

fertile staminate zone ellipsoid. 

General distribution: Central and South 

America, Puerto Rico and the Lesser Antilles. 

Distribution in Puerto Rico: Naturalized, 

known from Añasco, Mayagüez, and San Juan. 


San Juan: Río Piedras, Stevenson 2496 (US). 

3. Xanthosoma sagittifolium (L.) Schott in 

Schott & Endicher, Melet. Bot. 19. 1832; 

Arum sagittifolium L., Sp. Pl. 966. 1753. 

Lectotype: Jamaica: Sloane, Voy. Jamaica 1: 

t. 106, f. 2. 1707, designated by Nicolson in 

A.C. Sm., Fl. Vit. Nov. 1: 459. 1979. 

Xanthosoma atrovirens K. Koch & Bouché, App. 

Gen. Sp. Nov. 1854: 3. 1854-55. Lectotype: 

Jamaica: Sloane, Voy. Jamaica 1: t. 106, f. 2. 

1707, here designated. 

Fig. 59. G, H 

Glabrous, erect, herb to 2 m tall, acaulescent 

when young, mature plants with a thick, erect, 

fleshy stem to 1 m long, these with numerous leaf 

scars and sometimes with aerial roots, the base 

enlarged, ovoid, producing lateral, elongated 

subterranean, edible tubers. Leaves several, nearly 

in a rosette in acaulescent plants, or in a distal 

crown in mature plants; blades horizontal to 

slightly nodding, with the posterior lobes 

ascending, 40-100 × 40-70 cm, simple, sagittateovate 

or subcordate, chartaceous, upper surface 

dark green with light green primary and secondary 

veins, lower surface light green, pruinose, with 

dark green venation, the apex obtuse, ending in an 

acute point, the base cordate with non-overlapping 

lobes, the lowest pair of secondary veins 

surrounded by marginal tissue at their insertion 

with the petiole, the margins undulate; petioles 

erect, 1-1.5 m long, green, invaginate on lower 

2 /3 , with straight, wavy or sometimes involute 

margins. Inflorescences 1-3, axillary, ascending; 

peduncles to 20 cm long; spathe chartaceous, 13- 

15 cm long, the tube 6-7 cm long, grayish green, 

oblong-ovoid, the blade elliptic, erect, concave, 

adaxially cream to white, shortly acuminate at 

apex; spadix slightly shorter than the spathe, the 

pistillate zone cylindrical, the sterile staminate 

zone conical, pinkish, the fertile staminate zone 

elongated, ellipsoid, cream. 

General distribution: Native to the Neotropics, 

cultivated throughout the tropics for its edible 

starchy tubers. 

Distribution in Puerto Rico: Cultivated and 

naturalized in moist or wet, disturbed vegetations. 

Arecibo, Salinas, and Utuado. Cultivated in St. 


Common names: Puerto Rico: Yautía, Yautía 

blanca, Yautía amarilla. 


Arecibo: Río Abajo State Forest, along Juan Ruiz 

trail, Acevedo-Rdgz. 10684 (US); Along Rd. 123, 

Acevedo-Rdgz. 13414 (US). Salinas: Bo. Lapa, 

vicinity of Las Tetas de Cayey, Proctor 45303 

(SJ). 

4. Xanthosoma undipes (K. Koch) K. Koch, 

Bonplandia 4: 3. 1856; Alocasia undipes K. 

Koch, App. Gen. Sp. Nov. 1854: 5. 1854-55. 

Type: Cultivated material grown in Berlin and 

Potsdam (Sanssouci) [probably destroyed]. 

Xanthosoma jacquinii sensu Schott, Syn. Aroid. 

57. 1856, non Schott in Schott & Endicher, 

Melet. Bot. 19. 1832. nom. illeg., non Kunth 

1841. 

Figs. 59. I; 60. A, B, D 

Glabrous, erect herb to 2 m tall, producing 

milky sap, acaulescent when young, mature plants 

with a thick, decumbent, starchy stem 1-1.5 m 

long, with numerous leaf scars and aerial roots at 

the base. Leaves several, in a distal crown in 

mature plants; blades horizontal to nodding, with 

the posterior lobes ascending, 40-70 × 30-50 cm, 

simple, cordate-sagittate, chartaceous, upper 

surface dark green, lower surface grayish green, 

pruinose, the apex widely obtuse, mucronate, the 

base cordate with wide sinus, the lowest pair of 

secondary veins naked (not surrounded by


marginal tissue) at their insertion with the petiole, 


long, green, invaginate on lower 1 /3-½, with 

involute margins. Inflorescences axillary, in 

groups of 3, ascending; peduncles to 30 cm long; 

spathe 25-31 cm long, the tube ellipsoid, 11-13 cm 

long, thickened, abaxially and adaxially burgundy, 

the blade ephemeral, ovate to elliptic, chartaceous, 

erect, concave, adaxially whitish, abaxially deeppink, 

acuminate at apex; spadix shorter than the 

spathe, the pistillate zone (basal), yellowish, 3-4 

cm long, conical, the sterile zone pink, fertile 

staminate zone (distal) light pink, ellipsoid, 12-14 

cm long. Berries not collected in Puerto Rico. 

General distribution: Native to tropical 

America. Known from Central America, South 

America, Puerto Rico and the Lesser Antilles 

Distribution in Puerto Rico: In moist 

disturbed habitats. Arecibo, Florida, Isabela, 

Luquillo, Mayagüez, Ponce, Quebradillas, Río 

Grande, and Salinas. 

Common names: Puerto Rico: Yautía rascana, 

Yautía de palma. 


Arecibo: Bo. Río Arriba, just E of Río Abajo State 

Forest, Proctor et al. 49497 (SJ-3); Río Arriba, ca. 

2 km N of Dos Bocas lake, along Rd. 123 (old road 

10), Acevedo-Rdgz. 13413 (US). Isabela: Bo. 

Coto, Highway 2, km 105.3, Proctor 45967 (SJ). 

Quebradillas: Bo. Cacao, along E side of Río 

Guajataca, Proctor 50209 (SJ-2). Río Grande: 

Guzmán Arriba, Acevedo-Rdgz. 13752 (US). 

Salinas: Bo. Lapa, vicinity of Las Tetas de Cayey, 


5. Xanthosoma violaceum Schott, Oesterr. Bot. 

Wochenbl. 3: 370. 1853. Type: Unknown; 

representative material of this species and a 



4288). Possibly from cultivated material 

collected by Schott in Brazil (1817-1821). 

Arum nigrum Vell., Fl. Flumin. 9. t. 107. 1831 

[“1827”], nom. nud.*; Arq. Mus. Nac. Rio de 

Janeiro 5: 386. 1881, non Schott 1857; 

Xanthosoma nigrum Stellfeld., Tribuna Farm. 

12: 201. 1944. Type: Brazil. Vellozo, Fl. 

Flumin. 9, t. 107. 1827 [1831]. 

Fig. 60. C 

Glabrous herb, acaulescent when young, 

developing a thick, starchy caudex (> 1 m long) 

with age. Leaves few, in a loose rosette when 

acaulescent or in a distal crown in mature plants; 

blades pointing downward or less often horizontal, 

20-65 × 15-40 cm, simple, sagittate-ovate, 

chartaceous, upper surface green, lower surface 

purple-tinged, especially along veins, the apex 

obtuse, with short acumen, the base cordate with 

non-overlapping or slightly overlapping sinuses, 

the lowest pair of secondary veins surrounded by 

marginal tissue at their insertion with the petiole, 


long, purple-black, invaginate on lower ½, with 

revolute margins. Inflorescences axillary, 

ascending, 6-8 clustered; peduncles to 20 cm long; 

spathe chartaceous, 30-40 cm long, the tube 5-15 

cm long, abaxially greenish, adaxially yellowish, 

ellipsoid-ovoid, the blade ovate or oblonglanceolate, 

erect, concave, adaxially yellowish to 

whitish abaxially, yellowish to yellowish brown, 

shortly acuminate at apex; spadix slightly shorter 

than the spathe. 

General distribution: Central America, South 

America, and the West Indies. 


naturalized. In moist disturbed habitats. Adjuntas, 

Arecibo, Luquillo, and Río Grande. 

Common names: Puerto Rico: Yautía 

morada, Yautía lila. 

Note: * When volume 9 was published in 

1831, it did not contain plant descriptions, and 

because figure 107, representing Arum nigrum 

did not contain any analytical sub-figures, the 

name is invalidly published (art. 44. 1 ICBN). 


Arecibo: Bo. Sabana Hoyos, Acevedo-Rdgz. et al. 

13370 (MAPR, UPRRP, US). Río Grande: Sierra 

de Luquillo, Sintenis 1587 (US); Guzmán Arriba, 

Acevedo-Rdgz. 13753 (US); Luquillo Mts., along 

service road to El Yunque Peak, Acevedo-Rdgz. 

14453 (UPRRP, US). 


Xanthosoma caracu K. Koch & Bouché, 

known as Rollisa blanca or Manola, was reported 

as cultivated in Puerto Rico by Engler & Krause 

(1920).

52 


Cultivated Genera 

The calla lily (Zantedeschia aetiopica (L.) Link) is cultivated in the central mountains of Puerto Rico 

for cut flowers. This operation exists only on a very small scale. Aglaonema commutatum Schott, 

Aglaonema pictum Kunth, and Calla palustris L. were cited by Martorell et al. (1981) as cultivated in 

Puerto Rico. 

Excluded Genus 

Montrichardia arborescens (L.) Schott, has been treated as occurring in Puerto Rico by Urban (Symb. 

Antill. 4: 133-134. 1903) and by Britton & P. Wilson (Bot. Porto Rico 5: 125. 1923) based on a report by 

Bello (Anales Soc. Esp. Hist. Nat. 12: 115. 1883). More recently it has been reported by Liogier and 

Martorell (Fl. Puerto Rico, Syst. Synop., 2000) as occurring along borders of the Río Blanco in Naguabo. 

However, based on the fact that we have not been able to locate any collections of this species for Puerto 

Rico, and that the species is otherwise known from South America north only to Guadeloupe, we conclude 

that the species does not occur in our area. Bello’s record was apparently a misidentification of another 

shrubby Araceae occurring in Puerto Rico. The fact that he provided the common name Yautía madera for 

this species in Puerto Rico suggests that whatever he identified as such may have been relatively common. 

In studying Agustin Stahl’s unpublished watercolors, we came across an illustration of Philodendron 

giganteum bearing the common name Yautia madera. The two men knew each other, and it is quite 

possible that Bello’s Montrichardia referred to Stahl’s illustration of Yautia madera, especially since 

these species are superficially similar. 

Family 2. LEMNACEAE Duckweed Family 

Lemnaceae Gray, Nat. Arr. Brit. Pl. 2: 729. 1821, nom. conserv. 

by G. R. Proctor 

Monoecious or rarely dioecious, small to minute aquatic, leafless, annual herbs, floating on or 

submerged in fresh to slightly brackish water. Vegetative part consisting of a thallus, either solitary or 

connected in small clusters, symmetric or asymmetric, flat or minutely hemispheric, varying in outline 

from reniform, round, elliptic, oblong or lanceolate, usually green but red or brown pigments sometimes 

also present; roots several, one per frond, or wanting. Vegetative propagation frequent from budding 

pouches at margins of frond. Flowers also produced in small pouches or cavities, 1 or 2 per frond; 

inflorescence with or without a spathe, otherwise consisting of a pistillate and 1 or 2 staminate flowers. 

Staminate flower consisting of a single stamen, the anther 1- or 2-locular. Pistillate flower consisting of 

a sessile globular, 1-locular ovary with a short apical style. Ovules 1-4. Fruit a 1- to 4-seeded utricle; seeds 

ribbed or smooth. A small cosmopolitan family of 5 genera and 32 species. These are the world’s smallest 

flowering plants. 

TYPE: Lemna L. 

Reference: Landolt, E. 1986. The family Lemnaceae -a monographic study. Veröff. Geobot. Inst. 

ETH Stiftung Rübel Zürich 71: 7-566. Les, D.H. & D.J. Crawford. 1999. Landoltia (Lemnaceae), a new 

genus of Duckweeds. Taxon. 9: 530-533. 


1. Roots present ……......................................................................................................................…….. 2 

2. Each thallus with a single root ……..............................................................................…. 1. Lemna 

2. Each thallus with several roots (2-21) …........................................................……….. 2. Spirodela 

1. Roots absent ……….......................................................................................................................….. 3


3. Thallus floating, very minute, hemispheric or globular ………...............................…… 3. Wolffia 

3. Plants submerged, flat, membranous ……............................................................…… 4. Wolffiella 

Lemna L., Sp. Pl. 970. 1753, nom. conserv. 

1. LEMNA 

Plants floating or submerged (if submerged, rising to the surface during flowering periods); roots 1 

per thallus or rarely absent. Thallus solitary or in small, connected groups of 2-10, symmetric or slightly 

asymmetric, round, elliptic, obovate or lanceolate, flat or slightly swollen beneath, containing raphides, 

the margins entire or rarely denticulate; stomata on upper surface; nerves usually 1-3; budding pouch 

opening at margin of thallus, rarely slightly dorsal or ventral. Seeds longitudinally ribbed, rarely smooth. 

A cosmopolitan genus of 13 species. 

TYPE: Lemna minor L., Sp. Pl. 970. 1753, typ. conserv. 

Key to the species of Lemna 

1. Thallus 3-nerved, broadly oblong, obovate or suborbicular, not much longer than broad 

................................................................................................................................1 . L. aequinoctialis 

1. Thallus 1-nerved or apparently nerveless, oblong (often narrowly so) usually at least twice as long as 

broad when mature ………............................................................................................................…. 2 

2. Thallus mostly 1-1.5 mm long, equilateral, always floating on water surface; epidermis often with 

minute, linear, whitish cystoliths ……….........................................................…. 2. L. minuscula 

2. Thallus mostly 2-4.5 mm long, inequilateral and often subfalcate, floating or submerged; 

epidermis lacking evident cystoliths ………………..................................……. 3. L. valdiviana 

1. Lemna aequinoctialis Welw., Ann. Cons. 

Ultramar. (Portugal), Parte Não Off. ser. 1, 

55: 578. 1859 [“1858”] Type: Angola; 

Luanda. Welwitsch 206 (lectotype: STU; 

isolectotypes: BM, G, K, ZT), designated by 

Landolt, Veröff. Geobot. Inst. ETH Stiftung 

Rübel Zürich 71: 445. 1986. 

Lemna perpusilla var. trinervis Austin ex A. Gray, 

Manual ed. 5, 479. 1867; Lemna trinervis (A. 

Gray) Small, Fl. s.e. U.S. 230. 1903. Type: 

United States; Pennsylvania. C. F. Austin s.n. 

(NY). 

Lemna minor sensu Griseb., 1864, non L., 1753. 

Lemna perpusilla sensu Britton & P. Wilson, 

1923, non Torr., 1843. 

Fig. 10. A, B 

Thallus solitary or 2-5 connected, mostly 1.5- 

3.3 × 0.5-2.5 mm; upper surface smooth except for 

a small papule near the tip and another at the node; 

nerves usually 3 but sometimes indistinct. Ovary 

with a single ovule; seed oblong to ovoid, 0.45-0.8 

mm long, brownish, with 8-26 longitudinal ribs. 

General distribution: A pantropical species 

extending both north and south into temperate 

areas. 


Islands: Floating in quiet or sheltered fresh water 

at low elevations (0-84 m). Recorded from 

Aguadilla, Cabo Rojo, Caguas, Cataño, Coamo, 

Dorado, Guánica, Humacao, Juncos, Lajas, 

Quebradillas, Salinas, San Juan, and Vega Baja; 

St. Croix, St. John, Tortola, and Virgin Gorda. 

Common names: Puerto Rico: Lentejilla de 

agua, Yerba de pato. 


Cataño: Bo. Palmas, Ciénaga de las Cucharillas, 

Proctor & Rivera 46204 (US). Dorado: Liogier & 

Liogier 29856 (UPR). Guánica: N.L. Britton & 

Shafer 1886 (US). San Juan: Bo. Puerto Nuevo, 

wetlands along Ave. Kennedy near crossing of Río 

Puerto Nuevo, Proctor 46207 (US); Río Piedras, 

Liogier 37226 (UPR). Vega Baja: sea level, 

Liogier 31988 (UPR). ST. CROIX: Crequis Dam 

Resevoir, Fosberg 53950 (US); South River, in 

upper estuary, Fosberg 59042 (US); Estate Mt. 

Pleasant (Colquehoun), Rd 72, Cruzan Gardens, 

Proctor et al. 44992 (US). ST. JOHN: Coral Bay 

Quarter, Road 107, Acevedo-Rdgz. et al. 4022 

(US); Reef Bay Quarter, Lameshur, Acevedo- 

Rdgz. 5260 (US).

54 


Fig. 10. A, B. Lemna aequinoctialis. A. Habit (colony), top and lateral views. B. Pistillate and staminate flower and utricle. From 

Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78. 

2. Lemna minuscula Herter, Revista Sudamer. 

Bot. 8: 185. 1954. Type: Chile; Santiago. 

Philippi 5 (lectotype: STU; isolectotypes: 

BM, G, GOET, K, LE, MEL, MO, S, SGO, 

UPS, ZT), designated by Landolt, Veröff. 

Geobot. Inst. ETH Stiftung Rübel Zürich 71: 

447. 1986. 

Lemna minuta Kunth in Humb., Bonpl., & Kunth, 

Nov. Gen. Sp. 1 [quarto ed.]: 372. 1816, nom. 

nud. 

Lemna perpusilla sensu Liogier & Martorell, 

1982, non Torr., 1843. 

Thallus solitary or sometimes 2-4 cohering, 

mostly 0.8-1.5 × 0.5-1.5 mm, up to 2 times longer 

than wide, rounded at both ends. Nerves indistinct 

and relatively short. Plants seldom flowering or 

fruiting; seeds ovoid, 0.4-0.55 mm long, with 12- 

15 longitudinal ribs. 

General distribution: Indigenous primarily in 

the warm-temperate parts of North and South 

America and at rather high elevations throughout 

the Andes, very rare in the West Indies; introduced 

in western Europe and Japan. 


Islands: Rare, sea level to 300 m, sometimes in 

artificial pools. Recorded for Barceloneta and 

Cayey; St. Thomas. 


Barceloneta: Bo. Garrochales, Caño Matos, 

2. SPIRODELA 

Spirodela Schleid., Linnaea 13: 391. 1839. 

Proctor 46146 (SJ). Cayey: Liogier et al. 33219 

(UPR). ST. THOMAS: Estate Neltjberg, vicinity of 

Ruy Point, overlooking Neltjberg Bay, Proctor et 

al. 45290 (SJ, US). 

3. Lemna valdiviana Phil., Linnaea 33: 239. 

1864. Type: Chile; Valdivia. Philippi 1, 

(lectotype: STU; isolectotype: MO), 

designated by Landolt, Veröff. Geobot. Inst. 

ETH Stiftung Rübel Zürich 71: 447. 1986. 

Thallus floating or frequently submerged, 

often clustering in groups of 6-20, mostly 2.5-3.5 × 

1.3-3 mm, 1.3-3 times longer than wide, often 

somewhat pointed at apex; nerve usually rather 

prominent and relatively long (extending at least 

76% of the distance between node and thallus tip). 

Occasionally flowering and fruiting; seeds ovoid, 

0.6-0.8 mm long, with 16-29 longitudinal ribs. 

General distribution: Confined to the 

Western Hemisphere, where it occurs in tropical 

and warm-temperate areas. 

Distribution in Puerto Rico: Very rare, 

confined to the north side of Puerto Rico at or near 

sea level. Recorded for Barceloneta, San Juan, and 

Vega Baja. 


Barceloneta: Bo. Garrochales, Caño Matos, 

Proctor 46147 (SJ). San Juan: Hioram s. n. (GH, 

cited by Landolt). 

Thallus lanceolate, ovate, or suborbicular, always floating on water surface, solitary or several


cohering in a rosette-like group; nerves mostly 5-16 per thallus; tissue containing anthocyanin pigments 

(visible in dead thallus as brown dots) and thin crystals in bundles (raphides). Roots multiple (2-21) per 

thallus. Ovary with 1-5 ovules. A cosmopolitan genus of 3 species. 

TYPE: Spirodela polyrhiza (L.) Schleid. (≡ Lemna polyrhiza L.) 

Key to the species of Spirodela 

1. Thallus 1-1.5 times longer than broad, the upper surface smooth; roots 7-21 per thallus 

…….............................................................................................................................. 1. S. polyrhiza 

1. Thallus 1.5-2 times longer than broad, the upper surface with a median line of small papules; roots 

mostly 2-7 per thallus …………...........................................................................…….. 2. S. punctata 

1. Spirodela polyrhiza (L.) Schleid., Linnaea 13: 

392. 1839; Lemna polyrhiza L., Sp. Pl. 970. 

1753; Lenticula polyrhiza (L.) Lam., Fl. 

Franç. 2: 189. 1779 [“1778”]; Thelmatophace 

polyrhiza (L.) Godr., Fl. Lorr. 3. 1843. 

Neotype: Canada; Vancouver Is. Macoun 

88277 (US!; isoneotypes: C, GH, NY), here 

designated. 

Thallus solitary or in rosette-like clusters, 

rounded or sometimes pointed at tip, 1.5-10 × 1.5- 

5 mm, sometimes with a red spot above the node. 

Flowers and fruits rarely produced. Ovary with 1- 

2 ovules. Fruit 1-1.5 mm long and wide, with 

winged edges near the top; seeds 1 or rarely 2, 0.7- 

1 mm long, with 12-20 distinct longitudinal ribs. 

General distribution: A widely distributed 

species common in North America and Europe, 

frequent to common in eastern and southern Asia, 

northern and eastern Australia, and eastern Africa, 

but virtually absent from South America. 


Islands: Uncommon to rare, mostly at or near sea 

level, recorded from Aguada, Coamo, Guayanilla, 

Manatí, and Vega Baja; St. Croix. 

Note: Landolt (1986) was unable to locate the 

type of Spirodela polyrhiza, although he stated 

that the identity of the species is unmistakable. 

Because he failed to designate a type, we propose 

as neotype a specimen that was cited in his 

monograph and is represented in several herbaria. 


Coamo: Coamo River, N.L. Britton et al. 5805 

(US); Coamo, Sintenis 3189 (US). Guayanilla: 

N.L. Britton & Shafer 1795 (US). Manatí: Río 

Arriba Saliente, Sintenis 6656 (US). Vega Baja: 

Bo. Cabo Caribe, 1 km due SE of Escuela Cabo 

Caribe, Proctor et al. 45624 (US). ST. CROIX: 

Estate Mt. Pleasant (Colquehoun), Rd 72, Cruzan 

Gardens, Proctor et al. 44993 (US). Cruzan 

Gardens, Colquehorn, Fosberg 58961 (US). 

2. Spirodela punctata (G. Mey.) C. H. Thomps., 

Rep. (Annual) Missouri Bot. Gard. 9: 28. 

1898; Lemna punctata G. Mey., Prim. Fl. 

Esseq. 262. 1818; Landoltia punctata (G. 

Mey.) D.H. Les & D.J. Crawford, Novon 9: 

532. 1999. Neotype: Chile; Tierra de Fuego 

Is. Wilkes Expedition s.n., 1838 (US!; 

isoneotypes: BM, GH, KANU, MO), 

designated by Les & Crawford, Novon 9: 532. 

1999. 

Thallus solitary or in small clusters, ovate to 

lanceolate, often somewhat pointed, 1.5-8 × 1-5 

mm, 1.5-2 times longer than wide. Plants 

occasionally flowering and fruiting. Ovary with 1- 

2 ovules. Fruit 0.8-1 mm long, winged; seeds 

usually 1, ca. 0.8 mm long, with 10-15 longitudinal 

ribs. 

General distribution: Widely but sporadically 

distributed, most common in California, S. E. 

United States, S. E. Africa, S. E. Asia, Japan, and 

E. and S. E. Australia. It appears to be absent from 

Europe, northern Asia, Central America and 

nearly so from South America, except for the 

neotype, a few records from southeastern Brazil, 

and one record each from Colombia and Guyana. It 

has not previously been reported from the West 

Indies. 

Distribution in Puerto Rico: Occurs mostly 

at or near sea level. Recorded from Arecibo, 

Florida, Río Grande, Vega Alta, and Vega Baja. 

Note: Meyer’s original type from Essequibo 

River is believed to be lost. Les & Crawford (1999) 

proposed the new genus Landoltia based on

56 


Spirodela punctata as allozyme and cpDNA 

(rbcL) sequence data in this species differ from 

other species of Spirodela and Lemna. The 

recognition of this genus based on morphological 

characters however, is rather difficult as it is 

intermediate between other species of Spirodela 

and Lemna. 


3. WOLFFIA 

Wolffia Horkel ex Schleid., Beitr. Bot. 1: 233. 1844, nom. conserv. 

Arecibo: Bo. Santana, S side of Ciénaga 

Tiburones, c. 1.5 km N of Aeropuerto de Arecibo, 

Proctor & Rivera 46273 (US). Río Grande: Bo. 

Herreras, just N of Caño San Luis, Proctor & 

Morris 45993 (US). Vega Baja: Bo. Cibuco, 

mangrove thicket 1.2 km due ENE of Cerro 

Cibuco, Proctor et al. 45571 (US). 

Minute floating plants often occurring in vast numbers. Thallus globular, ovoid, cylindrical or boatshaped, 

mostly free or 2 cohering. Veins and roots lacking, the tissue without air spaces. Flowering thallus 

similar to vegetative ones. Inflorescence one per plant, originating in a cavity more or less median on the 

upper surface of the thallus; pistillate flower consisting of a pistil; staminate flower with a single stamen. 

A cosmopolitan genus of 8 species. 

TYPE: Wolffia michelii Schleid., typ. conserv. 

1. Wolffia brasiliensis Wedd., Ann. Sci. Nat. 

Bot., ser. 3, 12: 170. 1849; Grantia 

brasiliensis (Wedd.) Mac Mill., Metasp. 

Minnesota Valley 134. 1892. Type: Brazil; 

Mato Grosso. Weddell s. n. (lectotype: STU; 

isolectotypes: K, L, MO) designated by 

Landolt, Veröff. Geobot. Inst. ETH Stiftung 

Rübel Zürich 71: 452. 1986. 

Wolffia punctata Griseb., Fl. Brit. W. I. 512. 1864. 

Type: Jamaica. Wullschlaegel, s. n. (holotype: 

GOET?, isotype: STU). 

Plants densely gregarious on still, fresh water. 

Thallus broadly ovate to suborbicular, 0.5-1.6 × 

0.7-1.5 mm, rounded at the tip, with a prominent 

papule in the middle of the upper surface 

4. WOLFFIELLA 

Wolffiella (Hegelm.) Hegelm., Bot. Jahrb. Syst. 21: 303. 1895. 

(sometimes lacking on smaller thallus). Plants 

occasionally flowering but rarely fruiting. Seeds 

ca. 0.4 mm long. 

General distribution: Commonly in the 

southeastern United States, southern Mexico, 

Guatemala, El Salvador, the West Indies, northern 

and S. E. South America. First report for Puerto 

Rico. 

Distribution in Puerto Rico: Known from a 

single collection in Cabo Rojo. 


Cabo Rojo: Bo. Llanos Costa, small pond 1 km. 

due ESE of intersection Roads 301 & 303, elev. 35 

m, coll. 19 Sept. 1987, Proctor & McKenzie 43935 

(SJ). 

Thallus floating suspended under the surface in still fresh or slightly brackish water, some species said 

to rise to the surface when flowering or fruiting; tissue thin and translucent, the thallus solitary or several, 

cohering together, sometimes forming a loose ball; pigment cells occurring in some species at the nodal 

end of the thallus, there developing a flat triangular pouch out of which daughter thallus emerge. Flowers 

1 or 2, produced from a cavity at the side of the median line of the upper thallus surface. Fruit a slightly 

compressed utricle with a persistent style; seeds smooth, with spongy outer coat. A widely distributed 

genus of 8 species. 

LECTOTYPE: Wolffiella oblonga (Phil.) Hegelm. (≡ Lemna oblonga Phil.), designated by Britton & A. 

Brown, Ill. Fl. N. U. S. ed. 2., 1: 449. 1913.


1. Wolffiella welwitschii (Hegelm.) Monod, 

Mém. Soc. Hist. Nat. Afrique N., Hors Sér 2: 

229. 1949; Wolffia welwitschii Hegelm., J. 

Bot. 3: 114. 1865; Wolffiopsis welwitschii 

(Hegelm.) Hartog & Plas, Blumea 18: 366. 

1970. Type: Angola; Quizambo. Welwitsch 

209 (holotype: STU; isotypes: BM, C, G, H, 

K, L, as cited by Landolt, 1986). 

Wolffiella lingulata sensu Liogier & Martorell, 

1982, non Hegelmaier, 1895. 

Thallus submerged, with the basal part often 

near the water surface, the tip bent downward, 2 or 

3 cohering together to form a saddle-shaped 

configuration; individual thallus broadly tongueshaped, 

rounded at the tip, 4-7 × 2.5-5 mm, often 

1.2-2 times longer than wide; a few stomata (to 12) 

occurring especially along the lateral margins of 

the base. Plants often flowering and fruiting. 

Flowers 2 per thallus. Seeds ca. 0.45 mm long, 0.3 

mm thick. 

General distribution: Widely distributed in 

tropical America and Africa; it occurs on all of the 

Greater Antilles and also on Guadeloupe in the 



Islands: Uncommon (or unobserved) in still, 

sheltered, fresh or slightly brackish water at or 

near sea level. Recorded from Loíza, Manatí, and 

Vega Baja; collected long ago in St. Croix. 


Vega Baja: Bo. Cibuco, mangrove thicket 1.2 km 

due ENE of Cerro Cibuco, Proctor et al. 45572 

(US). 

Family 3. LIMNOCHARITACEAE Water-Poppy Family 

Limnocharitaceae Takht. ex Cronquist, Integr. Syst. Class. Fl. Pl.: 1048. 1981. 


Rhizomatous or stoloniferous perennial aquatic herbs producing milky sap, of fresh water habitats; 

roots fibrous, not septate. Stems erect, unbranched, fleshy. Leaves submersed and floating, primarily 

basal, sessile or petiolate with sheathing base, glabrous; blades orbicular to lanceolate with entire margins, 

obtuse or subacute at apex with an apical pore, cordate to attenuate at base, venation reticulate, primary 

veins parallel from base of blade to apex, secondary veins reticulate. Inflorescence a terminal, involucrate 

umbel, erect to floating, bracteate, with vegetative buds in some species; bracts few to several, ovate, 

membranous, acuminate. Flowers bisexual, actinomorphic, hypogynous, pedicellate; sepals 3, persistent 

and enclosing flower and fruit; petals 3, often delicate, deciduous; stamens (6-) 20-25, distinct, basifixed; 

anthers 4-locular, dehiscing longitudinally; pistils (3-) 10-12, coherent proximally, 1-locular with 

numerous ovules, the placentation laminar; style short or absent, the stigma linear. Fruit a follicle, 

dehiscing adaxially; seeds numerous, U-shaped, glandular-pubescent or costate. A family of 3 genera and 

8 species with pantropical distribution. 

Type: Limnocharis Humb. & Bonpl. 

References: Haynes, R. R. & L. B. Holm-Nielsen. 1992. The Limnocharitaceae. Fl. Neotrop. Monogr. 

56: 1-34. Haynes, R. R. Fam. 220. Limnocharitaceae. Pp. 5-6. In: Flora of North America Editorial 

Committee (eds.), Fl. North Amer., vol. 22. 1993. 

1. HYDROCLEYS 

Hydrocleys Rich., Mém. Mus. Nat. Hist. 1: 368. 1815. 

Submersed, stoloniferous herbs with submersed or floating leaves. Stems short, erect. Leaves basal, 

the submersed ones phyllodia, sessile, the floating leaves long-petiolate; petioles terete, septate, sheathing 

at base; blades orbiculate to oblong-lanceolate, rounded to cordate at base, mucronate to obtuse at apex. 

Inflorescences few- to many-flowered, occasionally proliferating with leaves and stolons; scape elongate, 

septate; bracts subtending pedicel, distinct, elliptic to lanceolate, shorter than pedicel. Flowers on elongate

58 


pedicels; sepals erect, lanceolate, coriaceous, green, midvein present or absent, hood like at apex, 

persistent; petals erect to spreading, orbicular to oblong-obovate, fugacious, yellow to white; stamens 6 

to many, in 1 to several series, the outer often sterile; carpels 3-8, terete, linear-lanceoloid, attenuate into 

style; style curved inward, papillose at apex. Follicles ± terete, linear-lanceoloid, membranous, dehiscing 

along adaxial margins; seeds numerous, sparsely to densely glandular-pubescent or glabrous. Five species 

with the center of distribution in South America; one species introduced in North America. 

TYPE: Hydrocleys commersonii Rich. 

1. Hydrocleys nymphoides (Willd.) Buchenau, 

Index Crit. Butom. Alism. Juncag. 2, 7. 1868; 

Stratiotes nymphoides Willd., Sp. Pl. 4: 821. 

1805. Type: Venezuela. Humboldt s.n. 

(holotype: B-Willd.). 

Plants 10-45 (-60) cm tall; stolons to 45 cm 

long. Leaves with petioles 1.5-40 (-70) cm × 

(0.9-) 1.7-10 mm, the sheathing base to 9 cm long; 

blades with 5-9 veins, broadly ovate to orbicular, 

(1.4-) 2-10 (-12) × (0.9-) 2-9 (-11) cm, obtuse to 

slightly mucronate at apex, cordate at base. 

Inflorescence 1- to 6-flowered, proliferating with 

stolons and leaves; peduncles 4-30 cm × 1.5-6 

mm; bracts elliptic, 2-4.5 × 0.4-1 cm, obtuse at 

apex; pedicels spreading, 3.5-17.5 cm × 1.5-6 mm. 

Flowers 5.5-6.5 cm broad; sepals ovate-elliptic, 

Family 4. ALISMATACEAE Water-plantain Family 

Alismataceae Vent., Tabl. Règne Vég. 2: 157. 1799, nom. conserv. 


Perennial (sometimes annual) herbaceous, acaulescent plants with milky sap, aquatic or growing in 

wet or marshy places. Leaves petiolate, sheating at base, the blades flat and several-veined. Flowers 

bisexual or unisexual, symmetrical, whorled on terminal, erect racemes or panicles. Calyx of 3 persistent 

green sepals. Petals 3, white, delicate and soon falling. Stamens 6 or more, the filaments distinct, the 

anthers 2-locular. Carpels free, few or many, borne in heads in our species, each normally containing 1 

ovule; style usually persistent, appearing as a beak on the fruit. Fruit a head of achenes. Seeds curved, the 

embryo horseshoe-shaped. A semi cosmopolitan family of 11 genera and ca. 75 species (Haynes & Holm- 

Nielsen, 1994), of which two occur in the Neotropics. 

TYPE: Alisma L. 

Reference: Haynes, R. R. & Holm-Nielsen, L. B. 1994. The Alismataceae. Fl. Neotrop. Monogr. 64: 

1-112. 


1.3-2.8 × 0.7-1.3 cm, narrowly scarious on 

margins; petals spreading, 2.3-3.5 × 3.5-4.4 cm, 

pale yellow to white with yellow base; stamens 20- 

25, the anthers 4-6 mm long; staminodes 

numerous; pistils 5-8, ca. 10 mm long. Follicle 10- 

18 × 2-3.5 mm, the beak 2.5-5.5 mm long. Seeds 

oblong-obovate, 1-1.3 × 0.7-0.8 mm, glabrous or 

glandular-pubescent to sparsely so, reticulate on 

surface, reddish. 

General distribution: United States (Florida 

and Texas), Guatemala, Puerto Rico, Curaçao, 

Trinidad, and South America. 

Distribution in Puerto Rico: Stagnant waters. 

Recorded from Añasco and Vega Alta. 


Añasco: Sintenis 5779 (U); Villa Espinosa, N.L. 

Britton & E.G. Britton 9941 (NY). 

1. Flowers all bisexual; fruits terete, usually ribbed, with glands between the ribs ……….. 1. Echinodorus 

1. Flowers, at least the basal ones, unisexual; fruits not terete, often with a curved wing …… 2. Sagittaria


1. ECHINODORUS 

Echinodorus Rich. ex Engelm. in A. Gray, Manual 460. 1848. 

Annual or perennial herbs, glabrous to stellate-pubescent, growing emersed in shallow, fresh or 

slightly brackish water. Stems short-rhizomatous. Leaves petiolate; petioles trigonous, sheathing at base; 

blades linear to broadly ovate, often with pellucid dots or lines, the margins entire or undulating, the apex 

acute to acuminate, the base attenuate to cordate. Inflorescence erect, emersed, racemose or paniculate, 

rarely umbelliform; bracts free or partly connate, glabrous or papillose along the ribs, obtuse to acute. 

Flowers bisexual, subsessile to pedicellate; pedicels often elongating after anthesis; sepals herbaceous to 

coriaceous, reflexed or spreading; petals white, larger than the sepals; stamens 9-many, the filaments 

glabrous, the anthers versatile or basifixed; gynoecium of many free carpels, distributed evenly over the 

receptacle, each with a single ovule, the styles terminal or lateral, persistent on the fruits. Fruit a terete 

achene, often longitudinally ribbed, glandular, and beaked. A genus of 26 species distributed from the 

northern United States to Chile. 

LECTOTYPE: Echinodorus rostratus (Nutt.) Engelm. (≡ Alisma rostrata Nutt.; = Echinodorus berteroi 

(Spreng.) Fassett), designated by Small, N. Amer. Fl. 17(1): 46. 1909. 

Key to the species of Echinodorus 

1. Leaf blades elliptic or ovate, usually truncate or subcordate at base, with 3-11 longitudinal veins; sheath 

at base of petiole to 7 cm long; inflorescence with up to 9 whorls of flowers; achenes with erect, 

terminal, beak 0.6-1.3 mm long …..............................................................…………… 1. E. berteroi 

1. Leaf blades linear-lanceolate, attenuate at base, with 1-3 longitudinal veins; sheath at base of petiole not 

over 2 cm long; inflorescence umbelliform or with two whorls of flowers; achene with a lateral beak, 

0.1-0.3 mm long ……............................................................................................….. 2. E. bolivianus 

1. Echinodorus berteroi (Spreng.) Fassett, 

Rhodora 57: 139. 1955; Alisma berteroi 

Spreng., Syst. Veg. 2: 163. 1825. [as 

“berterii”]. Type: Lesser Antilles; 

Guadeloupe. Bertero s. n. (holotype: GOET; 

isotype: TO). 

Alisma rostratum Nutt., Trans. Amer. Philos. 

Soc. ser. 2. 5: 159. 1837. Type: Arkansas (or 

Oklahoma ?). Nuttall s. n. (holotype: PH; 

isotype: BM). 

Echinodorus cordifolius sensu Bello, 1883, and 

sensu Britton & P. Wilson, 1923, non (L.) 

Griseb., 1857. 

Fig. 11. A-D 

Annual or short-lived glabrous herb of wet 

ditches and marshy sites; rhizomes erect, to 2.5 

cm long; plants to 70 cm tall. Leaves variable in 

size and shape; petioles to 20 cm long; submerged 

leaves linear or ribbon-like; emersed leaves 

ovate, with pellucid lines. Inflorescence erect and 

overtopping the leaves, paniculate or sometimes 

a raceme of flowers in whorls; pedicels 0.6-2.8 

cm long. Flowers 6-11 mm wide; petals clawed, 

spreading, 2.5-4.8 mm long; stamens ca. 15, with 

filaments to 7 mm long, the anthers versatile, ca. 8 

mm long. Carpels numerous on an ovoid receptacle 

that enlarges in fruit. Achene terete, 2-keeled, 

glandular, to 3.2 mm long with slender erect beak 

0.6-1.3 mm long. 

General distribution: Northern United States 

to Mexico, Central America, the West Indies, 

northern South America and Peru. 


Islands: At or near sea level in fresh-water ditches 

and marshes, and occasionally in habitats that are 

slightly brackish. Recorded from Añasco, Cabo 

Rojo, Cataño, Coamo, Guánica, Guayama, 

Guayanilla, Juana Díaz, Lajas, and Vega Baja; St. 

Croix and St. Thomas. 

Common name: Puerto Rico: Llantén de agua. 


Juana Díaz, Proctor 43361 (US). 

2. Echinodorus bolivianus (Rusby) Holm-Niels., 

Brittonia 31: 276. 1979; Alisma bolivianum 

Rusby, [as “boliviana”] Mem. New York Bot. 

Gard. 7: 208. 1927. Type: Bolivia; Reyes.

60 


White 1540 (lectotype: NY; isolectotypes: 

GH, K, NY, US!), designated by Haynes & 

Holm-Nielsen, Fl. Neotrop. Monogr. 64: 14. 

1994. 

Alisma tenellum f. latifolium Seub. in Martius, Fl. 

Bras. 3(1): 105. 1848; Echinodorus tenellus 

var. latifolius (Seub.) Fassett, Rhodora 57: 

202. 1955. Echinodorus latifolius (Seub.) 

Rataj, Stud. Ceskoslov. Akad. Ved. 2: 18. 

1975. Lectotype: Brazil; Minas Gerais. 

Claussen 50 (BR), designated by Haynes & 

Holm-Nielsen, Brittonia 38: 326. 1986. 

Helianthium tenellum sensu Britton & P. Wilson, 

Bot. Porto Rico 5: 14. 1923, non Alisma 

tenellum Mart., 1830. 

Small annual, glabrous herb of fresh-water 

marshes and pools. Rhizomes short, stoloniferous. 

Leaves erect, often numerous in a loose rosette; 

petioles varying greatly according to habitat, from 

ca. 1.5 cm long on plants growing on wet soil to 35 

cm or more on plants submerged under water, in 

the latter case both leaf blades and inflorescence 

emersed above the water surface; leaf blades 

linear-lanceolate to lanceolate, 1-6(7.4) × 0.2-1.8 

Sagittaria L., Sp. Pl. 993. 1753. 

2. SAGITTARIA 

cm, pellucid lineolate, the apex acute to 

acuminate, the base long-tapering, veins 1 or 

usually 3, two of them located parallel to and close 

to the margins, the margins entire. Inflorescence 

erect and overtopping the leaves, umbelliform or 

sometimes a raceme with two whorls of 5-6 

flowers; pedicels 1.1-6.2 cm long, bracteate at 

base, spreading. Sepals spreading, 2.8-3.6 × 2.8- 

3.6 mm, 7-veined; petals clawed, 2.5-4.1 mm long; 

stamens 9, the anthers basifixed, 0.5-0.6 mm long; 

carpels 15-20. Achene obovoid, 0.6-1.8 mm long, 

3-or 4-ribbed, not keeled, glandless, the beak 

lateral, 0.1-0.3 mm long. 

General distribution: Southernmost Mexico, 

Central America, Greater Antilles (except Cuba), 

and South America. 

Distribution in Puerto Rico: Although known 

only from near sea level in Puerto Rico, this 

species occurs at various elevations up to 2000 

meters elsewhere in its range. Collected in San 

Juan, Caja de Muerto Island, Salinas, and Vega 

Baja. 


Vega Baja: Laguna Yeguada, N.L. Britton et al. 

6773 (US). 

Monoecious or rarely dioecious, perennial or rarely, glabrous to sparsely pubescent herbs of fresh or 

slightly brackish waters, the plants submerged, floating, or emerged. Stems often forming rhizomes, these 

sometimes terminated by tubers. Leaves sessile or petiolate; petioles terete or triangular; leaf blades 

present or sometimes absent, with or without basal lobes, lacking pellucid lines. Inflorescence racemose, 

paniculate or less often umbelliform with flowers usually arranged in paired whorls of 3; staminate 

flowers distal, pistillate flowers basal on inflorescence; sepals herbaceous to coriaceous, reflexed in 

staminate flowers, appressed or reflexed in pistillate flowers; petals usually white, rarely tinged with pink; 

stamens whorled, 7 to numerous, the anthers basifixed, linear to orbicular, the filaments linear or subulate, 

glabrous or pubescent. Carpels numerous, spirally arranged, each with 1 ovule, the styles terminal. 

Achene flattened, eglandular, winged, beaked, membranaceous. A primarily Western Hemisphere genus 

of about 25 species, 3-4 of which occur in Europe and Asia. 

LECTOTYPE: Sagittaria sagittifolia L., designated by Small, N. Amer. Fl. 17: 51. 1909. 

Reference: Bogin, C. 1955. A Revision of the genus Sagittaria (Alismataceae). Mem. New York Bot. 

Gard. 9: 179-233. 

Key to the species of Sagittaria 

1. Leaf blades linear, elliptic or ovate, attenuate at base, lacking basal lobes …….......….. 2. S. lancifolia 

1. Leaf blades sagittate or hastate, with 2 large divergent lobes at base ……................................…….. 2


2. Sepals appressed around fruiting aggregate; rhizome short, not stoloniferous, lacking tuberous 

corms; achenes 1.5-2.2 mm long …………....................................................…. 1. S. intermedia 

2. Sepals reflexed below fruiting aggregate; rhizomes producing slender stolons to 30 cm long, often 

bearing tuberous corms; achenes 2.5-3.5 mm long …….................................……. 3. S. latifolia 

1. Sagittaria intermedia Micheli in Alph. de 

Candolle & C. de Candolle, Monogr. Phan. 3: 

80. 1881. Type: Cuba. Wright 3199 (lectotype: 

K; isolectotypes: G, GH, GOET, MO, NY, S, 

US!, W), designated by Bogin, Mem. New 

York Bot. Gard. 9: 201. 1955. 

Sagittaria acutifolia sensu Griseb., Cat. Pl. Cub. 

218. 1866, and sensu Bello, 1883, non L. f., 

1782 [“1781”]. 

Fig. 11. E-H 

Perennial, glabrous herb with short rhizomes. 

Leaves emersed; petioles 17-45 cm long, 

trigonous, with basal sheath to 10.5 cm long; 

blades hastate to sagittate, 5-15 (-21) × 2-9 cm, the 

apex acute, the base hastate with slightly spreading 

acute to subacuminate lobes. Inflorescences 

simple or branching from lowest whorls, with 3-8 

whorls of flowers; peduncles trigonous, mostly 

15-25 cm long. Sepals ovate, 4-8 mm long; petals 

white, ca. 2.5 cm long in staminate flowers, ca. 1 

cm long in pistillate flowers; stamens numerous, 

the filaments ca. 0.7 mm long, glabrous, the 

anthers terete, ca. 0.7 mm long. Achene obovoid, 

beaked, non-keeled, tuberculate, 1.5-2.2 mm long, 

grouped into a spherical aggregate, 0.6-1.5 cm in 

diam. 

General distribution: Primarily on all four of 

the Greater Antilles, with a few isolated records 

from southernmost Mexico and northernmost 

Colombia. 

Distribution in Puerto Rico: Uncommon in 

wet, marshy sites at or near sea level. Recorded 

from Cabo Rojo, Dorado, Hatillo, Loíza, 

Naguabo, and Río Grande. 


Cabo Rojo: Sintenis 679 (US). Dorado: N.L. 

Britton et al. 6690 (US). 

2. Sagittaria lancifolia L., Syst. Nat. ed. 10, 1270. 

1759. Lectotype: Jamaica. Browne s.n. 

(LINN-1124.61), designated by Haynes & 

Holm-Nielsen, Fl. Neotrop. Monogr. 64: 89. 

1994. 

. 

Perennial, glabrous to sparsely pubescent 

herb; rhizomes to 19 cm long and 4 cm thick. 

Leaves emersed, erect; petioles terete to trigonous, 

to 58 cm long, with basal sheaths to 26 cm long; 

blades linear, elliptic or ovate, 20-35 × 0.7-16 cm, 

chartaceous, pale green, with 7-9 longitudinal 

veins, the apex acute, the base obtuse. Inflorescence 

a raceme or panicle with up to 13 whorls, usually 

containing 3 flowers; peduncle 75-125 cm long; 

bracts striate, lanceolate to ovate-lanceolate. 

Staminate flowers with reflexed sepals; petals 

clawed, 0.8-1.5 cm long; stamens numerous, the 

filaments pubescent, 2.2-4 mm long, the anthers 

linear, 1.7-2.3 mm long. Pistillate flowers with 

reflexed or spreading sepals; petals clawed, 0.6- 

1.5 cm long. Achene obovate, beaked, keeled, 1.6- 

2.5 mm long, smooth, grouped into aggregates, 

0.5-1.2 cm in diam. 

General distribution: Coastal southeastern 

North America, Mexico, Central America, the 

Bahamas, Greater Antilles, Cayman Islands, and 

northern coastal South America from Colombia to 

Brazil. 

Distribution in Puerto Rico: Frequent or 

common in wet marshy sites from sea level to 

middle elevations (ca. 500 m). Recorded from 

Aguada, Aibonito, Arecibo, Bayamón, Cabo Rojo, 

Cataño, Humacao, Manatí, Mayagüez, San Juan, 

Toa Alta, Toa Baja, Utuado, and Vega Baja. 

Common names: Puerto Rico: Flecha de 

agua, Saeta de agua. 


Aibonito: Sintenis 2697 (US). Arecibo: Bosque de 

Río Abajo, Los Puercos, Acevedo-Rdgz. et al. 

2268 (NY, US, VINPS). Bayamón: Johnston 912 

(US); Heller & Heller 416 (US); Stevenson 114 

(US); Johnston & Stevenson 1149 (US). Cabo 

Rojo: Sintenis 680 (US). Humacao: Playa de 

Humacao, Eggers s.n. (US). Manatí: Laguna 

Tortuguero, N.L. Britton et al. 3844 (US). 

Mayagüez: Sargent B31 (US); N.L. Britton 2370 

(US). Toa Alta: Puerto Flaco, Goll et al. 917 (US). 

Toa Baja: Candelaria, Goll et al. 281 (US). Vega 

Baja: N.L. Britton et al. 6757 (US).

62 


Fig. 11. A-D. Echinodorus berteroi. A. Habit. B. Flower at anthesis. C. Flower at fruiting stage. D. Achene. E-H. Sagittaria 

intermedia. E. Habit. F. Flower at anthesis. G. Flowers at fruiting stage, side and top views. H. Achene. (A-B, from Proctor 

43934; C-D, from Proctor 46370; E, from Proctor 46984; F-H, from Proctor 45331).


3. Sagittaria latifolia Willd., Sp. Pl. 4: 409. 1805. 

Type: “Canada at Carolinam”. Muhlenberg s. 

n. (holotype: B-Willd., fragm. US!). 

Perennial glabrous or pubescent herb 

proliferating by stolons and/or corms, the stolons 

to 30 cm long and ca. 0.3 cm thick, the corms ca. 2 

cm in diam. Leaves emersed, erect; petioles 

trigonous, 6.5-51 cm long with basal sheath 1.5- 

17.5 cm long; leaf blades pale green, sagittate or 

rarely hastate, 15-30 × 2-17 cm, with 7-13 

longitudinal veins, the apex acute, or rarely obtuse, 

the base sagittate, with acuminate lobes 0.5-16.5 

cm long. Inflorescences racemose or sparingly 

paniculate, with up to 9 whorls of flowers; 

peduncles trigonous, 10-59 cm long. Staminate 

flowers with reflexed or spreading sepals; petals 

clawed, 6-9 mm long; stamens 16-18, the 

filaments glabrous 1-3 mm long, the anthers linear, 

1.2-1.9 mm long. Pistillate flowers with reflexed 

sepals; petals clawed, ca. 18 mm long. Achene 

oblanceolate, beaked, not keeled, 2.5-3.5 mm 

long, non-tuberculate, sometimes with 1 or 2 

glands, grouped into an aggregate 1-1.7 cm in 

diam. 

General distribution: North America to 

northern South America, and Puerto Rico. 

Distribution in Puerto Rico: Recorded by a 

single specimen collected in marshy lands near 

San José Lagoon, San Juan by Hioram in 1909. 

Perhaps a chance introduction, and very likely 

now extirpated from Puerto Rico. 


San Juan: San José Lagoon, Hioram s.n. (GH). 

Family 5. HYDROCHARITACEAE Frog’s bit Family 

Hydrocharitaceae Juss., Gen. Pl. 67. 1789, nom. conserv. 


Annual or perennial aquatic herbs, monoecious or dioecious, having either a creeping monopodial 

rhizome with roots at the nodes and distichous leaves, or an erect axis with roots at the base and leaves 

opposite, spirally arranged, or apparently whorled. Leaves eligulate, submerged or floating, linear, 

lanceolate, elliptic, ovate, or orbicular, with margins entire, serrulate or toothed, and either sessile and 

sheathed at base or else petiolate with leaf blade more or less expanded. Flowers symmetrical, unisexual 

or bisexual, sessile or pedicellate, solitary or in cymose inflorescences in a spathe of two, free or connate 

leaves; perianth absent or of one or two whorls, each of 3 free segments; stamens 1 or 3-several, the 

filaments slender or absent; ovary inferior, 1- to 15-carpellate, the ovules solitary or several; style 1-15. 

Fruit indehiscent, with membranous or fleshy pericarp; seeds fusiform, ellipsoid, or globose. A 

cosmopolitan family of marine and fresh-water plants with 17 genera and ca. 130 species. 

TYPE: Hydrocharis L. 

References: den Hartog, C., 1970. The sea grasses of the world, Verh. Kon. Ned. Akad. Wetensch., 

Afd. Natuurk., Tweede Sect. 59(1): 212-275. Guard, B. J., 1995. Wetland plants of Oregon & Washington. 

Long Pine Publishing, Canada. Les, D. H. & M. A. Cleland, 1997. Phylogenetic studies in Alismatidae II: 

Evolution of marine Angiosperms (sea grasses) and hydrophily. Syst. Bot. 22(3): 443-463. Phillips, R. C. 

& E. G. Meñez. 1988. Seagrasses. Smithsonian Contr. Marine Sci. 34: 1-104. 


1. Submerged marine plants (only in salt water). …………..................................................................... 2 

2. Leaves less than 4 cm long, petiolate, with oblong to elliptic blades ……........……. 2. Halophila 

2. Leaves 6-60 cm long, lacking petioles, with strap-shaped blades ……............…….. 5. Thalassia 

1. Submerged or floating plants of fresh or brackish water. …………….........................................…… 3 

3. Plants floating (rarely rooted on mud in very shallow water); leaves petiolate, entire, orbicular 

…..................................................................................................................…….. 3. Limnobium 

3. Plants submerged; leaves oblong or narrowly linear ………...................................................…. 4

64 


4. Stems few-branched; leaves oblong; flowers long-pedicellate, with conspicuous perianth 

………............................................................................................................……. 1. Elodea 

4. Stems much-branched; leaves narrowly linear; flowers sessile, without perianth …. 4. Najas 

Elodea Michx., Fl. Bor.-Amer. 1: 20. 1803. 

1. ELODEA 

Perennial submerged, aquatic herbs of fresh water; stems slender, elongate, few-branched, rooted or 

often breaking free. Lowest leaves alternate, elsewhere opposite or whorled, sessile, linear or narrowly 

oblong, translucent, bright green, 1-nerved, the margins finely serrulate. Flowers borne from small 

axillary spathes, usually unisexual, produced on filiform stalks that raise them just above the water 

surface; sepals 3, petals 3; stamens 3-9. Fruit a several-seeded capsule. A New World genus of about 14 

species, mostly occurring in temperate regions. Two species are widely cultivated in aquaria and have 

escaped and often become naturalized far from their original habitats. 

TYPE: Elodea canadensis Michx. 

Reference: St. John, H. 1967. The pistillate flower of Egeria densa Planch. Darwiniana 14(2-3): 571- 

573. 

Key to the species of Elodea 

1. All spathes with a solitary flower; leaves 0.6-1.2 cm long; petals narrower than the sepals 

….................................................................................................................................. 1. E. canadensis 

1. Staminate spathes with 2-several flowers; leaves up to 2 cm long; petals wider than the sepals 

….....................................................................................................................................… 2. E. densa 

1. Elodea canadensis Michx., Fl. Bor.-Amer. 1: 

20. 1803; Anacharis canadensis (Michx.) 

Planch., Mag. Nat. Hist., ser. 2. 1: 86. 1848. 

Type: Canada. Michaux s. n. (holotype: P). 

Stems dichotomously few-branched; lower 

leaves alternate, opposite, or in whorls of 3; upper 

leaves in whorls of 3-7, linear-oblong, translucent 

green and 1-nerved, the margins denticulate, the 

apex blunt to acutish. Staminate flowers sessile, in 

a globose spathe, emitting an elongate filamentous 

floral tube with hypanthium at water surface; 

stamens 3-9. Pistillate flowers solitary, in a tubular 

spathe, raised to the water surface by a slender 

hypanthial tube, bearing 3 green sepals and 3 white 

petals. Fruit a several-seeded capsule. 

General distribution: Eastern North America 

from Quebec to Virginia and west to California; 

widely introduced elsewhere and naturalized in 

Europe. 

Distribution in Puerto Rico: Cited by Liogier 

& Martorell (1982) as occurring “in a lake near 

Florida”; not seen by the author. Often used in 

aquaria because it increases water oxygen levels 

and thrives in artificial habitats (Guard, 1995). 

Common name: Puerto Rico: Tomillo de 

agua. 

2. Elodea densa (Planch.) Casp., Monatsber. 

Königl. Preuss. Akad. Wiss. Berlin 1857: 49. 

1857; Egeria densa Planch., Ann. Sci. Nat. 

Bot., ser. 3, 11: 80. 1849. Anacharis densa 

(Planch.) Vict., Contr. Lab. Bot. Univ. 

Montreal 18: 401. 1931. Type: Argentina; 

Buenos Aires. Tweedie 10 (holotype: K). 

Stems elongate, sparingly branched, 2-3 mm 

thick, rooted in the substrate or freely floating. 

Leaves linear-oblong, often 2-3 cm long, sessile, 

the lowermost ones opposite, the rest in whorls of 

3-6, translucent, bright green, mid rib sometimes 

with a few teeth beneath, the margins finely 

toothed. Staminate inflorescences borne in axils of 

upper leaf-whorls, 2-4 flowers developing one at a 

time, their pedicels raising them just above the 

water surface; sepals 3, green, 3-4 mm long; petals 

3, white, 8-10 mm long; stamens 9. Pistillate 

inflorescence axillary, 1-flowered, the spathe


tubular, 12-15 × 3.5-4 mm, bifid at tip with 

ciliolate teeth 1 mm long, the cleft ca. 6 mm down 

one side, through this slit issues the threadlike 

hypanthium tube; ovary 3.5 mm long, bearing at 

the tip the filiform elongate style that extends up 

through the hypanthium tube to the perianth and 

protrudes above the water surface; hypanthium 

tube white, 5.5 or more cm long; sepals 3-3.2 mm 

long, ovate, pale, greenish, reflexed at anthesis; 

petals 6-7 mm long, unequal, white, broadly 

elliptic, obtuse at apex, delicate, veinless; 

staminodes 3, at first white, becoming red, 

minutely glandular-papillose; style branched at 

apex into 3 stigmata, 3-3.3 mm long, these cleft 

into 3-4 finger-like, hairy lobes; nectaries present 

2. HALOPHILA 

Halophila Thouars, Gen. Nov. Madagasc. 2. 1806. 

at base of style-branches. Fruit and seeds 

unknown. 

General distribution: Native to South 

America. Cultivated in aquaria and fish ponds 

nearly worldwide, and often escaping and 

becoming naturalized in warm-temperate and 

tropical countries. Often classified and listed as 

Egeria densa, but the generic distinction from 

Elodea is based on characters that seem to be 

minor and unconvincing, of no more than 

subgeneric value. 

Distribution in Puerto Rico: Cited by Liogier 

& Martorell (1982) as occurring “in water near 

Corozal”; not seen by the author. 

Submerged (often deeply), rooted, monoecious or dioecious, marine herbs with slender, widely 

creeping rhizomes bearing erect, short shoots at nodes, and 2 scales, one clasping the shoot, the other the 

rhizome, sometimes with an extra pair of scales between the base of shoot and petioles. Leaves two or 

more in pairs or pseudowhorls, sessile or petiolate, linear to elliptic or ovate, entire or serrulate, glabrous 

or minutely pubescent. Flowers unisexual, solitary or sometimes a staminate and a pistillate flower 

together in a single spathe; spathes sessile, consisting of 2 overlapping membranous bracts, usually 

axillary to erect shoots. Staminate flowers pedicellate; tepals 3; stamens 3, the anthers 2- or 4-locular; 

pollen grains globose, attached together in moniliform chains. Pistillate flowers sessile, with elongated 

hypanthium, crowned by 3 reduced tepals; ovary ellipsoid, unilocular, the styles 3-5, linear. Fruit ovoid, 

rostrate, with membranous pericarp; seeds globose, few to several. A cosmopolitan genus of 8 species, at 

least some occurring in all tropical and subtropical seas, a few extending into warm temperate areas. 

LECTOTYPE: Halophila ovalis (R. Br.) Hook. f. ( ≡ Caulinia ovalis R.Br.), designated by Rydberg, N. 

Amer. Fl. 17: 67. 1909. 

Reference: Short, F. L. & M. L. Cambridge. 1984. Male flowers of Halophila engelmannii: 

Description and flowering ecology. Aquatic Bot. 18(4): 413-416. 

Key to the species of Halophila 

1. Leaves in pairs only; shoots with a pair of scales at base ……..................................…. 2. H. decipiens 

1. Leaves mostly in pseudowhorls of 2-4 pairs; shoots with 2 pairs of scales, one at the base and another 

between the base and the petioles …………................................................................................…… 2 

2. Individual leaves distinctly petiolate, with 3-5 pairs of cross veins …………..... 1. H. baillonis 

2. Leaves subsessile, with 6-8 pairs of cross veins ………….........................….. 3. H. engelmannii 

1. Halophila baillonis Asch. in H. Moseley, J. 

Linn. Soc., Bot. 14: 317. 1874. Syntypes: St. 

Thomas, U. S. Virgin Is. Moseley s. n. (B, 

destroyed, K?); place unknown. Bertero s.n. 

(B, destroyed); Martinique. Hahn s.n. (B, 

destroyed). 

Halophila aschersonii Ostenf., Bot. Tidsskr. 24: 

239. 1902. Syntypes: St. Croix, U. S. Virgin 

Is. Lassen s. n. (C); Börgeson s. n. (C). 

Dioecious marine herb; rhizomes slender, 

branching, 0.8-1 mm in diam., with a single 

unbranched root at each node. Lateral erect shoots 

0.6-2 (-4) cm long, with 2 pairs of scales, one at the

66 


base and another about halfway up; leaves 

petiolate, in a pseudowhorl of 2-3 pairs, at the 

shoot’s apex; petioles 2-5 mm long; blades elliptic, 

5-22 × 2-5 (-8) mm, the margins finely spinulose. 

Spathes lanceolate, 5-8 mm long. Staminate 

flower on pedicel ca. 3 mm long; anthers ca. 4 mm 

long. Pistillate flower 6-7 mm long with minute 

perianth; ovary sessile, the styles 2-5, each 10-30 

mm long. Fruit spherical, 2-3 mm in diam. with a 

beak 4-5 mm long; pericarp membranous; seeds 

10-20, subglobose, apiculate at both ends. 

General distribution: Pacific coast of Panama, 

Cayman Islands, Jamaica, Puerto Rico, Virgin 

Islands, Guadeloupe, Curacao, and Brazil 

(Pernambuco). 


Islands: Recorded from Culebra, Guánica, Lajas, 

Loíza, and Mayagüez; St. Croix and St. Thomas. 

Note: The spelling “ baillonis” is herein 

retained as the International Code of Botanical 

Nomenclature (Greuter et al., 2000) only 

recommends that treating Latinized modern 

names as if they were in third declension should be 

avoided, but does not mandate their standardization 

to the ii ending. The original description made 

reference to four specimens, but we have not been 

able to examine any of them for this treatment. 

Designation of a lectotype would be desirable, 

however, it must be designated only after a careful 

study of the original material. 


Lajas: Media Luna Reef; Parguera., Aponte & 

Vélez JN-15052 (US). Loíza: Sur de la Cáncora, 

Vicente s.n. (US). Mayagüez: Punta Arenas, Díaz- 

Piferrer 2050, 2744 (US). ST. CROIX: West of Buck 

Island, Abbott 11893 (US); Salt River, Dill & 

Geology class 14185 (US). 

2. Halophila decipiens Ostenf., Bot. Tidsskr. 24: 

260. 1902. Lectotype: Thailand; Koh Kahdat. 

J. Schmidt 540 (C), designated by R.A. 


Halophila baillonis sensu Britton & P. Wilson, 

1923 (and other American authors), not 

Ascherson, 1874. 

Monoecious marine herb; rhizome threadlike, 

branching, 0.2-0.4 mm in diam., fragile, with a 

single unbranched root at each node. Lateral erect 

shoots each with 1 pair of leaves at apex, and a pair 

of scales at the base. Leaves opposite; petioles 3- 

15 mm long; blades oblong-elliptic, 10-18 (-25) × 

3-6 mm, membranous, the margins finely 

serrulate, adaxially minutely pubescent, abaxially 

glabrous. Spathes ovate, acuminate, 3-6 mm long, 

with a staminate and a pistillate flower. Staminate 

flower on pedicel 3 mm long, with tepals 1-1.5 mm 

long; anthers ca. 1 mm long. Pistillate flower 

subsessile; hypanthium 1-2 mm long; ovary ca. 1 

mm long, the 3 styles each 2.5 mm long. Fruit 

ellipsoid, 2.5 mm long, the beak 1.5-2 mm long; 

pericarp scarious; seeds ca. 30, ovoid, ca. 0.2 mm 

long. 

General distribution: Pantropical. 


Islands: Often occurs in shallow water (less than 1 

m deep), but it has been found as deep as 90 m. 

Recorded from Cabo Rojo, Culebra, Guánica, 

Guayanilla, Lajas, Mayagüez, and Yabucoa; 

Guana Island, St. Croix, and St. Thomas. 


Guánica: Cayos de Caña Gorda, 1 mi. offshore 

near Guánica, Gittins 7557 (US). Guayanilla: 

Guayanilla Bay, Vicente s.n. (US). Lajas: ca. 3 mi. 

S of La Parguera, Earle 66219 (US). ST. CROIX: 

Christiansted Lagoon, Ostenfeld 290 (NY). 

3. Halophila engelmannii Asch. ex Neumayer, 

Anl. Wiss. Beobacht. Reisen, 368. 1875. 

Type: United States; coast of Florida. 

Unknown collector (B?). 

Dioecious marine herb; rhizome slender, 

branching, 1-1.3 mm in diam., with a single 

unbranched root at each node. Lateral erect shoots 

mostly 2-4 (-10) cm long, bearing 2 pairs of 

lanceolate to obovate scales, one at the base, the 

other about half way to the top, and at the apex 2- 

4 pairs of subsessile leaves, arranged in a 

pseudowhorl; blades oblong or elliptic-oblong, 

10-25 (-30) × 3-6 mm, coriaceous, with obtuse 

apex and cuneate base, the margins minutely 

serrulate. Spathe lanceolate, acuminate, sessile in 

the axil of a leaf, enclosing a single flower. 

Staminate flowers on 4-10 mm long pedicels; 

tepals elliptic, reflexed; anthers 4 mm long. 

Pistillate flowers consisting of a hypanthium 3-5 

mm long, a minute 3-parted perianth and a sessile 

or subsessile, ovoid ovary 3-4 mm long, the styles 

3, ca. 3 cm long. Fruit and seeds not known. 

General distribution: Coasts of Florida and 

the Gulf of Mexico to Texas; Bahamas, northern


Cuba, Cayman Islands, and Puerto Rico. 

Distribution in Puerto Rico: Recorded from 

two localities, Cayo Pájaros, Salinas (Vicente s.n. 

US); and Bahia de Tallaboa, Peñuelas (Britton & 

Cowell 1325). The latter specimen had originally 

been misidentified and reported as Halophila 

aschersonii Ostenf. (H. baillonis Asch.). 


Salinas: Cayo Pájaros, Jobos Bay, Vicente s.n. 

(US). 

3. LIMNOBIUM 

Limnobium Rich., Mém. Cl. Sci. Math. Inst. Natl. France 1811(2): 32, 66. 1814. 

Perennial, monoecious herbs of fresh water, usually floating, spreading by means of stolons, and 

forming mats; sometimes rooted in mud, the roots elongate. Leaves strongly and unequally petiolate, 

forming rosettes that bud from the ends of stolons and remain attached to them; leaf blades ovate to 

suborbicular. Flowers unisexual, arising from leaf axils. Staminate flowers often 2 or 3 together, maturing 

sequentially, the pedicels elongating at anthesis; sepals and petals 3 each (rarely 4); stamens variable in 

number, with filaments united into a column, the anthers linear, attached at different levels; staminodes 

usually present. Pistillate flowers solitary, enclosed in 2-cleft spathes; sepals rudimentary; petals 

rudimentary or wanting; ovary unilocular, usually with 6 placentas; styles 6, with deeply divided linear 

stigmas. Fruit a globose or ellipsoid, many-seeded berry. A unispecific genus of the western hemisphere, 

its single species divided into two subspecies. 

TYPE: Limnobium spongia (Bosc) Steud. ( ≡ Hydrocharis spongia Bosc). 

Reference: Lowden, R. M. 1992. Floral variation and taxonomy of Limnobium L. C. Richard 

(Hydrocharitaceae). Rhodora 94: 111-134. 

1. Limnobium spongia subsp. laevigatum 

(Humb. & Bonpl. ex Willd.) Lowden, 

Rhodora 94: 129. 1992; Salvinia laevigata 

Humb. & Bonpl. ex Willd., Sp. Pl. 5: 537. 

1810; Limnobium laevigatum (Humb. & 

Bonpl. ex Willd.) Heine, Adansonia, ser. 2, 8: 

315. 1968; Hydromystria laevigata (Humb. & 

Bonpl. ex Willd.) Hunz., Lorentzia 4: 5. 1981. 

Type: Colombia; Bogotá. Humboldt & 

Bonpland s. n. (holotype: B-Willd. 20251). 

Hydromystria stolonifera G. Mey., Prim. Fl. 

Esseq. 153. 1818; Limnobium stoloniferum 

(Meyer) Griseb., Fl. Brit. W. I. 506. 1864. 

Type: Guyana; Essequibo River. Meyer s. n. 

(holotype: GOET). 

Interconnected rosette-plants, either floating 

or rooted in mud. Leaves unequally petiolate, the 

petioles 0.3-3 cm long (rarely to 15 cm); blades 

ovate to suborbicular, 1.5-4 (-8) × 1.3-5 (-6) cm, 

the apex obtuse or rounded, the base short-cuneate 

or truncate, the margins entire, glabrous, fleshy or 

spongy. Staminate flowers with lanceolate sepals 

and white, often bilobed linear petals; fertile 

stamens (3-) 6 (-10); staminodes 0-3; filaments 

shorter than the anthers. Pistillate flowers with a 

subulate hypanthium and 0-8 staminodes; stigmatic 

lobes commonly 8-12. Fruits ellipsoid, 10-15 × 3.5 

mm, with 3-6 protruding placentas; seeds 0.1-0.3 

mm long, beaked and covered with minute 

trichomes. 

General distribution: Mexico, Central 

America, Greater Antilles (except Jamaica), 

Lesser Antilles, Trinidad, and South America as 

far south as Argentina but rare or lacking in most 

of the Amazon basin. 

Distribution in Puerto Rico: Very rare, in 

shallow ponds, sluggish shaded streams, and in 

fresh-water ditches or swamps at or near sea level. 

It is noteworthy that this species grows at 

elevations of up to more than 2, 000 meters in 

South America. Recorded from Añasco, Loíza, 

Mayagüez, Río Grande, and Vega Alta. 

Common name: Puerto Rico: Cuchara. 


Añasco: Sintenis 5779 (US). Mayagüez: N.L. 

Britton 2383 (US). Río Grande: Bo. Herreras, 

Road 187, Proctor & Thomas 45238 (US).

68 

Najas L., Sp. Pl. 1015. 1753. 


4. NAJAS 

Monoecious or dioecious submerged, aquatic herbs of fresh or brackish water, often rooted with 

fibrous roots: stems very slender, often much branched. Leaves linear or linear-lanceolate, opposite or 

crowded and apparently whorled, sheathing at base, the margins minutely spinescent or sometimes 

toothed. Flowers solitary, axillary, very small, lacking a true perianth. Staminate flower subtended by an 

inner hyaline scale and an outer spathe-like bract; stamen solitary, sessile, the anther 1- to 4-locular. 

Pistillate flowers consisting of a single naked ovary; style short or lacking; stigmata 2-4. Fruit a 1-seeded 

drupelet with a membranous coat. A cosmopolitan genus of about 50 species. 

TYPE: Najas marina L. 

References: Les, D. H., Cleland, M. A., & Waycott, M. 1997. Phylogenetic studies in Alismatidae, II: 

Evolution of marine Angiosperms (Seagrasses) and hydrophyly. Syst. Bot. 22: 443-463. Lowden, R. M. 

1986. Taxonomy of the genus Najas (Najadaceae) in the Neotropics. Aquatic Bot. 24: 147-184. 

Note: This taxon has until recently been treated as a separate family Najadaceae. However, cladistic 

studies (Les et al., 1997) suggest that Najas belongs in the Hydrocharitaceae, and it is so treated here. 

Key to the species of Najas 

1. Leaves flaccid, with minutely spinescent margins; plants monoecious ……....….. 1. N. guadalupensis 

1. Leaves stiff, with coarsely toothed margins; plants dioecious ……………..........…… 2. N. marina 

1. Najas guadalupensis (Spreng.) Magnus, Beitr. 

Kenntn. Najas 8. 1870; Caulinia guadalupensis 

Spreng., Syst. Veg. 1: 20. 1824. Type: 

Guadeloupe. Bertero s. n. (lectotype: TO; 

isolectotype: M), designated by Lowden, 

Aquatic Bot. 24: 169. 1986. 

Najas major sensu Bello, Anales Soc. Esp. Hist. 

Nat. 12: 115. 1883, non All., 1785. 

Fig. 12. A-D 

Plants forming dense tangles rafts in fresh or 

brackish water. Leaves opposite or clustered; 

blades linear, 10-25 × 0.5-1.5 mm, the apex acute 

or obtuse, the margins minutely spinescent or 

rarely entire; sheaths widened, with 5-10 small 

teeth. Stamens 2-3 mm long; pistil 1-2.5 mm long. 

Fruit ca. 2 mm long, spinulose-tipped; seeds 

narrowly ellipsoid, conspicuously reticulate. 

General distribution: Southeastern United 

States west to California; Mexico to Panama, 

Bahamas (Inagua), Greater Antilles, Antigua, 

Guadeloupe, Aruba, Curaçao, and South America 

as far south as Argentina. 

Distribution in Puerto Rico: In ponds, 

ditches, slow-moving streams and rivers at low 

elevations (sea level to ca. 250 m). Recorded from 

Bayamón, Cabo Rojo, Caguas, Ciales, Coamo, 

Dorado, Guánica, Utuado, and Vega Baja. 


Bayamón: N.L. Britton et al. 3830 (US). Cabo 

Rojo: Bo. Llanos Costa, Proctor & McKenzie 

43936 (US). Caguas: Sintenis 2533 (US). Coamo: 

Coamo Springs and river, N.L. Britton et al. 5803 

(US). Dorado: N.L. Britton et al. 6707 (US). 

Guánica: Guánica Lake, N.L. Britton & Shafer 

1869 (US). Utuado: N.L. Britton & Cowell 435 

(US). Vega Baja: Bo. Cibuco, in marshy pond area 

just SW of playa Cerro Gordo, Proctor et al. 45570 

(US). 

2. Najas marina L., Sp. Pl. 1015. 1753. Lectotype: 

“Habitat in Europae maribus.” (LINN- 

1156.1), designated by Viinikka, Ann. Bot. 

Fenn. 13 : 128. 1976. 

Fig. 12. E-I 

Submerged dioecious herb, branching 

profusely and forming dense masses; stems brittle, 

armed with numerous brownish spinulose teeth on 

the internodes; branches many, often 20-40 cm 

long. Leaves opposite, linear, 1-4.5 cm long, stiff, 

the margins beset with coarse triangular teeth, 1 

mm long or more; basal leaf sheaths widened, 

without teeth. Staminate flower subsessile, with 

outer envelope and stamen enclosed in a thin, 

tubular perianth. Pistillate flower solitary in upper


Fig. 12. A-D. Najas guadalupensis. A. Habit. B. Leaf. C. Infructescence. D. Fruit. E-I. Najas marina. E. Habit. F. Leaf. G. 

Infructescence. H. Pistillate flower. I. Seed. (A, from Proctor 45570; B-D, from Proctor 44116; E, G-I, from Proctor 45529; F, 

from Proctor 46143).

70 


axils. Fruits 3-4 mm long, reticulate; seeds 

ellipsoid, ca. 3 mm long, pitted with many 

irregular rows of polygonal areoles. 

General distribution: North America from 

New York to California and southward; Mexico 

and Central America (rare); Bahamas, Greater 

Antilles (very rare in Cuba and Puerto Rico); 

South America south to Argentina (apparently 

absent from the Amazon basin); widespread in 

Eurasia, and Australia; apparently absent from 

Africa. 

Distribution in Puerto Rico: Recorded only 

from Arecibo and Manatí. 


Arecibo: Bo. Garrochales, Ciénaga Tiburones, 

Proctor et al. 46143 (US). Manatí: Western end of 

Laguna Tortuguero, N.L. Britton 7878 (GH, NY, 

US). 

5. THALASSIA 

Thalassia Banks & Sol. ex K. D. Koenig, Ann. Bot. (König & Sims) 2: 96. 1805. 

Perennial dioecious marine herbs with elongate, scaly, often contorted rhizomes, producing short 

erect shoots at nodes. Leaves tufted, 2-6 together, eligulate, linear, strap-shaped, often somewhat falcate 

and sheathing at the base. Inflorescence solitary, arising from leaf axils, bearing 1(2) staminate and 1 

pistillate flowers; peduncles enclosed at base within a tubular spathe that is 2-cleft toward the apex; 

perianth segments 3, strap-like; stamens (3-) 9 (-12); filaments very short or obsolete; ovary muricate, 

unilocular, the 6-8 styles, each split into 2 filiform segments with stigmas longer than the styles. Fruit 

globose, echinate, with fleshy pericarp, splitting irregularly; seeds few, pyriform. A genus of 2 species, 

one occurring in the Caribbean area, the other found in the Indian Ocean and western Pacific Ocean 

regions, including the East Indian Archipelago and the Red Sea. 

LECTOTYPE: Thalassia testudinum Banks & Sol. ex K. D. Koenig, designated by Rydberg, N. Amer. Fl. 

17: 73. 1909. 

1. Thalassia testudinum Banks & Sol. ex K. D. 

Koenig, Ann. Bot. (König & Sims) 2: 96. 

1805. Type: Antigua. Smeathman s. n. (BM). 

Fig. 13. A-E 

Plants submerged except sometimes at lowest 

tides; rhizomes branching dichotomously, 3-6 mm 

thick with internodes 4-7 mm long, clothed with 

loosely appressed ovate scales; erect leafy shoots 

produced at intervals of 5-10 internodes, with 

persistent sheaths 6-10 cm long. Leaf linear, (4-) 

10-50 × 0.4-1.2 cm, the apex obtuse, the margins 

entire to minutely serrulate. Staminate flower: 

peduncle 3-7 cm long; spathe 1.5-3 cm long; tepals 

2-10 mm long; stamens 9, the anthers oblong, 8-9 

mm long. Pistillate flower: peduncle 3-4 cm long; 

spathe 1.5-2 cm long; ovary ca. 1 cm long, bearing 

7 or 8 styles, each 1.5-2 cm long. Fruit ellipsoid or 

globose, l.5-2 cm long with a beak 4-7 mm long, 

the pericarp echinate, splitting irregularly; seeds 

usually 3, pyriform, to 10 mm long. 

General distribution: Sheltered marine habitats 

throughout the Caribbean Sea and Gulf of Mexico, 

also Bermuda and the Atlantic coast of Florida and 

in the Bahamas. 


Islands: In shallow marine waters from low tide 

level down to a depth of ca. 10 meters, forming 

extensive dense colonies. Common in sheltered 

coastal waters all around Puerto Rico and the 

Virgin Islands but recorded by voucher specimens 

only from Ceiba, Culebrita, Fajardo, Guánica, 

Guayanilla, Mayagüez, and Río Grande; Anegada, 

Guana Island, St. Croix, St. Thomas, St. John, 

Tortola, and Virgin Gorda. 

Common names: Puerto Rico: Hierba de 

Manatí, Palma de mar. 


Guánica: Playa Tamarindo, Bahia Ballena, S of 

Guánica, Gittins 7598 (US). Guayanilla: Playa 

Tamarindo, Bahia Ballena, Gittins 7598 (US). ST. 

CROIX: Smuggler’s Cove, Phillips s.n. (US-3); 

Phillips s.n. (US-3); Phillips s.n. (US-2); Romney 

Point, N shore, E end, Ogden 624 (US). ST. JOHN: 

Reef Bay Quarter, Gentry Bay, Acevedo-Rdgz. 

2732 (US).


Fig. 13. A-E. Thalassia testudinum. A. Habit and detail of retiulate venation. B. Staminate flower. C. Stamen. D. Pistillate flower 

and l.s. pistillate flower. E. Fruit. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78. 

Family 6. CYMODOCEACEAE Manatee-grass Family 

Cymodoceaceae N. Taylor, N. Amer. Fl. 17: 31. 1909, nom. conserv. 


Monoecious or dioecious, submerged, perennial herbs with slender, elongate, monopodial, rhizomes 

rooted in marine sand. Leaves closely distichous, flattened-linear and several-nerved or cylindrical and 

lacking an evident midvein; sheathed at base. Flowers unisexual, either solitary and axillary or else few 

to several on stalked axillary cymes. Perianth absent. Staminate flower consisting of two partially fused 

filaments each with a 2- or 4-locular anther that are distally fused and longitudinally dehiscent; pollen 

threadlike. Pistillate flowers consisting of 2 free carpels with long, slender styles, these simple or 2- or 3branched, 

terminating in slender stigmas; carpels with a single apical pendulous ovule. Pollination 

effected underwater. Fruit indehiscent, with a hard, nutlike endocarp and a solitary seed. A family of 5 

genera and 16 species, distributed around the world in mostly shallow marine waters of tropical, 

subtropical, and warm temperate regions. 

TYPE: Cymodocea K.D. Koenig, nom. conserv. 

References: den Hartog, C. 1964. An approach to the taxonomy of the sea-grass genus Halodule Endl. 

(Potamogetonaceae). Blumea 12: 283-312. den Hartog, C., 1970. The sea grasses of the world. Verh. Kon. 

Ned. Akad. Wetensch., Afd. Natuurk., Tweede Sect.59 (1): 144-186.

72 


Note: The author follows Dahlgren et al. (1985) and Haynes & Holm-Nielsen (2003) in treating 

Halodule within the family Cymodoceaceae instead of Potamogetonaceae. 


1. Leaf blades flat, toothed at the apex; flowers solitary; style single ….................….. 1. Halodule 

1. Leaf blades subulate, terete or nearly so; flowers in cymose inflorescences; styles 2 …. 2. Syringodium 

Halodule Endl., Gen. Pl. 1368. 1841. 

1. HALODULE 

Dioecious marine herbs with rooted, creeping rhizomes, at each node these producing a short upright 

stem bearing 1-4 leaves. Leaf sheaths persistent; leaf blades narrowly linear, flat. Flowers solitary, each 

enclosed in a leaf-sheath. Staminate flower stalked, consisting of two anthers at different levels. Pistillate 

flower subsessile, consisting of two free carpels with long styles. Fruit drupe-like, subglobose-ovoid, 

slightly compressed and with a short beak. A genus of 6 species widely distributed in shallow tropical seas. 

TYPE: Halodule tridentata (Steinh.) Endl. ex Unger (≡ Diplanthera tridentata Steinh.). 

Key to the species of Halodule 

1. Leaf apex acute and mucronate (one large central tooth and two, minute lateral ones) …. 1. H. beaudettei 

1. Leaf apex emarginate (with two equal lateral projecting teeth) ……...........….. 2. H. wrightii 

1. Halodule beaudettei (Hartog) Hartog, Blumea 

12: 303. 1964; Diplanthera beaudettei 

Hartog, Pacific Naturalist 1(15): 4. 1960. 

Type: Panama; Jicarón Is. Dawson 21104 

(holotype: US!). 

Rhizomes creeping, with 2-4 roots and a 

short, erect stem at each node; internodes 2-4 cm 

long. Leaf sheaths 0.5-6 cm long; blades narrowly 

linear, 5-20 cm × 0.5-1.5 mm, the apex acute, 

mucronate, the mucron 1-10 times as long as the 

narrow lateral teeth. Flowers born within sheaths 

of the branches just above the rhizome. Staminate 

flower consisting of 2 anthers, inserted on a stalk 

that contains tannin cells on either side of the stalk 

just below the higher anther. Pistillate flower 

consisting of 2 carpels on a stout stalk, one inserted 

obliquely above the other, and tannin cells on 

either side of the stalk just below the upper carpel, 

ovary asymmetrical, with a single, filiform, lateral 

style. Nearly mature fruit yellow, bulb-like. 


States from North Carolina to Florida, the Gulf of 

Mexico, the Caribbean coast of Central America, 

Bahamas, Cuba, Cayman Islands, Jamaica, 

Hispaniola, Virgin Islands, Guadeloupe, Marie 

Galante, Trinidad, Curacao, and Venezuela. 


Islands: Coastal marine areas. Not collected yet in 

Puerto Rico, but expected there; St. Croix and St. 

Thomas. 

Selected specimens examined: ST. CROIX: 

Krause Lagoon, Børgesen s. n. (K, C). ST. THOMAS: 

Saen ven Tutu, Krebs s. n. (C); idem Hornemann s. 

n. (C); Mosquito Bay, Ostenfeld 182 (C). 

2. Halodule wrightii Asch., Sitzungsber. Ges. 

Naturf. Freunde Berlin 1868: 19. 1868; 

Diplanthera wrightii (Asch.) Asch. in Engler 

& Prantl., Nat. Pflanzenfam., Nachtr. 1: 37. 

1897. Type: Cuba. Wright 3720 (holotype: K; 

isotype: US!). 

Rhizomes jointed, creeping. Leaf sheaths 1.5- 

4 cm long; blades 5-8 cm × 0.3-1 mm, the apex 

appearing emarginate (two lateral projecting 

teeth). Staminate flower on stalk 12-23 mm long; 

anthers 3.5-5 mm long, containing tannin cells. 

Pistillate flower consisting of an ellipsoid, ovoid 

or globose ovary, 1.5-2 mm long, the style apical


or sub apical, 10-28 mm long. Mature fruit brown 

or blackish, ovoid or globose, 1.5-2 × 1.5 mm, 

farinaceous, apiculate. Seldom seen in flower or 

fruit. 

General distribution: Greater Antilles, 

Cayman Islands, Lesser Antilles, the coast of East 

Africa (Kenya, Tanzania, Mozambique), 

Madagascar, Mauritius, and the west coast of 

Australia. 


Islands: In most coastal marine areas. Collected 

only from Cabo Rojo, Ceiba, Culebrita Island, 

Guánica, Guayanilla, and San Juan; St. Croix, St. 

2. SYRINGODIUM 

Syringodium Kütz. in Hohenacker., Alg. Mar. Exsicc. 9: 426. 1860. 

John, and St. Thomas. 


Cabo Rojo: Joyuda Lagoon, Vicente s.n. (US-3). 

Ceiba: Roosevelt Roads Naval Research Station, 

Bo. Guayacán: Pelican Bay, between Punta 

Algodones & Punta Cascajo, Proctor et al. 48039 

(US). Guayanilla: Guayanilla Bay, Phillips s.n. 

(US-5). ST. CROIX: Christiansted Lagoon, Børgesen 

s.n. (US); Long Point, Phillips s.n. (US-3). ST. 

JOHN: Lameshur Bay, Phillips s.n. (US); Randall 

s.n. (US). ST. THOMAS: Mosquito Bay, Ostenfeld 

s.n. (US). 

Dioecious marine herbs with branched, widely creeping, monopodial rhizomes, rooted in sand, 

bearing at frequent intervals short, upright shoots bearing 2 or 3 leaves with persistent sheaths at base. Leaf 

blades subulate. Inflorescence cymose, the lower branches dichasial. Individual flowers enclosed by the 

bases of a pair of reduced bract-like leaves. Staminate flower stalked, consisting of two basally connate 

anthers. Pistillate flowers sessile, consisting of two, free ovaries, each with a short style which divides into 

two linear stigmas. Fruit drupe-like, obliquely ellipsoid, with a short bifid apex. A genus of two species, 

one in the Caribbean, the other occurring in the Indian and western Pacific Oceans. 

TYPE: Syringodium filiforme Kütz. 

Reference: Tomlinson, P. B. & U. Posluszny. 1978. Aspects of floral morphology and development 

in the seagrass Syringodium filiforme (Cymodoceaceae). Bot. Gaz. 139: 335-345. 

1. Syringodium filiforme Kütz. in Hohenacker., 

Alg. Mar. Exsicc. 9: 426. 1860. Lectotype: St. 

Thomas, U. S. Virgin Islands. Hohenacker 

426 (BM), designated by Dandy & Tandy, J. 

Bot. 77: 115. 1939. 

Cymodocea manatorum Asch., Sitzungsber. Ges. 

Naturf. Freunde Berlin 1868: 19. 1868. Type: 

To be sought among several syntypes. 

Fig. 14. A-E 

Rhizomes tough and wiry. Leaf sheaths 2.5-6 

cm long; blades 10-30 cm long, 0.8-2 mm thick, 

narrowed at the base. Staminate flowers on stalks 

5-10 mm long; anthers ovate, 4-5 mm long. 

Pistillate flowers: ovary ellipsoid, 3-4 mm long; 

style 2-3 mm long; stigmas 4-6 mm long. Fruit 

obliquely obovoid, 6-7 mm long, the bifid apical 

portion (rostrum) 2-3 mm long. 

General distribution: Bermuda and throughout 

the Caribbean Sea and Gulf of Mexico, but absent 

from the shores of South America. 


Islands: Collected in Cabo Rojo, Ceiba, Culebrita, 

and Río Grande, but undoubtedly occurs along 

most coasts of Puerto Rico and adjacent islands; 

Guana Island, Peter Island, St. Croix, St. John, and 

St. Thomas. 

Common name: Puerto Rico: Yerba manatí. 


Culebra: Culebrita, Proctor 45063 (US). Guánica: 

Acevedo-Rdgz. 3687 (US). ST. CROIX: Knight Bay, 

ca. ½ km from shore, Fosberg 56809 (US); Eggers 

s.n. (US). Frederiksted: Børgesen s.n. (US). ST. 

JOHN: Tektite Project, transect 1, Earle s.n. (US); 

Great Lameshur Bay, Earle s.n. (US).

74 


Fig. 14. A-E. Syringodium filiforme. A. Habit. B. L.s. bracteal leaf showing stamen, and portion of inflorescence with staminate 

flower. C. Portion of inflorescence showing pistillate flowers. D. L.s. pistil, l.s. bracteal leaf showing pistils, and pistillate 

flower. E. Fruit with subtending bracts, and l.s. fruit. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. 

Gard. 78. 

Family 7. RUPPIACEAE Ditch-grass Family 

Ruppiaceae Horan., Fam. Pl. 2: 48. 1934, nom. conserv. 

by G. R. Proctor & P. Acevedo-Rodríguez 

Submerged aquatic herbs of fresh, brackish, or sometimes saline water, with long, thread-like, forking 

stems. Leaves alternate or opposite; blade long-linear; sheath loose, auriculate. Inflorescence of spikes, 

usually 2-flowered, initially produced in leaf-axils within the basal sheath, then becoming protruded by 

elongation of the peduncle. Flowers bisexual; perianth absent; stamens 2, the filaments very short, with 2locular 

anthers, the locules separated by a connective; carpels 2-19, free, sessile, each with a single, 

pendulous or campylotropous ovule, the style lacking, the stigma peltate. Fruit a minute drupe, at first 

sessile, usually becoming long-stipitate as it ripens. A family of a single genus and ca. 10 species, of 

cosmopolitan distribution. 

TYPE: Ruppia L. 

Note: Some authors, including Dahlgren et. al. (1985) and Strong in Acevedo-Rodríguez (1996), have 

included Ruppia in the Potamogetonaceae. However, Ruppiaceae differs from Potamogetonaceae in (1) 

lacking a perianth; (2) having 2 instead of 4 stamens; (3) having unique isobilateral instead of globose 

pollen; (4) having ovules pendulous from the apex of the ovary instead of ventromarginal; (5) having long-


stipitate instead of sessile fruits; (6) having a chromosome count of 8-10 instead of 13-15; and (7) growing 

chiefly in brackish, saline, or alkaline water instead of chiefly in fresh water. Because of these differences, 

the authors are maintaining two separate families. 

Ruppia L., Sp. Pl. 127. 1753. 

1. RUPPIA 

Characters as given above for the family. A genus of nearly world-wide distribution, with ca. 10 

species. 

TYPE: Ruppia maritima L. 

Reference: Jacobs, S. W. L. & M. A. Brock. 1982. A revision of the genus Ruppia 

(Potamogetonaceae) in Australia. Aquatic Bot. 14: 325-337. 

Key to the species of Ruppia 

1. Peduncle of inflorescence 0.6-5(11.5) mm long, not becoming greatly elongated after fruiting; druplet’s 

stipe 0-2 mm long ……................................................................................................….. 1. R. didyma 

1. Peduncle of inflorescence 15-50 mm long, sometimes up to 30 cm after fruiting; druplet’s stipe (3)5-30 

mm long ………............................................................................................................… 2. R. maritima 

1. Ruppia didyma Sw. ex Wikstr., Kongl. 

Vetensk. Acad. Handl. 1825: 427. 1825. 

Type: St. Barthelemy. Forsström s. n. 

(holotype: S). 

Ruppia anomala Ostenf., Bull. Torrey Bot. Club. 

42: 659. 1915. Type: Puerto Rico. Britton & 

Shafer1870 (holotype: C; isotypes: NY!, 

US!). 

Submerged plants, rooted in mud, the stems 

0.5-1(1.4) mm in diam. Leaf sheaths 7-10 mm 

long; blades linear or filiform, 2-7 cm × 0.5-0.9 

mm, acute at apex. Peduncles 0.6-5(11.5) mm 

long, not elongating after anthesis. Drupelets 

ovoid, 2-3 mm long, with pedicel-like stipes 0-2 

mm long. 

General distribution: Mexico (Yucatan), 

scattered throughout the West Indies. 


single collection from Guánica lagoon. 


Guánica: border of Guánica Lake, Britton & 

Shafer 1870 (type of Ruppia anomala Ostenf., 

Bull. Torrey Bot. Club. 42: 659. 1915, holotype: 

NY!; isotype: US!). 

2. Ruppia maritima L., Sp. Pl. 127. 1753. 

Lectotype: Italy. Micheli, Nov. Pl. Gen. , t. 35. 

1729, designated by Setchell, Proc. Calf. 

Acad. Sci., ser. 4. 25: 470. 1946. 

Fig. 15. A-F 

Submerged plants, rooted in mud, with light 

green to whitish stems to 1 m long. Leaf sheaths 6- 

10 mm long; blades linear or filiform, 3-11 cm × 

0.3-0.8 mm, narrowly acuminate at apex. 

Peduncles 1.5-5 cm long, elongating after anthesis 

to 30 cm long. Druplets ovoid, 2-3 mm long, with 

pedicel-like stipes usually (0.3)0.5-3 cm long in 

umbellate clusters. 

General distribution: Cosmopolitan. 


Islands: Near sea level in brackish or saline ponds, 

lagoons, estuaries, and in sheltered mangrove 

habitats. Recorded from Arecibo, Bayamón, Cabo 

Rojo, Ceiba, Culebra, Guánica, Guayanilla, and 

Vieques; Anegada, Guana Island, St. Croix, St. 

Thomas, and St. John. 

Common name: Puerto Rico: Yerba de zanja. 


Arecibo: Bo. Garrochales, Ciénaga Tiburones, in 

immediate outflow of artesian well beside 

centerline road 0.4 km S of Caño Tiburones, 

Proctor et al. 46136 (US). Bayamón: Sintenis 

1020 (US). Culebra: Flamingo Bay, Howe 195 

(US). Guánica: Guánica Lake, N.L. Britton et al. 

5510 (US). Guayanilla: Tamarindo Beach, S of 

Guánica State Forest, Strong et al. 459 (US). 

GUANA ISLAND: White Bay, Proctor 48402 (US). 

ST. CROIX: Great Pond, McMillan 56792 (US); 

Buck Island, N.L. Britton & Shafer 657 (US). ST. 

THOMAS: Hornbeck s.n. (US).

76 


Fig. 15. A-F. Ruppia maritima. A. Habit. B. Portion of flowering culm. C. Detail of culm with inflorescence hidden by leaf 

sheaths. D. Inflorescence, stamen, and carpel. E. Infructescence. F. Drupelet. From Acevedo-Rdgz., P. 1996, Flora of St. John, 

Mem. New York Bot. Gard. 78. 

Family 8. POTAMOGETONACEAE Pondweed Family 

Potamogetonaceae Dumort., Anal. Fam. 59, 61. 1829, nom. conserv. 


Perennial aquatic herbs, chiefly of fresh water, proliferating not only by seeds but also by rhizomes, 

tubers, or detached winter buds. Leaves alternate, sheathing at base, sometimes dimorphic; submerged 

leaves linear or nearly filiform; floating leaves much wider. Flowers bisexual, in pedunculate, axillary 

spikes, originating between stipular sheaths; perianth of 4 clawed valvate segments; stamens 4, attached 

to the claws of the perianth-segments; anthers 2-locular, sessile, each with a connective produced into a 

broad sepaloid appendage; ovary of 4 free carpels, each with a single ovule and a sessile stigma. Fruit a 

sessile, indehiscent, 1-seeded drupe, with fleshy or spongy exocarp and crustaceous endocarp. A 

cosmopolitan family of two genera and ca. 100 species. 

TYPE: Potamogeton L. 

Reference: Haynes, R. R. & L. B. Holm-Nielsen. 2003. Potamogetonaceae. Fl. Neotrop. Monogr. 85: 

1-52. 

Potamogeton L., Sp. Pl. 126. 1753. 

1. POTAMOGETON 

Characters as given above for the family. A genus of about 100 species of subcosmopolitan 

distribution with center of distribution in temperate regions. 

LECTOTYPE: Potamogeton natans L., designated by N. Taylor, N. Amer. Fl. 17(1): 14. 1909.


Key to the species of Potamogeton 

1. Leaves all submerged, all similar, narrowly linear; spikes 0.2-7 mm long, cylindrical or capitate 

…...................................................................................................................... 1. P. foliosus 

1. Leaves of 2 kinds, submerged and floating, the submerged ones linear or narrowly lanceolate, the 

floating ones elliptic; spikes 20-70 mm long, cylindrical …………….. 2 

2. Floating leaves coriaceous, with entire margins, slightly inconspicuous venation, and petioles 2- 

12.5 cm long ………...........................................................................................…. 3. P. nodosus 

2. Floating leaves membranous, with crenate margins, conspicuous venation, and petioles 0.5-3 cm 

long ………......................................................................................................…….. 2. P. illinoensis 

1. Potamogeton foliosus Raf., Med. Repos., ser. 

2, 5: 354. 1808; Potamogeton pauciflorus 

Pursh, Fl. Amer. Sept. 121. 1814, nom. illeg. 

Type: United States; South Carolina. Michaux 

s. n. (holotype: P, photo at GH). 

Potamogeton gramineus sensu Michaux, Fl. Bor.- 

Amer. 1: 102. 1803, non L., 1753. 

Plants rooted, entirely submerged; stems 

filiform, flattened, branched. Leaves narrowly 

linear, 2-6 cm × 0.5-2.5 mm, acute or subacute at 

apex, slightly tapering at base, 3- to 5-nerved with 

prominent mid rib, the margins entire; young 

stipules free from the leaf base, 7-18 mm long, 

soon deciduous. Peduncles 0.5-1.8 mm long; 

flower spikes capitate, few-flowered. Fruits 

lenticular or nearly orbicular, 1.8-2 × 2-2.5 mm, 

the keel crested; embryo an incomplete spiral. 

General distribution: Throughout North 

America, Central America, Greater Antilles, parts 

of South America, and Hawaii. 


Islands: Frequent in slow-flowing streams and 

rivers at low to middle elevations (sea level-400 

m). Recorded from Aguas Buenas, Bayamón, 

Coamo, Isabela, Mayagüez, Jayuya, Lares, 

Vieques, and Yauco; St. Croix. 


Aguas Buenas: Sintenis 2536 (US). Coamo: 

vicinity of Coamo Springs, N.L. Britton & Cowell 

1334 (US). Isabela: Bo. Coto, W side of Río 

Guajataca, from hwy 2 bridge to 1.2 km upstream, 

Proctor 44117 (US). Jayuya: Sintenis 6306 (US). 

Lares: Sintenis 5818 (US). Yauco: Sintenis 3489 

(US). 

2. Potamogeton illinoensis Morong, Bot. Gaz. 5: 

50. 1880. Type: United States; Illinois. 

Patterson s. n. (lectotype: NY; isolectotype: 

BM), designated by González, Taxon 36: 112. 

1987. 

Potamogeton pulchelliformis Hagstr., Ark. Bot. 

27(7): 12. 1922. Type: Haiti. Ekman H326 

(holotype: S). 

Rooted plants with both submerged and 

floating leaves; stem erect, terete. Submerged 

leaves elliptic to lanceolate or rarely linear, 5-20 × 

0.2-4.5 cm, 7- to 19-nerved, sessile or petiolate. 

Floating leaves elliptic to oblong-elliptic, 4-19 × 

2-6.5 cm, 13- to 29-nerved, the apex rounded, 

mucronate, the base cuneate, the margins entire; 

petioles 2-9 cm long; blade sheath brownish, 3-9 

cm long. Spikes cylindrical, 2.5-7 cm long; 

peduncles 4-30 cm long. Fruits laterally 

compressed, obovate, 2.5-3.6 mm long, crested, 

the embryo forming a complete spiral. 

General distribution: North and Central 

America, Greater and Lesser Antilles, and South 

America. 

Distribution in Puerto Rico: Uncommon, 

native herb, recorded from Toa Baja, Tortuguero 

and Utuado. 


Toa Baja: Río Dorado, Liogier et al. 33773 (UPR). 

Utuado: Sintenis s.n. (BM), Sintenis 6501 (MO); 

Caonillas Arriba, Sintenis 6310 (GH), 

3. Potamogeton nodosus Poir. in Lamarck, 

Encycl. Suppl. 4: 535. 1816. Type: Canary Is. 

Broussonet s. n. (P). 

Potamogeton occidentalis Sieber ex Cham. & 

Schltdl., Linnaea 2: 224. 1827. Type: 

Martinique. Sieber 275 (lectotype: LE; 

isolectotypes: G, GOET, HAL, P), designated 

by Haynes & Holm-Nielsen. Fl. Neotrop. 

Monogr. 85: 26. 2003. 

Potamogeton pennsylvanicus var. portoricensis 

Graebn. in Urban, Symb. Antill. 4: 73. 1903; 

Potamogeton nuttallii var. portoricensis

78 


(Graebn.) Graebn. in Engler, 

Potamogetonaceae. Pflanzenr. IV. 11 (Heft 

31): 56. 1907. Syntypes: Puerto Rico. Sintenis 

1025 & 2537 (S). 

Potamogeton insulanus Hagstr., Kongl. Svenska 

Vetenskpsakad. Handl. 55(5): 154. 1915. 

Type: Puerto Rico. Sintenis 2537 (S). 

Potamogeton epihydrus sensu Britton & P. 


Rafinesque, 1808. 

Potamogeton plantaginea sensu Bello, Anales 

Soc. Esp. Hist. Nat. 12: 115. 1883, non Roem. 

& Schult., 1818. 

Potamogeton fluitans sensu Britton & P. Wilson, 

Bot. Porto Rico 5: 10. 1923, and Graebn. in 

Engler, Potamogetonaceae. Pflanzenr. IV. 11 

(Heft 31): 61. 1907, non Roth, 1788. 

Rooted plants with both submerged and 

floating leaves; stem slender, nearly terete, 

branched. Submerged leaves linear- to ellipticlanceolate, 

9-20 × 1-3.5 cm, pellucid-punctate, 7to 

15-nerved, petiolate. Floating leaves broadly 

elliptic, 3-11 × 1.5-4.5 cm, 9- to 21-nerved, the 

apex acute or acuminate, the base acute to 

rounded, the margins entire; petioles 3.5-26 cm 

long; blade sheath brownish, 3.5-6.5 cm long. 

Spikes cylindrical, 2-7 cm long; peduncles 3-15 

cm long. Fruits obovoid, 2.7-4.3 mm long, crested, 

the embryo forming a complete spiral. 

General distribution: North and Central 

America, Greater and Lesser Antilles, South 

America, Europe, Africa, and Asia. 

Distribution in Puerto Rico: In slow-flowing 

streams and rivers, rarely in ponds, at low 

elevations. Recorded from Arecibo, Bayamón, 

Cabo Rojo, Dorado, Hatillo, Isabela, Loíza, 

Manatí, Maunabo, Río Grande, San Juan, San 

Sebastián, Toa Baja, Utuado, and Vega Baja. 

Common name: Puerto Rico: Lino de agua. 


Arecibo: N.L. Britton & Cowell 305 (US). 

Bayamón: Sintenis 1025, 1025b (US). Dorado: 

Woodbury s.n. (UPR). Hatillo: Sargent B86 (US). 

Loíza: Stevenson 5472 (US). Manatí : Laguna 

Tortuguero, Sintenis 6670 (US). Río Grande: Río 

del Espiritu Santo, Eggers 1191 (US). San 

Sebastián: Guajataca Gorge, Liogier et al. 30349 

(UPR). Utuado: Sintenis 6310 (US). Vega Baja: 

N.L. Britton et al. 6763 (US). Vega Baja: Bo. 

Cibuco, along Río Cibuco, ca. 1 km due SE of 

Cerro Cibuco, Proctor et al. 45628 (US). 

Family 9. SMILACACEAE Sarsaparilla Family 

Smilacaceae Vent., Tabl. Règne Vég. 2: 146. 1799, nom. conserv. 

by P. Acevedo-Rodríguez 

Herbaceous to slender woody vines aided by tendrils, or less often erect herbs or shrubs, arising from 

creeping, starchy rhizomes. Leaves alternate or sometimes opposite; petioles usually with a pair of tendrils 

at junction with short stipule-like flanges. Flowers small, commonly unisexual (the plant dioecious) or less 

often bisexual, actinomorphic, 3-merous, produced in axillary umbels, racemes, or spikes bearing lateral 

umbels; staminate flowers lacking pistillodes; pistillate flowers bearing staminodes. Perianth of 6, equal, 

petaloid tepals in 2 series, free or connate at base into a short tube; nectaries commonly at the inner base 

of the tepals or at the base of the stamens or staminodes; stamens commonly 6 or less often more numerous 

or only 3, the filaments distinct, sometimes adnate to the perianth tube, or more or less connate into a 

column, the anthers opening by longitudinal slits; gynoecium 3-pistillate, ovary superior, trilocular or 

unilocular, the placentation accordingly axile or parietal, the ovules 1 to many per locule, the styles 

terminal, distinct, or basally connate. Fruit a 1- to 6-seeded berry. A family of about 12 genera and ca. 375 

species, mostly with tropical and subtropical distribution, but occurring in subtemperate zones as well. 

TYPE: Smilax L. 

Reference: Howard, R. A. 1979. The genus Smilax L. in the Lesser Antilles. Taxon 28: 55-58. 1979.

Smilax L., Sp. Pl. 1028. 1753. 


1. SMILAX 

Dioecious, herbaceous or woody vines, with starchy rhizomes; stems cylindrical, wiry, armed or 

unarmed. Leaves alternate, simple, with arching parallel veins; petioles with a pair of filiform tendrils at 

junction with the open sheaths. Flowers unisexual, 3-merous, actinomorphic, with 6 perianth segments, 

produced in axillary umbels. Staminate flowers with 6 stamens, the anthers basifixed; pistillate flowers 

usually bearing staminodes, the ovary 3-pistillate with 1 or 2 ovules per carpel. Fruit a fleshy berry; seeds 

1-6. A genus of about 350 species from primarily tropical and subtropical areas but extending into 

temperate zones. 

LECTOTYPE: Smilax aspera L., designated by Britton & A. Brown, Ill. Fl. N. U. S. ed. 2. 1: 527. 1913. 

Key to the species of Smilax 

1. Leaves coriaceous or thick-coriaceous, usually with spiny margins, the apex obtuse to rounded; stems 

spiny ..............................................................................................................………… 1. S. coriacea 

1. Leaves chartaceous, with entire margins, the apex acuminate; stems not spiny ……… 2. S. domingensis 

1. Smilax coriacea Spreng., Syst. Veg. 2: 103. 

1825. Neotype: Puerto Rico; Cayey. Sintenis 

2252 (US), here designated. 

Smilax ilicifolia Desv. ex Ham., Prodr. 58. 1825; 

Smilax coriacea var. ilicifolia (Desv. ex 

Ham.) O. E. Schulz in Urban, Symb. Antill. 4: 

150. 1903. Type: West Indies. Desvaux s. n. 

(P). 

Smilax havanensis var. portoricensis A. DC. in 

Alph. de Candolle & C. de Candolle., 

Monogr. Phan. 1: 124. 1878. Syntype: Puerto 

Rico. Wydler 341 (FI); Dominican Republic. 

Rob. Schomburgk 71 (TO). 

Smilax coriacea Bello, Anales Soc. Esp. Hist. Nat. 

12: 120. 1883, nom. illeg., non Sprengel, 

1825. Type. Puerto Rico. Bello s.n. (destroyed). 

Smilax rotundifolia Bello, Anales Soc. Esp. Hist. 

Nat. 12: 120. 1883, nom. illeg., non L., 1753. 

Type: Puerto Rico. Bello s.n. (destroyed). 

Smilax guianensis var. subarmata O. E. Schulz in 

Urban, Symb. Antill. 4: 149. 1903; Smilax 

subarmata O. E. Schulz in Urban, Symb. 

Antill. 5: 28. 1904. Type: Puerto Rico. 

Sintenis 4943 (syntype: B, destroyed); 

Sintenis 5932 (syntype: B, destroyed). 

Smilax lappacea sensu O. E. Schulz in Urban, 

Symb. Antill. 5: 37. 1904, non Willd., 1806. 

Smilax ilicifolia var. sublappacea sensu O. E. 

Schulz in Urban, Symb. Antill. 5: 37. 1904, 

non A. DC., 1878. 

Figs. 16. A-F; 60. H 

Slightly woody vine, to 5 m long; stems 

slender, cylindrical, wiry, puberulent or slightly 

scabrous, with scattered curved spines. Leaves 

alternate, simple; blades 2-17 × 1-12 cm, ovate, 

elliptic, oblong, lanceolate or linear, coriaceous or 

thick-coriaceous, glabrous, with 3-7 parallel 

veins, the margins spiny or entire, the apex acute to 

rounded, usually mucronate, the base rounded, 

obtuse or cordate; petioles 0.5-2 cm long, slightly 

swollen, with a pair of filiform tendrils near the 

base. Inflorescence a flexuous spike bearing 

lateral umbels, or less often a solitary, axillary 

umbel. Perianth yellowish, in staminate flowers, 

2-3 mm long, in pistillate flowers 1.5-1.8 mm long. 

Berry globose or depressed-globose, 5-7 mm 

diam., turning from green to black at maturity. 


Rico, Virgin Islands, and the Lesser Antilles. 


Islands: Common in open disturbed areas, from 

low to middle elevations. Reported from Arecibo, 

Barranquitas, Bayamón, Cabo Rojo, Cayey, 

Ciales, Coamo, Fajardo, Guánica, Isabela, Jayuya, 

Manatí, Maricao, Maunabo, Mayagüez, Río 

Grande, Sabana Grande, Salinas, San Germán, San 

Juan, San Lorenzo, Vieques, and Yauco; Guana 

Island, St. Croix, St. John, St. Thomas, Tortola, 

and Virgin Gorda. 

Common names: Puerto Rico: Dunguey, 

Dunguey blanco; Virgin Islands: Greenbriar. 

Note: A neotype is here designated because

80 


the original material used by Spreng. in describing 

this species is apparently lost. In a previous 

publication (Acevedo-Rdgz., 1986), Smilax 

coriacea Spreng was treated as a synonym of S. 

havanensis Jacq, following Liogier’s (1982) 

Synoptical Flora of Puerto Rico and adjacent 

islands. However, they are distinct species easily 

recognized by the venation pattern. Smilax 

coriacea has interprimary veins borne at an angle 

of 45° to 90° from the primary vein, while those of 

S. havanensis are ascending, being borne at 25° to 

35°. Smilax havanensis occurs in Cuba and 

Hispaniola but does not reach Puerto Rico or the 

Virgin Islands. 


Arecibo: Bosque de Río Abajo, Acevedo-Rdgz. 

183 (SJ). Barranquitas: Monte Torrecilla, N.L. 

Britton et al. 5604 (US). Bayamón: Underwood 

859, 860 (US). Cabo Rojo: Sintenis 708 b (US). 

Cayey: Sintenis 2252 (US). Ciales: along trail 

Camino de la Ceiba towards Quebrada del Pozo 

Azul, Acevedo-Rdgz. & Vicens 11848 (MAPR, 

UPRRP, US). Coamo: Coamo Springs, N.L. 

Britton & Marble 2256 (US). Fajardo: playa de 

Fajardo, N.L. Britton & Shafer 1566 (US). 

Guánica: Sintenis 3798 (US). Isabela: Bo. Coto, 

Proctor 45930 (SJ). Jayuya: Toro Negro State 

Forest. Upper W summit ridge of Cerro de Punta, 

Proctor & Rivera 47081 (SJ). Manatí: Bo. Tierras 

Nuevas Saliente, S of Laguna Tortuguero, Proctor 

& McKenzie 44039 (SJ, US). Maricao: Maricao 

Insular Forest, Otero & Gregory 802 (US). 

Maunabo: Punta Tuna, Sintenis 5116 (US). 

Mayagüez: Mayagüez to Joyuda, Underwood 157 

(US). Río Grande: along Road 191 to El Yunque 

Forest Reserve, Acevedo-Rdgz. & Siaca 9330 

(UPR, US). Sabana Grande: Sargent 436 (US). 

Salinas: Bo. Lapa, summit of east peak, Las Tetas 

de Cayey, Proctor 43732 (SJ). San Germán: 

Maricao State Forest, Bo. Minillas, W side trail to 

Santana Peak S of Rd 362, Proctor et al. 48069 

(SJ). San Juan: Río Piedras, Stevenson 678 (US). 

San Lorenzo: Bo. Espino, Carite Forest Reserve, 

Cerro La Santa, forest to E of radio towers, 

Axelrod & Kay 11160 (US). Vieques: Lighthouse 

Peninsula, limestone thicket, Shafer 2809 (US). 

Yauco: Susúa Forest Reserve, Acevedo-Rdgz. 

11418 (UPRRP, US). GUANA ISLAND: near head of 

shallow ravine just NE of Palm Ghut, Proctor 

47244 (SJ). ST. JOHN: Lameshur, N.L. Britton & 

Shafer 522 (US). ST. THOMAS: Bordeaux, E.G. 

Britton & Marble 1376 (US). TORTOLA: High 

Bush, N.L. Britton & Shafer 821 (US). Yauco: 

Susúa, Liogier 9888 (US). 

2. Smilax domingensis Willd., Sp. Pl. 4: 783. 

1806. Type: Hispaniola: Richard s. n. 

(holotype: P). 

Figs. 16. G-K; 60. G 

Slightly woody vine, 5-10 m long; stems 

slender, cylindrical, wiry, glabrous, smooth, 

without spines. Leaves alternate, simple; blades 5- 

12(15) × 2-6.5 cm, ovate, oblong, elliptic, or 

widely elliptic, chartaceous, involute, glabrous, 

with 5 parallel veins, the margins entire, the apex 

acuminate, the base obtuse, rounded, or subcordate, 

upper surface slightly lustrous, lower surface with 

prominent veins; petioles 1-1.5 cm long, articulate 

at base and forming a sheath that has a pair of 

filiform tendrils, 8-12 cm long. Inflorescence 

axillary, umbelliform, solitary. Perianth greenish, 

in staminate flowers, 3-3.5 mm long; pedicel 5-9 

mm long. Berry globose, fleshy, ca. 1 cm diam., 

turning from green to black or purple at maturity; 

seed solitary. 

General distribution: Mexico, Guatemala, 

Belize, Greater Antilles and the Virgin Islands. 


Islands: Occasional in open disturbed areas, from 

low to middle elevations. Reported from Arecibo, 

Canóvanas, Ciales, Manatí, Maricao, Moca, 

Naguabo, Río Grande, San Juan, Utuado, Vega 

Alta, and Vega Baja; Tortola. 


membrillos, Raíz de zarzaparilla, Zarzaparilla. 


Arecibo: Bo. Río Arriba, area to NW of temporary 

S end of Rt. 10 (near Rt 621), adjacent to Río Abajo 

Forest Reserve, Axelrod et al. 9492 (US). 

Canóvanas: Sierra de Luquillo, Caribbean 

National Forest, W end of El Toro trail, first 1.5 km 

from Rd 186, Proctor & Taylor 46291 (SJ). Ciales: 

along trail Camino de la Ceiba towards Quebrada 

del Pozo Azul, Acevedo-Rdgz. & Vicens 11835 

(FTG, MAPR, NY, UPRRP, US). Manatí: Laguna 

Tortuguero, Acevedo-Rdgz. & Cedeño 9488 (US). 

Maricao: Maricao State Forest, Bo. Maricao 

Afuera, N side Alto del Descanso, Proctor et al. 

48067 (SJ). Moca: Sargent 362 (US). Naguabo: 

Loma Icaco, Shafer 3449 (US). Río Grande: Sierra 

de Luquillo, Bo. Jiménez. Finca San Pedrito, ca.


Fig. 16. A-F. Smilax coriacea. A. Leaf with spiny margin. B. Fertile branch. C. Branch with non-spiny leaves, and detail of 

inflorescence. D. Staminate flower, top view and longitudinal section. E. Pistillate flower, gynoecium, and longitudinal section 

of the gynoecium, showing ovule. F. Branch with fruit. G-K. Smilax domingensis. G. Vegetative branch with tendrils. H. Fertile 

branch. I. Staminate flower. J. Fruiting branch. K. Seed. From Acevedo-Rdgz., P. 2003, Bejucos y plantas trepadoras de Puerto 

Rico e Islas Vírgenes, Smithsonian Institution.

82 


0.9 km due ESE of Rd 186 at El Verde, Proctor 

50442 (SJ). Río Grande: Sierra de Luquillo, in 

monte Jiménez, Sintenis 1417 (US). San Juan: Río 

Piedras, Stevenson 3458 (US). Utuado: Bo. Don 

Alonso, Acevedo-Rdgz. et al. 7129 (UPRRP, US). 

Vega Alta: S of communication tower, Acevedo- 

Rdgz. 12288 (US). Vega Baja: Bo. Algarrobo, 


Excluded Species 

Smilax rotundifolia L., Sp. Pl. 1030. 1753, 

was listed for St. Croix by Britton & P. Wilson 

(1923) based on a citation of O. E. Schulz, but they 

were not able to verify this record. There is no 

evidence to substantiate this record for Puerto 

Rico or the Virgin Islands nor has it been recently 

collected there. It is a species of the United States, 

not otherwise known to occur in the West Indies. 

Family 10. DIOSCOREACEAE Yam Family 

Dioscoreaceae R. Br., Prodr. 294. 1810, nom. conserv. 


Twining vines or less often erect herbs, with thickened starchy tubers. Leaves alternate or opposite, 

simple or palmately compound; petioles usually jointed at base and forming a stipule-like flange. Flowers 

small, commonly unisexual (the plant dioecious) or less often bisexual, actinomorphic, 3-merous, in 

axillary racemes, spikes or panicles; perianth of 6 nearly equal tepals, usually connate at base into a short 

tube; stamens 6, in two cycles, sometimes the inner cycle reduced to staminodes or vestigial, the filaments 

distinct, sometimes adnate to the perianth tube, the anthers opening by longitudinal slits; gynoecium 3pistillate, 

ovary inferior, trilocular, with axile placentation, the ovules 2 to many per locule, the styles 3, 

terminal, distinct, or basally connate. Fruit commonly a capsule, or less often a berry or a samara. A family 

of 5 or 6 genera and ca. 650 species, mostly tropical and subtropical. 

TYPE: Dioscorea L. 

References: Al-Shehbaz, I. A. & B. G. Schubert. 1989. The Dioscoreaceae in the southeastern United 

States. J. Arnold Arbor. 70: 57-95; Ayala Flores, F. 1998. Dioscoreaceae del Peru. Amazonian Natural 

Products, Iquitos Peru. 60 pages. Pedralli, G. 2004. Dioscoreaceas. In: A. Reis, Fl. Ilustrada Catarinense. 

Itajaí, Santa Catarina, Brazil, 84 pages. 


1. Fruit a trivalved capsule with 3, flattened locules …………...........................…… 1. Dioscorea 

1. Fruit a unilocular, flattened samara with a distal wing ……....................................…… 2. Rajania 

Dioscorea L, Sp. Pl. 1032. 1753. 

1. DIOSCOREA 

Dioecious, twining, herbaceous to woody vines, with large, single or clustered tubers; stems 

cylindrical, angled or winged, armed or unarmed. Leaves alternate or opposite, simple or palmately lobed, 

with arching parallel veins, long-petiolate, usually with aerial bulbils in axils. Flowers unisexual, 3merous, 

actinomorphic, in axillary spikes, racemes or panicles. Perianth minute; staminate flowers with 

6 stamens, all fertile or the inner cycle modified into staminodes; pistillate flowers bearing staminodes and 

an inferior ovary. Fruit a dry, chartaceous to leathery, 3-winged capsule; seeds numerous, flattened, 

winged. A tropical and subtropical genus of about 600 species, 10 of which are the source of edible yams


or tubers. 

LECTOTYPE: Dioscorea sativa L., designated by Britton & A. Brown, Ill. Fl. U.S. ed. 2, 1: 535. 1913. 

References: Burkill, H. M. 1985. The useful plants of west tropical Africa. Vol. 1. Royal Botanic 

Garden, Kew. Coursey, D. G. 1967. Yams. An account of the nature, origins, cultivation and utilization 

of the useful members of Dioscoreaceae. Longmans, London. Prain D. & I. H. Burkill. 1919. Dioscorea 

sativa. Bull. Misc. Inform. Kew 9: 339-375. Caddick, L.R., P. Wilkin, P.J. Rudall, T.A.J. Hedderson, & 

M.W. Chase. 2002. Yam reclassified: a recircumscription of Dioscoreaseae and Dioscoreales. Taxon 51: 

103-114. 

Key to the species of Dioscorea 

1. Stems mostly quadrangular or 4-winged (cylindrical at base in D. alata) ……..........................…….. 2 

2. Leaves opposite, entire …….............................................................................………. 1. D. alata 

2. Leaves alternate, 3- to 6-lobed ……..............................................................………. 7. D. trifida 

1. Stems cylindrical or trigonous, not winged …....................................................................…………… 3 

3. Leaves abaxially pilose along veins …......................................………………. 5. D. pilosiuscula 

3. Leaves glabrous …………..............................................................................................………. 4 

4. Stems acutely triangular, spiny …………...................................................….. 2. D. altissima 

4. Stems cylindrical …………...............................................................................…………… 5 

5. Stems usually spiny; leaves opposite ………..........................……….. 4. D. cayennensis 

5. Stems not spiny; leaves alternate …………….................................................………. 6 

6. Staminate flowers in lateral cincinni; fertile stamens 3; bulbils angulate, smooth; 

petiole slightly winged at base, not clasping the stem to form a pseudostipule 

………….............................................................................…… 6. D. polygonoides 

6. Staminate flowers along raceme axes, not grouped in cincinni; fertile stamens 6; 

bulbils rounded, verrucose; petiole winged at base, the wing clasping the stem to 

form a circular pseudostipule ……...........................................…… 3. D. bulbifera 

1. Dioscorea alata L., Sp. Pl. 1033. 1753. Type: 

India. (LINN-1184.2). 

Rajania flexuosa Bello, Anales Soc. Esp. Hist. 

Nat. 12: 124. 1883. Type: Puerto Rico. Bello, 

s. n. (destroyed). 

Fig. 17. A-E 

Herbaceous twining vine, 10-15 m long; 

stems slender, glabrous, 4-winged; old basal stems 

slightly woody, cylindrical, and spiny. Leaves 

mostly opposite; blades 6-26 (-30) × 4-18 cm, 

widely ovate, coriaceous, glabrous, with 5-7 

parallel veins, upper surface lustrous, with sunken 

veins, lower surface dull, with prominent veins, 

the apex acute or acuminate, sometime reflexed, 

the base cordate, the margins entire; petioles 4-12 

cm long, 4-winged, forming an auriculate sheath 

(pseudostipules) around the stem; bulbils axillary, 

elongated, pendent, to 15 cm long. Inflorescences 

axillary, unisexual, hanging; staminate 

inflorescences a panicle 5-15 cm long, with 

numerous lateral, flexuous, densely-flowered 

spikes; pistillate inflorescences simple, 9-25 cm 

long, in distant- to approximate-flowered spikes. 

Perianth of staminate flowers 1-1.5 mm long; 

perianth of pistillate flowers 2-2.8 mm long. 

Capsule 3-winged, 2-3 cm diam.; each locule 2seeded. 

General distribution: Native of southeast 

Asia, but widely cultivated and naturalized 

throughout the tropics. 


Islands: Previously widely cultivated for its edible 

tubers, now commonly naturalized in disturbed 

and secondary forests of moist areas. Recorded for 

Adjuntas, Aguada, Arecibo, Cataño, Cayey, 

Isabela, Maricao, Mayagüez, Quebradillas, Río 

Grande, Sabana Grande, San Juan, Toa Baja, Vega 

Baja, and Villalba; St. John. 

Common names: Puerto Rico: Ñame, Ñame 

de agua, Ñame blanco; Virgin Islands: Red yam, 

Water yam, and White yam. 


Adjuntas: Sintenis 4509 (US). Aguada: Bo.

84 


Fig. 17. A-E. Dioscorea alata. A. Axillary bulbil. B. Sterile branch, with detail of the stem. C. Infructescences, with detail of 

pseudostipules. D. Immature fruit. E. Mature fruit. F-G. Disocorea altissima. F. Mature stem, with adventitious roots. G. Leaves. 

From Acevedo-Rdgz., P. 2003, Bejucos y plantas trepadoras de Puerto Rico e Islas Vírgenes, Smithsonian Institution.


Guaniquilla, just SSE of Rd 441, km 1.5, Proctor 

47513 (SJ). Arecibo: Santa Ana, Goll 137 (US). 

Cataño: Goll 1066 (US). Isabela: Agr. Exp. Sta., 

Almeyda s.n. (US). Maricao: Bo. Indiera Fría, 

along track near Río Lajas below El Salto de Curet, 

Proctor & Padrón 45679 (SJ). Mayagüez: Bo. 

Montoso. Road 119, km 21.3, Breckon & 

Pteridology class 6282 (US). Quebradillas: 

Sargent B48 (US). Río Grande: Caribbean 

National Forest, entrance to El Verde area, 

Acevedo-Rdgz. & Siaca 7076 (US). San Juan: Río 

Piedras, Underwood & Griggs 358 (US). Toa 

Baja: Bo. Candelaria, 0.4-0.7 km WSW of Pico 

Nevárez, Proctor & Thomas 45574 (SJ). ST. JOHN: 

Bordeaux Mountain, Woodbury 461/6915 

(VINPS). 

2. Dioscorea altissima Lam., Encycl. 3: 233. 

1789. Lectotype: Martinique (?). Surian s.n., 

(P 00307100), designated by Pedralli in A. 

Reis, Fl. Ilustr. Catarinense. 12. 2004. 

Fig. 17. F, G 

Semi-woody, glabrous, twining vine, 10-15 m 

long; stems smooth, angled or triangular, to 1.5. 

cm wide, with recurved spines ca. 1 cm long. 

Leaves alternate or less often subopposite; blades 

8-16 × 7-16 cm, ovate or sub-orbicular, 

coriaceous, glabrous, with 7-9 parallel veins, the 

apex shortly acuminate to caudate, the base 

cordate, the margins slightly undulate; petioles 

slender, 6-15 cm long, sometimes minutely spiny, 

swollen at both ends, forming a pair of spiny 

pseudostipules at base; bulbils axillary, ellipsoid, 

elongated, pendent, scaly. Inflorescences axillary, 

of simple spikes; pistillate inflorescences longer 

than the staminate ones. Perianth of oblong 

segments. Capsule 3-winged, ca. 3 × 1.5 cm 


Rico, Lesser Antilles, northern South America to 

southern Brazil and Bolivia. 

Distribution in Puerto Rico: Apparently 

native, occasional in moist forests in low or 

medium elevations. Cayey, Juncos, Luquillo, 

Manatí, Río Grande, and Yabucoa. 

Common names: Puerto Rico: Ñame dunguey, 

Dunguey. 


Luquillo: Bo. Sabana, Caribbean National Forest, 

Axelrod et al. 10051 (US). Juncos: Monte Santo de 

León, Sintenis 2495 (US). Manatí: Coto Sur, 

Poblado Polvorín, Acevedo-Rdgz. 12296 (US). 

Yabucoa: Sintenis 5176 (US). 

After studying an image of the type of 

Dioscorea altissima, discovered just as this project 

was concluding, I got the impression that what has 

traditionally been called D. altissima, is a different 

species from the type of that name. Although the 

available descriptions of D. altissima portray the 

species as quite variable, the image of its type 

suggests it to be conspecific with D. cayennensis. 

Further studies are needed to clarify whether the 

type of D. altissima represents a distinctive species 

or the same as D. cayennensis. If the type of D. 

altissinma is indeed conspecific with D. 

cayennensis, the species that traditionally has been 

called D. altissima in the West Indies needs a 

different name. Alternatively, the name D. 

altissima could be proposed for conservation with 

a new type that would preserve its traditional 

usage. 

3. Dioscorea bulbifera L., Sp. Pl. 1033. 1753. 

Type: India. Hermann, Parad. Bat. t. 217. 

1698, designated by Milne-Redhead in 

Polhill, Fl. Trop. E. Afr. Dioscor. 10. 1975. 

Fig. 18. A-E 

Herbaceous, glabrous, twining vine, 8-10 m 

long; stems terete, smooth, unarmed. Leaves 

alternate; blades 9-12.5(17) × 5.5-11(15.5) cm, 

ovate, chartaceous, glabrous, with 9-11 parallel 

veins, the apex acuminate to caudate, the base 

cordate, the margins slightly undulate; petioles 

(4)12-15 cm long, winged and projecting as a pair 

of pseudostipules surrounding the stem at base; 

bulbils axillary, rounded, 5-6 cm wide, verrucose. 

Inflorescences axillary, simple fasciculate; 

staminate inflorescences usually paired, 9-15 cm 

long, the flowers in lateral, stipitate cincinni; 

pistillate inflorescences ca. 12 cm long, with 

solitary flowers. Flowers diminutive, sessile, 

white or pinkish tinged; staminate flowers with 

perianth ca. 1.2 mm long and 6 fertile stamens; 

pistillate flowers with perianth ca. 1.4 mm long 

and a hypanthium ca. 2 mm long. Capsules 3winged, 

ca. 2.5 cm long. 

General distribution: Native to the Old World 

tropics, but introduced and naturalized in the 

Neotropics. 

Distribution in Puerto Rico: Locally common 

in secondary moist forests of low and medium

86 


elevations. Collected in Arecibo, Río Grande, and 

Vega Alta. 

Common name: Puerto Rico: Gunda. 


Arecibo: 0.5 km S of Biáfara, Acevedo-Rdgz. & 

Siaca 11701 (US). Vega Alta: Bo. Sabana, NE of 

Regadera, Proctor & Concepción 45791 (SJ-2). 

4. Dioscorea cayennensis Lam., Encycl. Méth. 

Bot. 3: 233. 1789. Holotype. French Guiana; 

Cayenne. Stoupy s.n. (P-LAM-270403). 

Fig. 18. F-K 

Slightly woody vine, twining (toward the 

right), glabrous, attaining 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. 

General distribution: 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 is has become naturalized. 

Distribution in Puerto Rico: In areas of 

disturbed or secondary vegetation. Collected in 

Bayamón, Mayagüez, and San Sebastián. 

Common name: Puerto Rico: Ñame de 

Guinea. 


Without locality, Goll s.n. (US). Bayamón: 

Underwood & Griggs 1003 (US). Mayagüez: 

Kinman s.n. (US). San Sebastián: Sargent 370 

(US). 

5. Dioscorea pilosiuscula Bertero ex Spreng., 

Syst. Veg. 2: 152. 1825. Type: Hispaniola. 

Bertero s. n. (holotype: probably at TO). 

Fig. 18. L-P 

Slender, herbaceous twining vine, 2.5-8 m 

long; stems slender, wiry, glabrous or puberulent, 

cylindrical to slightly angled, unarmed. Leaves 

alternate; blades 4.5-12.5 × 2.5-6.5 cm, oblongovate 

to ovate, chartaceous, minutely pilose along 

veins below, with 5-7 parallel veins, the apex 

acuminate, the base cordate, the margins entire; 

petioles 1-4 cm long, slender, furrowed and 

swollen at both extremities; bulbils paired in leaf 

axils, ovoid, verrucose, 2.5-3 cm long. 

Inflorescences axillary; staminate inflorescences 

of 1-4 clustered spikes, 15-30 cm long; pistillate 

inflorescence a solitary spike, 9.5-30 cm long. 

Perianth of pistillate flowers 1.5-2 mm long. 

Capsule oblong, 3-winged, 1-2.5 cm long. Seeds 

ca. 8 mm long, with a basal wing. 

General distribution: Greater Antilles, Lesser 

Antilles, and South America 


Islands: Occasional in open disturbed areas. 

Reported for Arecibo, Isabela, Luquillo, Moca, 

Naguabo, Río Grande, San Juan, Utuado, and 

Vega Baja, Yabucoa; St. John, St. Thomas and 

Tortola. 

Common name: Puerto Rico: Dunguey. 


Isabela: Guajataca State Forest, Raz 184 (NY). 

Moca: Sargent 401 (US). Naguabo: Sierra de 

Naguabo, Sintenis 5353 (US). San Juan: Río 

Piedras, Stevenson 2466 (US). ST. JOHN: Bethany, 

N.L. Britton & Shafer 348 (NY, US); Coral Bay 

Quarter; Center Line Road, Acevedo-Rdgz. & 

Siaca 3991 (COL, F, JBSD, MICH, MO, NY, 

UPR, TEX, US, VINPS). ST. THOMAS: St. Peter, 

Eggers s.n. (US); E.G. Britton & Marble 1242 

(US). Tutu, N.L. Britton & E.G. Britton 451 (US). 

TORTOLA: Potwood Pond to West End, Shafer 1183 

(US). 

6. Dioscorea polygonoides Humb. & Bonpl. ex 

Willd., Sp. Pl. 4: 795. 1806. Type: Venezuela; 

Orinoco River. Humboldt & Bonpland s. n. 

(holotype: B-W; isotype: P-HBK, photo at 

US!). 

Fig. 19. A-I 

Herbaceous twining vine, 8 or more m long; 

stems glabrous, cylindrical, unarmed. Leaves 

alternate; blades (3)8-21 × (2)5-17 cm, ovate, 

chartaceous, glabrous, with 7-11 parallel veins, the 

apex acuminate, the base cordate, the margins


Fig. 18. A-E. Dioscorea bulbifera. A. Sterile branch, with axillary bulbils. B. Pseudostipules. C. Fertile branch. D. Pistillate 

inflorescence. E. Pistillate flower. F-K. Disocorea cayennensis. F. Fertile branch. G. Inflorescences. H. Staminate flower. I. 

Infructescences. J. Immature fruit. K. Stigmas and staminodia. L-P. Dioscorea pilosiuscula. L. Fertile branch, with axillary 

bulbils. M. Detail of infructescence. N. Immature fruit, with detail of the flower, showing stigmas and staminodia. O. Stigmas, 

side and top views. P. Fruit, side view and cross section. From Acevedo-Rdgz., P. 2003, Bejucos y plantas trepadoras de Puerto 

Rico e Islas Vírgenes, Smithsonian Institution.

88 


entire; petioles (1.5)5-8 cm long, slender, 

marginate at base but not extending as a 

pseudostipule; bulbils angled, to 5 cm wide, 

smooth. Inflorescences axillary, hanging, 

unisexual; staminate inflorescences solitary or 

paired, simple or less often branched, to 50 cm 

long, with flowers in lateral cincinni; pistillate 

inflorescence a solitary, simple spike, with solitary 

flowers along the axis. Flowers ca. 1 mm long, 

sessile, white or greenish; staminate flowers with 3 

fertile stamens; pistillate flowers hypanthium ca. 2 

mm long. Capsule elliptic, 3-winged, 2-3.6 cm 

long; seeds 2 per locule, orbicular, ca. 5 mm in 

diam. 

General distribution: Tropical America 

including the Antilles. 

Distribution in Puerto Rico: Common in 

moist secondary forests and disturbed habitats. 

Recorded from Arecibo, Bayamón, Caguas, 

Ceiba, Guaynabo, Luquillo, Maricao, Naguabo, 

Río Grande, Toa Baja, and Vega Baja. 

Common names: Puerto Rico: Gunda, 

Hícamo, Matagallina. 


Bayamón: Bo. Guaraguao Arriba, along Rd 879 

ca. 1 km WSW of Cerro La Peña, Proctor & 

Thomas 44550 (SJ). Caguas: Beatriz de Caguas, 

Goll 441 (US). Ceiba, N.L. Britton & Shafer 1544 

(US). Guaynabo: Bo. Sonadora, upper N slopes of 

Cerro Marquesa, Proctor 43537 (SJ, US). 

Luquillo: Sierra de Luquillo, Caribbean National 

Forest, Bo. Sabana, Proctor & Thomas 44646 (SJ). 

Naguabo: Sierra de Naguabo, Shafer 3569 (US). 

Río Grande: Sierra de Luquillo, Caribbean 

National Forest, Rd 191, km 9.9, Quebrada Juan 

Diego, Proctor 46230 (US), 46233 (US), 46234 

(SJ, US). Toa Baja: Bo. Candelaria, Proctor 45456 

(SJ). 

7. Dioscorea trifida L. f., Suppl. Pl. 427. 1782 

[“1781”]. Holotype: Surinam. Allamand s.n. 

Rajania L., Sp. Pl. 1032. 1753. 

2. RAJANIA 

(LINN-1184.1). 

Fig. 19. J-L 

Herbaceous twining vine; stems glabrous or 

puberulent, obtusely 4-angled, striate or 4-winged, 

unarmed. Leaves alternate; blades palmate or 5lobed, 

12-21 × 14-25 cm, ovate in outline, 

membranous, with 9-11 parallel veins, upper 

surface glabrous, with slightly prominent venation, 

lower surface puberulent, especially along the 

prominent veins, the lobes ovate-lanceolate, 

acuminate at apex, the base cordate, the margins 

entire or slightly undulate; petioles 5-19 cm long, 

4-winged, glabrous, pulvinate at base not forming 

a pseudostipule; bulbils not known. Inflorescences 

axillary, unisexual; staminate inflorescences 

paniculate, with flowers in lateral, sessile cymes; 

pistillate inflorescence racemose, with solitary 

flowers along the axis. Staminate flowers cream, 

1.5-6 mm long, with 6 fertile stamens; pistillate 

flowers 2.5-3.4 mm long, with pubescent 

hypanthium. Capsule oblong-elliptic, 2.5-3.4 cm 

long. 

General distribution: Apparently native to 

South America but now spread throughout the 

Antilles. 

Distribution in Puerto Rico: Cultivated for its 

edible tubers and apparently naturalized. Recorded 

from Moca. 

Common names: Puerto Rico: Mapuey, Ñame 

mapuey, Ñame morado, Ñame vino. 


Mayagüez: Agricultural Experimental Station, 

cultivated, coll. unknown s.n. (US). Moca: Sargent 

500 (US). 


Dioscorea floribunda M. Martens & Galeotti, 

D. friedrichsthalli R. Knuth, D. latifolia Benth., D. 

rotundata Poir., and D. sativa L. are cultivated in 

Puerto Rico, but they do not appear to have 

become naturalized. 

Dioecious, twining, herbaceous vines, with elongated, deep-rooted tubers; stems cylindrical, 

unarmed. Leaves alternate, simple, with arching parallel veins; petiolate. Inflorescences axillary. Flowers 

unisexual, the perianth minute, 6-merous, actinomorphic; staminate flowers with 6 fertile stamens, 

usually produced in lateral cymes along a racemiform thyrse; pistillate flowers produced in axillary 

racemes. Fruit a dry, indehiscent samara with a distal wing. An Antillean genus of about 25 species.


Fig. 19. A-I. Dioscorea polygonoides. A. Branch with staminate inflorescences. B. Cincinnus with staminate flowers, side view. 

C. Cincinnus with staminate flowers, front view. D. Staminate flower, top view. E. Stamens, staminate flower, side view. F. 

Branch with pistillate inflorescence. G. Pistillate flower. H. Stigmas. I. Infructescence. J-L. Disocorea trifida. J. Leaf. K. Fertile 

branch. L. Staminate flower, and detail of the stamens and pistillode. From Acevedo-Rdgz., P. 2003, Bejucos y plantas trepadoras 

de Puerto Rico e Islas Vírgenes, Smithsonian Institution.

90 


Rajania was merged into Dioscorea by Caddick et al. (2002) based on cladistic analyses of molecular and 

morphological characters that suggest that Rajania is nested within Dioscorea. They argue for the 

recognition of strictly monophyletic groups and thus do not recognize Rajania at the generic level. The 

present writer prefers to treat Rajania at the generic level as it seems to be monophyletic (within 

Dioscorea) and it is distinguishable by its indehiscent samaras. 

LECTOTYPE: Rajania hastata L., designated by Britton & Millspaugh, Bahamas Fl. 80. 1920. 

1. Rajania cordata L., Sp. Pl. 1032. 1753. 

Lectotype: West Indies. Plate 491 of the 

Boerhaave set of Plumier illustrations at the 

library of Rijksuniversiteit, Groningen, here 

designated. 

Rajania sintenisii Uline in Urban Symb. Antill. 3: 

281. 1902. Type: Puerto Rico. Sintenis 109 

(holotype: B, destroyed). 

Rajania cordata var. microcarpa Uline ex R. 

Knuth, Notizbl. Bot. Gart. Berlin-Dahlem. 7: 

219. 1917. Syntypes: Puerto Rico. Sintenis 

109b, 1384 (B, destroyed); St. Thomas, U.S. 

Virgin Islands. Eggers 184, 2893 (C?); Cuba. 

Wilson & León 11551 (NY!). 

Rajania venosa R. Knuth, Notizbl. Bot. Gart. 

Berlin-Dahlem. 7: 219. 1917. Type: Puerto 

Rico. P. Wilson 163 (holotype: NY, isotype: 

US!). 

Figs. 20. A-L; 60. I 

Herbaceous twining vine; stems glabrous, 

wiry, cylindrical. Leaves alternate; blades 4- 

11.5(20) × 2-6.5(12) cm, ovate, lanceolate or less 

frequently hastate, coriaceous, with 5 to 9 parallel 

veins, glabrous, the apex acute or acuminate, the 

base cordate or subtruncate, the margins entire; 

petioles 2-7 cm long, cylindrical, pulvinate at base, 

not forming a pseudostipule. Inflorescences 

axillary, unisexual; staminate inflorescences 

solitary or fasciculate, 4-25 cm long, with 1-3 

lateral, stipitate cymes per node; pistillate 

inflorescence racemose, 4-30 cm long, with 

solitary flowers along the axes. Flowers white or 

cream; staminate flowers 0.9-1.1 mm long, with 6 

fertile stamens; pistillate flowers 1-1.3 mm long, 

the hypanthium ca. 1.5 mm long. Samara (1.1)1.6- 

3.2 cm long, laterally flattened, usually reddish 

tinged. 

General distribution: Widely distributed 

throughout the Antilles. 


Islands: Common in humid or wet forests 

throughout the Cordillera Central and the karst 

limestone region. Recorded from Adjuntas, 

Aguadilla, Arecibo, Barranquitas, Bayamón, 

Cayey, Ciales, Cidra, Fajardo, Guayama, Isabela, 

Jayuya, Lares, Luquillo, Manatí, Maricao, 

Mayagüez, Naguabo, Peñuelas, Ponce, 

Quebradillas, Río Grande, Sabana Grande, 

Salinas, San Germán, Utuado, Vega Alta, 

Vieques, Villalba, and Yauco; St. Croix, St. 

Thomas. 

Common names: Puerto Rico: Ñame gulembo, 

Guáyaro, Bejuco de guaraguao. 


Adjuntas: Cordillera Central, Bo. Saltillo, 0.8 km 

due SW of Alto de la Bandera, Proctor 42125 (SJ). 

Aguadilla: Bo. Caimital Bajo, Acevedo-Rdgz. et 

al. 13437 (MAPR, US). Arecibo: Río Abajo State 

Forest, Acevedo-Rdgz. 11381 (US). Barranquitas: 

Bo. Barrancas, upper NE slopes and summit of 

Monte La Torrecilla, Proctor & Colón 50330 (SJ). 

Bayamón: Bo. Guaraguao Arriba, upper slopes 

and summit of Cerro La Peña, Proctor & Díaz 

41862 (SJ); Hato Tejas, Goll 238 (US). Cayey: 

Carite Forest Reserve, Cerro La Santa, Acevedo- 

Rdgz. 7927 (US). Ciales: along Road 149, 

Acevedo-Rdgz. & Alvarez 2977 (US). Cidra: Bo. 

Salto, uppermost slopes and summit of El Peñon, 

Proctor et al. 47634 (SJ). Fajardo: N.L. Britton & 

Shafer 1732 (US). Guayama: Carite Reserve, 

vicinity of Guavate Recreational Area, Wasshausen 

1316 (US). Isabela: Reserva Forestal Bosque 

Guajataca, La Caballa Trail, Acevedo-Rdgz. & 

Siaca 11735b (US). Jayuya: Bo. Veguitas, upper 

NW slope of Piedra Blanca, Proctor & Haneke 

41031 (SJ). Lares: Underwood & Griggs 40 (US). 

Manatí: Bo. Tierras Nuevas Saliente, area just S of 

Laguna Tortuguero, Proctor 42698 (SJ); Mango 

Hill, at end of road from Adventist camp, Axelrod 

& Nir 11069 (US). Maricao: Bo. Indiera Fría, 

mountain ridge E of Río Postrero, just E of State 

Forest boundary, Proctor et al. 45221 (SJ). 

Mayagüez: Cerro de Las Mesas, N.L. Britton & 

Hess 2693 (US); Naguabo: Sierra de Naguabo, El 

Duque, NW side and summit, Shafer 3655 (US). 

Peñuelas: Sintenis 4526 (US). Ponce: Toro Negro 

State Forest, along headwaters of Río Inabón


Fig. 20. A-L. Rajania cordata. A. Staminate inflorescence. B. Fertile branch. C. Branch with racemose inflorescence. D. Distal 

portion of inflorescence. E. Detail of cincinnus. F. Staminate flower, top and side views. G. Branch with pistillate inflorescence. 

H. Pistillate flower, not fertilized and fertilized. I. Stigmata, top and side views. J-K. Infructescence. L. Samaroid fruit. From 

Acevedo-Rdgz., P. 2003, Bejucos y plantas trepadoras de Puerto Rico e Islas Vírgenes, Smithsonian Institution.

92 


above high falls, Proctor 40073 (SJ). Quebradillas: 

Guajataca Forest, Acevedo-Rdgz. & Chinea 5226 

(US). Río Grande: Caribbean National Forest, El 

Yunque, Acevedo-Rdgz. 7101 (UPRRP, US). 

Sabana Grande: Maricao State Forest, Bo. 

Santana, S slopes of Monte Padrón, Proctor & 

Padrón 41807 (SJ). Salinas: Bo. Lapa, summit of 

E peak, Las Tetas de Cayey, Proctor & McKenzie 

44073 (SJ). San Germán: Maricao State Forest, 

vicinity of Campamento Buena Vista, Proctor & 

Padrón 41906 (SJ). San Juan: Río Piedras, 

Stevenson 158 (US). Utuado: Bo. Don Alonso, 

Acevedo-Rdgz. 13412 (US). Vieques: Isabel 

Segunda to Martineau, Shafer 2631 (US). Vega 

Alta: Espinosa, Stevenson 3113 (US). Vega Baja: 

Goll 1033 (US). Villalba: Toro Negro Forest, crest 

of Cordillera Central SW of Cerro Doña Juana, 

Webster et al. 8738 (US). Yauco: Susúa Forest 

Reserve, Acevedo-Rdgz. 11406 (US). ST. THOMAS: 

Crown Mountain, Acevedo-Rdgz. 11208 (US). 

Family 11. BURMANNIACEAE Burmannia Family 

Burmanniaceae Blume, Enum. Pl. Javae 1: 27. 1827, nom. conserv. 


Saprophytic or sometimes autotrophic small herbs. Rhizomes often present, these covered with 

minute scale-like leaves; roots filiform, lacking root-hairs. Stems erect, usually unbranched, white or 

variously colored. Leaves alternate, sessile, simple, entire, in saprophytic species minute and scale-like, 

but in autotrophic species sometimes rather large and often rosulate. Inflorescence a terminal, 1- to manyflowered 

cyme, this open and few-flowered to compact or dense, usually bifurcate. Pedicels usually 

distinct. Flowers bisexual, sympetalous, usually actinomorphic, white or variously colored, consisting of 

a basal tubular part and 6 tepals, these arranged in 2 whorls; floral tube sometimes winged or ribbed. 

Stamens 3 or 6, erect or pendent, inserted in the floral tube just below and opposite the inner whorl of 

tepals, the filaments mostly very short; anthers transversely or longitudinally dehiscent. Style 3-branched 

at the apex, each branch opposite an outer tepal and bearing a variously-shaped stigma; nectaries often 

present. Ovules many, very small. Fruit a capsule, transversely or longitudinally dehiscent by slits or 

valves, or sometimes opening irregularly by withering of the fruit wall. Seeds many, very small. A 

pantropical family of 15 genera and ca. 125 species, occurring also in subtropical to temperate areas such 

as southern China, Japan, northern Australia, Tasmania, and New Zealand. The Neotropical area is 

credited with 10 genera, 54 species, and three subspecies. 

TYPE: Burmannia L. 

References: Jonker, E. P. 1938. A monograph of the Burmanniaceae. Utrecht. Maas, P. J. M. et al. 

1986. Burmanniaceae. Fl. Neotrop. Monogr. 42: 1-189. Maas, P. J. M. & H. Maas. 1990. Flora vascular 

de La Española: Burmanniaceae. Moscosoa 6: 134-139. 


1. Non saprophytic herbs with small or minute green leaves; flowers clustered in a small dense head 

........................................................................................................................................... 2. Burmannia 

1. Saprophytic herbs without chlorophyll; flowers solitary, loosely cymose, or scattered, not in dense 

heads ……...................................................................................................................................…… 2 

2. Flowers usually purple, funnel-shaped; tepals persistent in fruit …………. 1. Apteria 

2. Flowers white at anthesis, salverform, tepals deciduous, soon falling, leaving a naked floral tube 

………............................................................................................................…. 3. Gymnosiphon


1. APTERIA 

Apteria Nutall, J. Acad. Nat. Sci. Philadelphia (7): 64. 1834. 

Erect, saprophytic herb with small, slightly tuberous, cylindrical rhizome, densely covered with 

minute, ovate-triangular imbricate scales and filiform roots. Stems with one terminal flower or few to 

several, scattered, axillary, erect or nodding flowers, these funnel- to salver-shaped. Inner tepals on a 

crescent-shaped outgrowth of the floral tube, each filament abaxially bearing a 2-lobed wing; anthers 

dehiscing transversely. Placenta lacking glands. Floral tube completely persistent in fruit. A unispecific 

genus, with a single polymorphic species widely distributed throughout the Neotropics. 

TYPE: Apteria aphylla (Nutt.) Barnhart ex Small (≡ Lobelia aphylla Nutt.). 

1. Apteria aphylla (Nutt.) Barnhart ex Small, Fl. 

s.e. U. S. 309. 1903; Lobelia aphylla Nutt., 

Amer. J. Sci. Arts 5: 297. 1822; Apteria 

setacea Nutt., J. Acad. Nat. Sci. Philadelphia 

7(1): 64 1834. nom. illeg. Type: United States; 

eastern Florida. Ware s. n. (holotype: BM). 

Apteria hymenanthera Miq., Stirp. Surinam. 

Select. 216. 1850; Apteria aphylla var. 

hymenanthera (Miq.) Jonker, Monogr. Burm. 

208. 1938 and in Pulle, Fl. Suriname 1(1): 

186. 1938. Type: Surinam. Hostmann 959 

(holotype: U; isotypes: BM, DS, G, GH, K, 

LE, OXF, P, W). 

Fig. 21. F-I 

Stems filiform, 4-25 cm tall or more, simple or 

branched, purplish except toward base. Leaves 

few, purplish to white, more or less ovate, 0.5-3.5 

cm long. Flowers few, widely separated, with 

filiform pedicels up to 20 mm long; floral tube 4- 

16 mm long; tepals narrowly ovate to narrowly 

Burmannia L., Sp. Pl. 287. 1753. 

2. BURMANNIA 

oblong, 1.2-5.3 mm long; usually purple (rarely 

white) with stripes inside the throat. Stamens 

yellow, inserted 2-4.5 mm below the insertion of 

the inner tepals; style 2.4-4.5 mm long, crowned 

by a persistent floral tube; seeds brownish, more or 

less ellipsoid, mostly 0.3-0.4 mm long or 

sometimes longer. 


States, southern Mexico, Central America, 

Greater and Lesser Antilles, Trinidad, and South 

America as far as Paraguay and southern Brazil. 

Distribution in Puerto Rico: In moist, deeplyshaded 

humus at chiefly middle elevations (300- 

800 m). It is probably more common than its few 

records seem to indicate. Recorded from Bayamón 

(Cerro La Peña), Manatí, Maricao, Naguabo, Río 

Grande, and Yabucoa. 


Maricao: Indiera Fría, N.L. Britton et al. 4472 

(US). 

Erect autotrophic (rarely saprophytic), terrestrial (rarely epiphytic) herbs without rhizomes but with 

delicately filiform roots. Stems simple or branched. Leaves minute and scale-like to rather large, 

sometimes basal and rosulate. Inflorescence a terminal, bifurcate, open to capitate cincinnus, or else 

sometimes the plant with a solitary terminal flower. Flowers erect (rarely nodding), tubular to salverform, 

sessile to shortly pedicellate. Outer tepals entire, larger than the inner ones, all inserted at the same level; 

inner tepals sometimes incurved over the stamens. Floral tube cylindrical to trigonous, wingless to broadly 

3-winged. Anthers 3, sessile, inserted just below the inner tepals. Style 3-branched at the apex, each branch 

with a bilabiate (or sometimes a funnel-shaped) stigma. Ovary 3-locular, trigonous, with axile 

placentation; septal nectaries 3, sometimes present, opening at base of the floral tube. Capsule crowned by 

the persistent perianth, this ultimately dehiscent. Seeds yellowish to greenish brown, more or less 

ellipsoid, 0.2-1 mm long. A pantropical genus with 60 species, of which 30 occur in Asia, 11 in Africa, and 

19 in the Neotropics, extending from southeastern United States to Argentina, Paraguay, and southeastern 

Brazil. 

LECTOTYPE: Burmannia disticha L., designated by Britton & A. Brown, Ill. Fl. N. U.S., ed. 2, 1: 547. 

1913.

94 


Fig. 21. A-E. Burmannia capitata. A. Habit. B. Inflorescence. C. Flower and bract. D. L.s. flower showing axial placentation 

and ovules, stamens, and style. E. Seed. F-I. Apteria aphylla. F. Habit. G. Flower and bracts. H. L.s. flower showing stamens and 

style. I. Seed. (A-E, from Proctor 39469; F-I, from Proctor 42775). 

1. Burmannia capitata (J. F. Gmel.) Mart., Nov. 

Gen. Sp. Pl. 1: 12. 1823; Vogelia capitata J. F. 

Gmel., Syst. Nat. 2: 107. 1791; Tripterella 

capitata ( J. F. Gmel.) Michx., Fl. Bor.-Amer. 

1: 19. 1803. Type: United States; North 

Carolina. Walter s. n. (BM ?). 

Fig. 21. A-E 

Delicate, usually unbranched, erect herb 

mostly 4-34 cm tall (rarely more), stems yellowish 

to green. Leaves subulate to narrowly ovate, 1.6-9 

mm long (rarely longer), up to 2 mm wide, usually 

few and inconspicuous, sometimes apparently 

lacking. Inflorescence 1- to many-flowered, 

usually head-like consisting of 2 densely 

contracted cincinni (rarely the later upright and 

extended). Flowers sessile or subsessile, tubular, 

white to yellowish white, mostly 2.5-6 mm long; 

outer tepals deltate; inner tepals elliptic, sometimes 

lacking; floral tube mostly 0.7-1.5 mm long, the 

wings reduced to ribs; style 1-1.8 mm long. Ovary 

obovoid, ellipsoid, or globose, 1.3-2.5 mm long. 

Capsules white to yellow, 1.3-2.9 mm long,


dehiscent by transverse slits; seeds 0.2-0.4 mm 

long. 


States, southern Mexico, Central America, 

Greater Antilles, Trinidad and northern South 

America, eastern South America south to 

Paraguay and southeastern Brazil. 

Distribution in Puerto Rico: Grows on moist 

white silica sand at or near sea level, but has also 

3. GYMNOSIPHON 

Gymnosiphon Blume, Enum. Pl. Javae 1: 29. 1827. 

been found on roadside banks at lower middle 

elevations (to 200 m), very rare but also easily 

overlooked. Recorded from Las Marias, Manatí, 

and Vega Baja. 


Manatí: SE side of Laguna Tortuguero, Strong et 

al. 425 (US). Vega Baja: Bo. Algarrobo, just S of 

Laguna Tortuguero, Proctor & Thomas 45099 

(US); Woodbury s.n. (US). 

Erect saprophytic herbs with small, cylindrical, slightly tuberous rhizomes, these densely covered 

with minute, narrowly ovate, imbricate scales and also bearing filiform roots. Inflorescence a terminal 

double cincinnus, each branch 1- to 10-flowered, the base of the inflorescence bearing two stem-clasping 

bracts. Flowers white, erect, salver-shaped, pedicellate; outer tepals 3-lobed; inner tepals very small, 

inserted in the floral tube below the insertion of the outer tepals. Stamens sessile, the connective without 

appendages; anther-sacs dehiscing horizontally. Stigmas mostly with long filiform appendages. Each 

placenta with two globose apical glands. Upper part of the floral tube quickly deciduous from the 

developing fruit. A pantropical genus of 24 species, of which 14 are found in the Neotropics, 7 occur in 

tropical Asia, and 3 in tropical Africa. 

TYPE: Gymnosiphon aphyllus Blume. 

Key to species of Gymnosiphon 

1. Capsules broadly obovoid to ellipsoid or globose, 3-5.5 mm long including the persistent part of the 

floral tube; pedicels up to 5 mm long ……...................................................................….. 1. G. niveus 

1. Capsules mostly very broadly obovoid, 2.5-4.2 mm long including the persistent part of the floral tube; 

pedicels up to 1.5 mm long ………….................................................………….. 2. G. sphaerocarpus 

1. Gymnosiphon niveus (Griseb.) Urb., Symb. 

Antill. 3: 444. 1903; Ptychomeria nivea 

Griseb., Cat. Pl. Cub. 257. 1866. Type: Cuba. 

Wright 3285 (lectotype: GOET; isotypes: 

BM, G, GH, MO, NY, S), designated by Maas 

et al., Fl. Neotrop. Monogr. 41: 126. 1986. 

Gymnosiphon germaini Urb., Symb. Antill. 3: 

444. 1903. Type: Guadeloupe. Germain, s.n. 

(holotype: B, destroyed; isotype: G). 

Gymnosiphon portoricensis Urb., Symb. Antill. 

3: 445. 1903; Ptychomeria portoricensis 

(Urb.) Schltr., Repert. Spec. Nov. Regni Veg. 

17: 257. 1921. Syntypes: Puerto Rico. 

Sintenis 5170 (B, G, GH, GOET, K, M, P, S, 

U); Sintenis 5259; Sintenis 5707. 

Stems filiform, 4-28 cm tall, white, simple or 

rarely branched. Leaves few, ovate, 0.7-2 mm 

long. Inflorescence of 2 cincinni, 0.3-6 cm long, 1- 

to 9-flowered; pedicels mostly 1-4.5 mm long. 

Flowers white, 5-7 mm long; central lobe of outer 

tepals the largest, ovate, up to 1.1 mm long, the 

outer lobes smaller; inner tepals entire, much 

smaller than the outer ones. Stamens inserted 

below the insertion of the inner tepals. Style 

(including branches and stigmas) up to 2.6 mm 

long, the stigmas with filiform appendages. Ovary 

obconic, 1.2-2.1 mm long. Capsules ellipsoid, up 

to 3.7 mm long (including the persistent part of the 

floral tube, 3-5.5 mm long); seeds brownish 

yellow to yellowish white, irregularly and broadly 

ellipsoid. 

General distribution: Greater and Lesser 

Antilles. 


from the collections of Sintenis (5170, 5259, 

5707), dating from late 19th century, not recently 

collected. Reported for Aguada, Coamo, and

96 


Yabucoa. 

Note: Although Jonker cited Sintenis 5170 as 

the type for Gymnosiphon portoricensis Urb. in 

his monograph of Burmanniaceae, he failed to 

lectotypify this name because he did not select a 

particular herbarium. 


Yabucoa: Sintenis 5170 (photo at US ex B). 

2. Gymnosiphon sphaerocarpus Urb., Symb. 

Antill. 3: 442. 1903; Ptychomeria 

sphaerocarpa (Urb.) Schltr., Repert. Spec. 

Nov. Regni Veg. 17: 257. 1921. Syntypes: 

Guadeloupe. L’Herminier s. n. (B, G); Duss 

3942 (NY-2). 

Herb 6-16 cm tall with white, branched or 

unbranched stems, the small rhizomes beset with 

narrowly ovate scales 1.7-2 mm long. Leaves oval 

to broadly ovate 0.8-2.1 mm long. Inflorescence 

bifurcate, of paired cincinni, each branch 0.5-9 cm 

DOUBTFUL GENUS 

long and with 2-10 flowers; rarely each cincinnus 

branched again; pedicels mostly 0.3-1.5 mm long, 

sometimes longer. Flowers pale yellow in bud, 

white at anthesis, 3.2-5 mm long with floral tube 

3.2-3.5 mm long; outer tepals 3-lobed, the middle 

lobe larger than the others; inner tepals entire, very 

small. Stamens not described. Style (including 

branches and stigmas) 1.4-1.8 mm long; stigmas 

almost sessile, horseshoe-shaped, without 

appendages. Ovary obconic, 0.7-1.5 mm long. 

Capsules broadly obovoid, to 2.8 mm long. Seeds 

0.2-0.3 mm long. 


Antilles. 

Distribution in Puerto Rico: A tiny saprophytic 

plant, easily overlooked, known from a single 

record (Britton et al. 4471) from Indiera Fría, 

Maricao. 


Maricao: Indiera Fría, Britton et al. 4471 (F, NY, 

US). 

Cymbocarpa refracta Miers was cited by Liogier and Martorell (2000) as occurring in wet forests 

in the Luquillo mountains, however, no specimen have been located to confirm this report. 

Family 12. HYPOXIDACEAE Star Grass Family 

Hypoxidaceae R. Br. in Flinders, Voy. Terra Austr. 2: 576. 1814, nom. conserv. 


Perennial herbs with tuberous rhizomes or corms. Leaves mostly radical, prominently veined, often 

hairy. Flowers bisexual, actinomorphic. Perianth-tube very short or absent or consolidated into a long 

beak on top of the ovary; segments 6, spreading, more or less uniform. Stamens 6 or 3, inserted at the bases 

of the segments; anthers 2-locular, opening lengthwise. Ovary inferior, 3-locular; styles 1 or 3; ovules 

numerous in each locule, in 2 series on axile placentas. Fruit a capsule or fleshy and indehiscent. Seeds 

usually small, black; embryo enclosed by copious endosperm. A family of 7 to 10 genera and ca. 150 

species, mainly in the southern hemisphere. 

TYPE: Hypoxis L. 

Reference: Brackett, A. 1923. I. Revision of the American species of Hypoxis. Rhodora 25: 120-147. 


1. Ovary extended to form a long beak below the perianth; leaves plaited; fruit berry-like ….. 1. Curculigo 

1. Ovary not beaked; leaves flat, not plaited; fruit a capsule ……….................................…… 2. Hypoxis


1. CURCULIGO 

Curculigo Gaertn., Fruct. Sem. Pl. 1: 63. 1788. 

Perennial herbs with a vertical short, cylindrical or corm-like rhizome, bearing contractile roots, the 

stems and leaves with bifurcate or stellate hairs. Leaves tristichous, few, plicate-veined, elongate, linear. 

Inflorescences solitary or umbellate, short. Flowers borne among leaves, short-pedicelled, villose; 

perianth 6-parted, borne on a tube-like extension of the ovary, perianth segments unequal; stamens 6, the 

anthers linear; ovary 3-celled, with 2 to many ovules per cell, the style short, with 3 stigmas. Fruit slightly 

fleshy, indehiscent; seeds subglobose, black. A tropical genus of ten species. 

TYPE: Curculigo orchioides Gaertn. 

Key to the species Curculigo 

1. Flowers numerous, capitate; leaves elliptic-lanceolate, to 1.8 m long, 6-10 cm wide … 1. C. capitulata 

1. Flowers 1-4, each on a solitary peduncle; leaves linear, to 30 cm long, 1.5 cm wide 

................................................................................................................................... 2. C. scorzonerifolia 

1. Curculigo capitulata (Lour.) Kuntze, Revis. 

Gen. Pl. 2: 703. 1891; Leucojum capitulatum 

Lour., Fl. Cochinch. 1: 199. 1790. Type: 

“Cochinchina”. Loureiro, s.n. (probably at 

BM ). 

Curculigo recurvata Dryand. in W. T. Aiton, Hort. 

Kew. ed. 2, 2: 253. 1811. Type: Unknown, a 

plant brought into cultivation at Kew in 1805, 

originally collected from Bangladesh by W. 

Roxburgh. 

Fig. 61. A 

Plants acaulescent, commonly forming dense 

clumps; rhizome fleshy, erect, cylindrical. Leaves 

forming a rosette, with a pseudopetiole 30-60 cm 

long; blades elliptic-lanceolate, narrowed at both 

ends, 30-90 × 5-15 cm, abaxially pubescent, 

otherwise glabrous. Inflorescence capitate, erect, 

becoming curved or recurved in fruit; peduncles 

short to elongate, pubescent; bracts narrowly 

lanceolate; flowers with yellow perianth, 2-2.5 cm 

long. Fruit fleshy, globose or ellipsoid, whitish 

turning black with age. 

General distribution: Native to India, Sri 

Lanka, S. China, Malaysia and tropical Africa, 

now widespread in tropical regions through 

cultivation, sometimes adventive. 

Distribution in Puerto Rico: Commonly 

2. HYPOXIS 

Hypoxis L., Syst. Nat. ed. 10, 972, 986, 1366. 1759. 

cultivated in Puerto Rico. Reported as adventive in 

Arecibo and San Juan. 


Photo: Acevedo-Rdgz. s.n. 

2. Curculigo scorzonerifolia (Lam.) Baker, J. 

Linn. Soc., Bot. 17: 124. 1878; Hypoxis 

scorzonerifolia Lam., Encycl. 3: 183. 1789. 

Lectotype: West Indies. Plumier, Descr. Pl. 

Amér. t. 108, f. 2. 1693, designated by R.A. 


Fig. 22. A-H 

Underground stem fleshy, erect, cylindrical. 

Leaves few, linear; blades 10-30 × 0.5-1.5 cm, 

pilose, gradually or abruptly narrowed at the base. 

Flowers borne singly on short axillary peduncles; 

perianth tube 2 cm long, pilose, the lobes bright 

yellow inside, 1.1-1.2 cm long. 

General distribution: Tropical America. 


adventive in San Juan and Mayagüez. 

Common name: Puerto Rico: Gorgojo. 


Mayagüez: Cerro Las Mesas, Proctor & McKenzie 

43778 (US). 

Herbs with corms. Leaves linear, grass-like. Scapes slender, 1- to few-flowered; perianth 6-parted,

98 


Fig. 22. A-H. Curculigo scorzonerifolia. A. Habit. B. Apex of leaf blade. C. Flower. D. Style and stigmata. E. Stamen. F. Fruit 

showing seeds. G. Fruit. H. Seed. (from Proctor 43778).


the segments free, withering and persistent; stamens inserted on perianth, the filaments short, the anthers 

erect; ovary 3-celled, not extended or narrowed at apex, with numerous ovules, the style short, the stigmas 

3. Capsule thin-walled, elongate, circumcissile below the apex; seeds subglobose, tuberculate, black. A 

genus of about 100 species, occurring in the Americas, Africa, and Eastern Asia, through the Malay 

Peninsula to Australia. 

LECTOTYPE: Hypoxis hirsuta (L.) Coville, (≡ Ornitogalum hirsutum L.), designated by Britton & A. 

Brown, Ill. Fl. N. U.S. ed. 2, 1: 534. 1913. 

Key to species of Hypoxis 

1. Peduncles 2- or more-flowered; leaves more than 2 mm broad…………..……………. 1. H. decumbens 

1. Peduncles 1-flowered; leaves mostly less than 2 mm broad ………................…………. 2. H. wrightii 

Fig. 23. A-E. Hypoxis decumbens. A. Habit. B. Flower, lateral and top views. C. Stamen, lateral and frontal views. D. Capsule. 

E. Seed. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78.

100 


1. Hypoxis decumbens L., Syst. Nat. ed. 10, 986. 

1759. Lectotype: Jamaica. Browne s.n. (LINN 

427.2 ), designated by R.A. Howard, J. Arnold 

Arbor. 60: 300. 1979. 

Fig. 23. A-E 

Plants 10-30 cm tall; underground stems 

upright, cylindrical, 1-2 cm thick. Leaves linear, 

10-45 × 0.3-1 cm, flattened, sparsely pilose to 

glabrous. Inflorescence slender, ascending or 

recurving, 4-20 cm long, pilose near apex, 1- to 4flowered. 

Perianth 9-14 mm long, greenish, pilose, 

the inner perianth segments yellow, 4-9 mm long. 

Capsule cylindrical, ellipsoid or club-shaped, 7-12 

mm long, tipped with the persistent, withered 

perianth. Seeds 0.8-1.2 mm in diam., black. 


America, Greater Antilles, Lesser Antilles, and 

South America. 


Islands: Occasional in moist grounds at middle 

and high elevations (300-1200 m). Reported from 

Adjuntas, Aibonito, Arecibo, Bayamón, Dorado, 

Florida, Hatillo, Jayuya, Luquillo, Manatí, 

Maricao, Mayagüez, Naguabo, Orocovis, Río 

Grande, San Juan, San Sebastián, Villalba, 

Yabucoa, and Yauco; St. John, Tortola, and Virgin 

Gorda. 

Common name: Puerto Rico: Coquí. 


Adjuntas: Liogier 9981 (US). Arecibo: Río Abajo 

State Forest, Acevedo-Rdgz. 10561 (US). Hatillo: 

Bo. Bayaney, Proctor et al. 41878 (US). Luquillo: 

Caribbean National Forest, El Yunque Area, 

Acevedo-Rdgz. & Siaca 6227 (US). Manatí: SE 

side of Laguna Tortuguero, Strong et al. 421 (US). 

Maricao: Sintenis 488 (US). Mayagüez: Cerro Las 

Family 13. IRIDACEAE Iris Family 

Iridaceae Juss., Gen. Pl. 57. 1789, nom. conserv. 


Mesas, N.L. Britton & Hess 2698 (US). Naguabo: 

Loma La Mina, Shafer 3239 (US). Río Grande: 

Luquillo Mountains, Acevedo-Rdgz. & Siaca 6227 

(NY, UPRRP, US). San Juan: Río Piedras, 

Stevenson 44 (US). San Sebastián: Sargent 409 

(US). ST. JOHN: Bethania to Rosenberg, N.L. 

Britton & Shafer 279 (US). Bordeaux Mt., 

Woodbury 69a/6556 (VINPS). TORTOLA. Road 


(US). VIRGIN GORDA: Wet places on mountains, 

Fishlock 113 (US). 

2. Hypoxis wrightii (Baker) Brackett, Rhodora 

25: 140. 1923; Hypoxis juncea var. wrightii 

Baker, J. Linn. Soc., Bot. 17: 106. 1878. 

Type: Cuba. Wright 239 (isotype: GOET). 

Underground, tuberous stem cylindrical or 

ovoid, 5-15 mm long, up to 12 mm thick, covered 

with the fibrous remains of old leaf sheaths. 

Leaves 5-18 (-25) cm × 1.5-2 (-3) mm, thinly 

cottony-hairy. Peduncles 3-5 cm long in flower, 

longer in fruit, very slender. Perianth ca. 8 mm 

across, light yellow, the outer members hispid 

externally, the inner hispid in midline only; ovary 

pubescent. Capsule densely pilose, ellipsoidobovate, 

4-6 mm long. Seeds oblong, minutely 

muriculate, 1 mm long, glossy, black. 

General distribution: Southern United States 

and the Greater Antilles. 

Distribution in Puerto Rico: Uncommon, 

reported for Cataño, Manatí, Dorado, and Vega 

Baja. 

Selected specimens studied: PUERTO RICO: 

Cataño, Sintenis 1067 (US). Manatí / Vega Baja, 

Underwood & Griggs 956 (US). 

Chiefly perennial herbs from rhizomes, bulbs or corms. Leaves equitant, entire. Inflorescences 

axillary, scapose, racemose, or cymose; flowers bisexual, regular or irregular, solitary or in clusters arising 

from spathe-like bracts; perianth of 6 segments usually fused at the base, short-lived and fugacious or 

persistent; stamens 3, inserted on the perianth opposite the outer ones, the filaments separate or partially 

united; ovary inferior, 3-locular, with numerous ovules; styles distinct, entire or divided, sometimes 

flattened and petal-like. Fruit a loculicidally 3-valved capsule; seeds spherical, flattened, or angled. A


cosmopolitan family of 60 genera and ca. 800 species widely distributed throughout the world in both 

temperate and tropical regions. 

TYPE: Iris L. 


1. Perianth actinomorphic or nearly so ……..................................................................................…….. 2 

2. Robust plants with stout rhizomes and aerial stems to 1 m tall or more; perianth orange with purple 

spots, 3-5 cm wide ……….................................................................….. 1. Belamcanda 

2. Diminutive tufted plants less than 15 cm tall; flowers pale yellowish, not over 5 mm 

wide…………….............................................................................................… 5. Sisyrinchium 

1. Perianth more or less zygomorphic ……...............................................................................……….. 3 

3. Stem a rhizome; flowers produced on a flattened, leaf like scape; perianth showy, the outer 

segments white ………....................................................................................….. 4. Neomarica 

3. Stem a subterranean bulb or corm …............................................................................……….. 4 

4. Perianth-segments united at base to form a tube ……………....................................……. 5 

5. Spathes emarginate; perianth tube straight or nearly so; perianth lobes bright orange-red 

………............................................................................................… 2. Crocosmia 

5. Spathes lanceolate, acute; perianth-tube curved; Perianth lobes light pink to pale orange 

......................................................................................................... [Gladiolus] 

4. Perianth-segments free or nearly so …………..............................................................….. 6 

6. Style-branches undivided, subulate; perianth white; capsules enclosed by a spathe-like 

bract ………….........................................................................…… 3. Eleutherine 

6. Style-branches divided; perianth yellow; capsules exserted on long pedicels 

..................................................................................................................... 6. Trimezia 

1. BELAMCANDA 

Belamcanda Adans., Fam. Pl. 2: 60, 524. 1763, nom. conserv. 

Perennial caulescent herbs with short, stout rhizomes and alternate equitant leaves. Flowers in 

branched, terminal, bracteate clusters; perianth tube short, the segments equal, with mottled colors; 

stamens 3, the filaments free but adnate to base of alternate perianth segments; ovary 4-locular, the style 

slender, with flattened and emarginate branches. Capsules obovoid, loculicidally 3-valved, the valves 

becoming recurved and deciduous; central column persistent, bearing temporarily attached, globose, 

shiny black seeds. An eastern Asiatic genus of two species, one of them widely cultivated and often 

becoming naturalized in tropical countries. 

TYPE: Belamcanda chinensis (L.) A. De Candolle 

1. Belamcanda chinensis (L.) DC. in Redouté, 

Liliac. 3: t. 121. 1805; Ixia chinensis L., Sp. 

Pl. 36. 1753; Belamcanda punctata Moench., 

Methodus 529. 1794, nom. illeg. Lectotype: 

“India”. (LINN 58. 34), designated by R.A. 


Stems stout, erect, up to 1 m tall. Leaves 2ranked, 

with overlapping bases, lengthwise 

folded, ascending, linear-lanceolate, 15-50 cm 

long. Inflorescences to 50 cm long, with bracts 

similar to leaves. Flowers several in each cluster; 

perianth 3-5 cm long, the segments nearly alike, 

oblong, obtuse at apex, narrowed at base, bright 

orange-yellow mottled with crimson or purple 

spots. Capsules 2-2.6 cm long, dehiscent; seeds 

globose, black. 

General distribution: Native of tropical Asia, 

now widely distributed in warm countries as an 

escape from cultivation. 

Distribution in Puerto Rico: Uncommon in 

open, disturbed places between 200-800 m. 

Recorded from Adjuntas, Bayamón, Juncos, and 

Yabucoa. 

Common name: Puerto Rico: Maravilla. 

Selected specimens examined: PUERTO RICO:

102 


Adjuntas: Garzas, Sintenis 4761 (US). Bayamón: 

Stevenson 3045 (US); Underwood & Griggs 853 

2. CROCOSMIA 

Crocosmia Planch., Fl. Serres Jard. Eur. 7: 161. 1851-52. 

(US). Juncos: Sierra de Juncos, Sintenis 1882 

(US). 

Erect, perennial, corm-producing herbs that spread vegetatively by means of stolons. Leaves 

narrowly ensiform. Inflorescence a loosely-branched panicle bearing showy orange to scarlet flowers; 

outer bracts acute or obtuse and apiculate at the apex, 4-9 mm long, spreading. Capsules subglobose, as 

broad as or broader than long, 3-grooved or 3-lobed; seeds 1 or 2 in each locule. An African genus of 6 

species, often cultivated for their showy flowers. A single species of hybrid origin has become established 

in several mountainous areas of the West Indies. 

TYPE: Crocosmia aurea (Pappe ex Hook.) Planch. ( ≡ Tritonia aurea Pappe ex Hook.) 

1. Crocosmia × crocosmiiflora (Lemoine ex E. 

Morren) N. E. Br., Trans. Roy. Soc. South 

Africa 20: 264. 1932; Montbretia × 

crocosmiiflora Lemoine ex E. Morren [as 

“crocosmiaeflora”], Belgique Hort. 31: 299. 

1881; Tritonia crocosmiiflora (Lemoine ex E. 

Morren) Nicholson [as “crocosmiflora”], 

Dict. Gard. 4: 94. 1886. Lectotype: A garden 

hybrid. Lemoine ex E. Morren, Belgique 

Hort. 31: t. 14. 1881, here designated. 

Fig. 61. C 

Corms depressed-globose, ca. 2.5 cm in 

diam., clothed with a loose fibrous covering; stems 

up to 1 m tall; leaves as long as the stems, mostly 

0.8-2 cm wide, glabrous. Flowers sessile, spicate 

and distichous on the slightly zigzag paniclebranches, 

each solitary within a small 2-bracted 

spathe; spathe bracts 6-7 mm long, acuminate- 

3. ELEUTHERINE 

Eleutherine Herb., Edward’s Bot. Reg. 29: t. 57. 1843, nom. conserv. 

aristate; perianth bright orange-scarlet, the tube 

slender, straight or slightly curved and spreading, 

the limb 6-parted, the lobes ca. 2 cm long. 

Capsules 3-4 mm in diam. 

General distribution: This garden hybrid 

between Crocosmia aurea Planch. and C. pottsii 

(Baker) N. E. Br. has been cultivated nearly 

worldwide in warm countries outdoors, and 

indoors in colder areas, and has escaped and 

become naturalized in many places, including 

mountainous parts of the West Indies. 

Distribution in Puerto Rico: Adventive in the 

Cordillera Central between 700-1100 m. Recorded 

from Adjuntas, Aibonito, Cayey, Ciales, Jayuya, 

Ponce, and Villalba. 


Jayuya: Toro Negro Forest Reserve, Acevedo- 

Rdgz. & Angell 9424 (NY, UPR, UPRRP, US). 

Acaulescent herbs arising from tunicate bulbs. Leaves few, narrowly ensiform, long-acuminate at 

apex, strongly plicate or plaited. Inflorescences on long slender scapes terminated by leaf-like spathes and 

several, unequally-stalked, often recurved, bracteate flower-clusters. Flowers without perianth-tube, the 

segments separate and subequal, spreading. Stamens attached to base of the segments, with short free 

filaments. Ovary oblong, containing numerous ovules; style short, with 3 apically stigmatic branches. 

Capsules loculicidally 3-valved; seeds many, small, angulate. A Neotropical genus of 2 species of wide 

distribution. 

TYPE: Marica plicata Ker-Gawl., nom. illeg. (Moraea plicata Sw., nom. illeg., Sisyrinchium 

latifolium Sw.). 

Reference: Goldblatt, P. & N. Snow. 1991. Systematics and chromosome cytology of Eleutherine 

Herbert (Iridaceae). Ann. Missouri Bot. Gard. 78: 942-949.


Fig. 24. A-H. Eleutherine bulbosa. A. Bulb and leaves with detail of leaf base and transverse section of blade. B. Plant with 

inflorescence. C. Lateral view of part of inflorescence with an open flower. D. Lateral view of flower. E. L.s. of flower. F. Anther 

and apex of filament. G. Part of infructescence. H. Open fruit without seed. From Mori, S. et al. 1997. Vascular plants of central 

French Guiana. Mem. NYBG Vol. 76(1).

104 


1. Eleutherine bulbosa (Mill.) Urb., Repert. 

Spec. Nov. Regni Veg. 15: 305. 1918; 

Sisyrinchium bulbosum Mill., Gard. Dict. ed. 

8, Sisyrinchium no. 3. 1768; Galatea bulbosa 

(Mill.) Britton, Brooklyn Bot. Gard. Mem. 1: 

37. 1918. Neotype: Guadeloupe. Stehlé 2520 

(US-1955627), here designated. 

Sisyrinchium latifolium Sw., Prodr. 17. 1788; 

Moraea plicata Sw., Fl. Ind. Occid. 1: 92. 

1797, nom. illeg.; Eleutherine plicata Bold., 

Fl. Dutch W. Ind. Is. 59. 1909, nom. illeg.; 

Cipura plicata Griseb., Fl. Brit. W. I. 589. 

1864, nom. illeg. Type: West Indies. Plumier 

(Burman ed.), Pl. Amer., t. 46, fig. 2. 1756. 

Eleutherine anomala Herb., Edward’s Bot. Reg. 

29: t. 57. 1843. Type: Bot. Reg. 29: t. 57. 

1843. 

Fig. 24. A-H 

Bulb ovoid, 3 cm or more in diam., the tunica 

reddish brown. Leaves 1 or 2, linear-lanceolate, 

14-70 × 0.5-3.5 cm, long- and sharply-acuminate 

at apex. Scapes to 67 cm long; spathes oblong, 1.5 

4. NEOMARICA 

Neomarica Sprague, Bull. Misc. Inform. Kew 1928: 280. 1928. 

cm long; branched clusters of flowers with slender 

pedicels; perianth white, 2-2.5 cm broad, the 

spreading segments obovate. Capsules globose, 

ca. 1 cm in diam.; seeds black. 


Antilles, and northern and western South America 

(especially Peru) to southeastern Brazil, but 

mostly absent from Amazonia. 


Islands: Introduced as an ornamental. Recorded 

from Cayey and Juana Díaz; cultivated in St. Croix 

and collected on St. Thomas in 1881, cited by 

Britton & P. Wilson (1924) for St. John apparently 

from a cultivated plant. 

Common names: Puerto Rico: Lágrimas de la 

Virgen, Mariposa, Ninfa. 

Note: Miller in his original description of this 

species mentioned that it grew naturally in the 

West Indies, but did not cite any original element, 

therefore a neotype is here designated. 


Juana Díaz: Sabana Llana, Stevenson 2307 (US). 

ST. THOMAS: Eggers s.n. (US). 

Perennial herbs with short-creeping rhizomes. Leaves elongate, leathery, ensiform, glabrous. Scapes 

erect or slanted, broadly winged (appearing leaf-like), terminating in a single elongate spathe-like leaf, 

producing from the axil 1 to few pedunculate bracteate flowers. Flowers large, actinomorphic; with 3 

spreading outer segments and 3 smaller inner ones, all with distinctive colors and markings according to 

species; inner segments bearing nectaries near the folded base; filaments free, enlarged at base, thread-like 

above; anthers connivent around the style below the stigmatic branches; style-branches crested at apex 

with stigmatic lobes below the crests. Capsules obovoid or cylindrical, truncate. Seeds angular and 

brownish, or else globose, fleshy, and reddish. A Neotropical genus of about 15 species. 

TYPE: Neomarica northiana (Schneev.) Sprague ( ≡ Moraea northiana Schneev.). 

1. Neomarica northiana (Schneev.) Sprague, 

Bull. Misc. Inform. Kew 1928: 280. 1928; 

Moraea northiana Schneev., Icon. Pl. Rar. t. 

41, 42. 1793. Lectotype: Peru. Schneev., Icon. 

Pl. Rar. t. 41. 1793, here designated. 

Plants up to 1.2 m tall or more. Leaves often 1 

m long and up to 6 cm wide. Outer flower 

segments white, shading to yellow and mottled 

with deep red towards base; inner segments barred 

with light blue near tips. Fruits green. 

General distribution: Native of Peru; widely 

cultivated elsewhere. 

Distribution in Puerto Rico: Widely cultivated, 

reported by Liogier & Martorell (1982) to be 

naturalized in forests at Jajome, Cayey. 


Neomarica caerulea (Ker Gawl.) Sprague is 

also sometimes cultivated, but is not recorded as 

being naturalized in Puerto Rico. It is easily 

distinguished from N. northiana by its flowers 

with deep purple-blue outer segments. This 

species is native to Brazil, and it is naturalized in 

Jamaica.

Sisyrinchium L., Sp. Pl. 964. 1753. 


5. SISYRINCHIUM 

Perennial or sometimes annual, usually tufted, grass-like, scapose herbs mostly of small stature with 

fibrous roots and small or obsolete rhizomes. Leaves basal, with linear blades. Scapes 2-edged or 2winged, 

when branched, each node bearing a bract resembling a leaf blade. Flowers in terminal clusters 

arising from within spathes of usually 2 bracts. Perianth blue, white, or yellow, with 6 spreading lobes. 

Stamens 3, with filaments united nearly to the apex, the anthers clustered. Ovary 3-locular, the stylebranches 

3, filiform, alternating with the anthers or else the style scarcely divided; ovules few to many in 

each locule. Capsules more or less globose, sometimes depressed or sometimes longer than thick, 

sometimes angled, opening at the apex. Seeds smooth or pitted. A chiefly North American genus of more 

than 75 species. 

TYPE: Sisyrinchium bermudiana L. 

1. Sisyrinchium rosulatum E.P. Bicknell, Bull. 

Torrey Bot. Club 26: 228. 1899. Syntypes: 

United States; South Carolina. L. R. Gibbs s. 

n. (NY); United States; Alabama. C. Mohr s. 

n. (MO). 

Sisyrinchium exile E. P. Bicknell, Bull. Torrey 

Bot. Club 28: 573. 1901. Type: United States; 

Texas. J. E. Bodin s. n. (MIN, US). 

Small annual herb with fibrous roots only. 

Leaves erect, often densely tufted, mostly 2-8 cm 

long (rarely longer), 1-2 mm wide, with entire 

margins. Scapes erect, up to 11 cm long or more, 

very slender, the body 0.2-0.5 mm thick, the wings 

0.2-0.3 mm wide. Spathes narrowly foliaceous, 

with unequal bracts, the outer one longer and 

slightly falcate. Flowers urceolate, whitish or 

yellowish, sometimes with purple lines and a 

purple eye-ring; pedicels capillary, flexuous, up to 

6. TRIMEZIA 

Trimezia Salisb. ex Herb., Edward’s Bot. Reg. 30 misc. 88. 1844. 

1 cm long or more. Capsules globose or nearly so, 

light brownish with purple stripes on the sutures, 

2.5-3 mm in diam.; seeds many, very small (0.5- 

0.7 mm thick), asymmetrically angled. 


States, Hispaniola, and Puerto Rico. 

Distribution in Puerto Rico: Confined to 

higher parts of the Cordillera Central at elevations 

of 950-1250 m. Recorded from Jayuya, Ponce, and 

Villalba. 

Note: The closely related S. micranthum Cav. 

occurs widely in Mexico, Central and South 

America, and has been attributed to Hispaniola. If 

these two annual species are combined, the correct 

name would appear to be S. micranthum. 


Jayuya: Cerro Maravillas, Liogier et al. 28916 

(UPR). Ponce: Monte Jayuya, Axelrod & Chavez 

4279 (US). 

Perennial cormous herbs, the tunica splitting and shredding into coarse fibers. Leaves flat, more or 

less linear, equitant, with prominent mid rib. Flowers stalked, actinomorphic, with more than one 

produced from each spathe; perianth tube lacking, the segments in two dissimilar whorls of 3, the outer 

ones obovate with broad claw, the inner ones smaller and more or less deflexed. Stamens short, erect, the 

filaments free; ovary clavate, the style basally subulate, 3-branched above, the branches ending in small 

tubercles overtopping the small stigmas. Capsules oblong, exserted beyond the bracts. A small 

Neotropical genus of 5 species occurring from Mexico to tropical South America and also in the West 

Indies. 

TYPE: Trimezia meridensis Herb.

106 


1. Trimezia steyermarkii R. C. Foster, Rhodora 

64: 310. 1962. Type: Guatemala. Steyermark 

49539 (holotype: GH; isotype: F). 

Trimezia martinicensis sensu Liogier & Martorell, 

Fl. Puerto Rico and Adjacent Is. 1982, non 

(Jacquin) Herbert, 1844. 

Fig. 61. B 

Corms ovoid, 2-4 cm in diam., with brown 

tunica fibers. Leaves rather thin-textured, 30-70 × 

2-4 cm, with a prominent mid rib. Scapes equaling 

or slightly overtopping the leaves, the inflorescence 

terminal, several-flowered; spathe-bracts unequal, 

the outer 2 cm long, the inner one to 3 cm long or 

more. Pedicels glabrous, more or less equaling the 

spathes at anthesis. Ovary glabrous, clavate, less 

than l cm long. Flowers yellow with mottled 

brownish purple bands toward base, ca. 3 cm wide 

when fully expanded. Capsules and seeds not 

CULTIVATED GENERA 

observed. 

General distribution: Mexico and Central 

America, cultivated elsewhere and sometimes 

naturalized. 

Distribution in Puerto Rico: Frequently 

cultivated as an ornamental. Recorded as 

naturalized in Arecibo, Patillas, and Río Grande. 

Note: This species resembles T. martinicensis, 

with which it has sometimes been confused. T. 

steyermarkii however is much larger than T. 

martinicensis. The latter species, common in the 

Lesser Antilles and Jamaica, is strangely absent 

from other parts of the Greater Antilles, but occurs 

widely in South America. 


Arecibo: Río Abajo State Forest, vicinity of 

Campamento Radley, Proctor 43015 (US). Río 

Grande: El Yunque Forest Reserve, Acevedo- 

Rdgz. & Angell 9455 (UPR, UPRRP, US). 

Gladiolus cuspidatus Jacq. (Jayuya: Cordillera Central, Bo. Saliente, Proctor et al. 43969, US) and 

G. x gandavensis Van Houtte (Orocovis: Doña Juana, Toro Negro, Liogier & Martorell 35081I, UPR and 

Ponce: La Carmelita, Liogier et al. 34392, UPR), both natives of South Africa, are planted as ornamentals 

in Puerto Rico. Gladiolus communis L., native to Europe, was cited by Martorell et al. (1981) as cultivated 

in Puerto Rico. 

Tigridia pavonia (L. f.) DC., native to Mexico, was cited by Martorell et al. (1981) as cultivated in 

Puerto Rico. 

Family 14. AMARYLLIDACEAE Amaryllis Family 

Amaryllidaceae J. St. Hil., Expos. Fam. Nat. 1: 134. 1805, nom. conserv. 

by G. R. Proctor, P. Acevedo-Rodríguez, & M. T. Strong 

Perennial herbs with usually bulbous or rarely rhizomatous stocks. Leaves few, basal, entire, usually 

rather soft. Flower bisexual, actinomorphic or subregular, often showy, solitary or umbellate and 

subtended by spathaceous bracts at the top of a naked simple scape. Perianth petaloid of 6 segments in 2 

similar series often united below into a tube. Stamens 6, opposite the perianth-segments, hypogynous or 

epipetalous; filaments free or joined together, often curved upwards in laterally directed flowers; anthers 

2-locular, basi- or medifixed, opening lengthwise or rarely by terminal pores. Ovary superior or more 

usually inferior, usually 3-locular with axile placentas; style slender; ovules mostly numerous, 

anatropous. Fruit a capsule or fleshy berry. Seeds with fleshy endosperm and small embryo. A family of 

about 90 genera and some 1100 species 

TYPE: Amaryllis L.



1. Flower solitary …….................................................................................................…. 5. Zephyranthes 

1. Flowers in umbels, not solitary ……..............................................................................................…… 2 

2. Flowers with a staminal cup, always white ………....................................................…………. 3 

3. Filaments short or apparently absent …………….......................................…….. 2. Eucharis 

3. Filaments very long, up to ca. 6 cm ……………....................................….. 4. Hymenocallis 

2. Flowers without a staminal cup, perianth white or variously colored ………...................……… 4 

4. Perianth orange, bearing a corona …………….....................................………3. Hippeastrum 

4. Perianth white, rose, crimson, or striped of similar colors, without a corona ……. 1. Crinum 

Crinum L., Sp. Pl. 291. 1753. 

1. CRINUM 

Plants bearing subterranean bulbs with short to long sheathing necks. Leaves succulent, linear or 

strap-shaped, somewhat broadened toward apex, spreading or arching, the margins entire or toothed. 

Inflorescence an umbellate cluster of few to many flowers subtended at base by 2 large, broad, spathe-like 

bracts, the scape solid, erect, green or brightly colored. Flowers large, showy, short-pedicellate; perianth 

white, rose, crimson or striped of similar colors, funnel-shaped or trumpet-shaped, with a long, cylindrical, 

straight or curved tube subtended by narrow bracts at base, the lobes linear, lanceolate, or oblong, 

subequal; stamens spreading or declinate, the filaments long, filiform, inserted in the throat of the perianth 

tube, the anthers linear versatile; ovary globose to oblong with 2 to several ovules in each locule, the style 

long, filiform, the stigma minute, capitate. Fruit a subglobose, tardily dehiscent capsule; seeds large, 

bulbiform, fleshy, green, with very thick endosperm. A genus of approximately 130 species, widely 

distributed in tropical, subtropical and warm-temperature regions worldwide. 

LECTOTYPE: Crinum americanum L., designated by Britton & P. Wilson, Bot. Porto Rico 5: 160. 1923. 

Key to species of Crinum 

1. Leaf margins minutely denticulate. …......................................................................................………. 2 

2. Perianth lobes pure white ...................................................................................... [C. americanum] 

2. Perianth white with a pink or reddish median stripe …………......................…….2. C. zeylanicum 

1. Leaf margins smooth. …………..................................................................................................…….. 3 

3. Perianth lobes rose, adaxially with a darker central stripe; filaments of stamens purple 

….......................................................................................................................... 1. C. asiaticum 

3. Perianth lobes red or rose throughout; filaments of stamens red ……..........…… [C. erubescens] 

1. Crinum asiaticum L., Sp. Pl. 292. 1753. 

Lectotype: Malabar or Ceylon. Hermann, 

Hort. Lugd.-Bat. Cat. 683, 1687, designated 

by Verdoorn in Killick (ed.), Fl. Pl. Africa 47 

: pl. 1875. 1983. 

Crinum amabile Donn ex Ker-Gawl., Bot. Mag. 

39: fig. 1605 A & B. 1814. Lectotype: East 

Indies. Bot. Mag. 39: fig. 1605A. 1814, here 

designated.* 

Crinum amabile Donn, Hort. Cantab. ed. 6, 83. 

1811, nom. nud. 

Bulb with a rather massive, stem-like neck, 

rising 30 cm or more above ground. Leaves 

numerous, spreading, narrowly lanceolate, 70-100 

× 9-15 cm, clasping at the slightly narrowed base 

and gradually tapering to the apex, the margins 

entire. Scape 40-80 cm tall, 2-keeled. Spathevalves 

deltate, reddish, 12-18 cm long. Umbel of 

18-30 pedicellate flowers; perianth with slender, 

cylindrical, deep rose or red tube, 7-12 cm long; 

lobes ca. 2 cm broad, equaling or exceeding the 

tube, pink outside and whitish with deep pink 

median stripe within; stamens and stigmas purple. 

Fruit not seen.

108 


General distribution: Native to India, but 

commonly cultivated in tropical and subtropical 

regions, and in greenhouses in temperate areas. 


cultivated in Puerto Rico and the Virgin Islands. 

Persisting after cultivation on Mona Island, 

vicinity of the light house. 

Note: * R.A. Howard lectotypified this name 

with Bot. Mag. 39: fig. 1605, wich includes figures 

A & B. Although his lectotypification does not 

represent a problem for the interpretation of this 

name, the present writers want to further restrict 

its typification to plate 1605A as it is a more 

representative illustration. 


Mona Island, Proctor 43131 (US-2). 

2. Crinum zeylanicum (L.) L., Syst. Nat. ed. 12, 

2: 236. 1767; Amaryllis zeylanica L., Sp. Pl. 

293. 1753; Crinum latifolium var. zeylanicum 

(L.) Hook. f. in Trimen, Handb. Fl. Ceylon 4: 

272. 1898. Lectotype: J. Commelijn, Horti 

med. amstelod. 1: tab. 73. 1697, designated by 

Dassanayake, Taxon 30: 481. 1981. 

Figs. 25. A, B; 61. H 

Perennial, acaulescent herb, with a bulbous 

base; bulb ovoid to subglobose, 9-13 × 5-11 cm, 

with a short neck; roots stout, coarse, 3-5 mm 

wide. Leaves many; blades 80-125 × 2.8-5.5 cm, 

strap-shaped, reflexed, finely veined, acuminate at 

apex, the margins with remote, blunt teeth. Scape 

25-50 cm long, glabrous, reddish tinged; spathe- 

2. EUCHARIS 

Eucharis Planch. & Linden in Linden, Cat. Pl. Exot. 8: 3. 1853, nom. conserv. 

like bracts deltate, membranaceous, 6-7 × 2-3 cm. 

Flowers 5-10, subsessile or short-pedicellate; 

perianth segments oblong-lanceolate, 8-12 × 2-3 

cm, white with a pink or reddish median stripe, the 

tube curved at maturity 8-13 cm × 3-4 mm; anthers 

1-2 cm long, greenish, the filament 6.5-8 cm long, 

whitish or pink; style 15-20 cm long. Fruit not 

seen. 

General distribution: Native to Sri Lanka and 

India, widely cultivated in tropical areas and in 

greenhouses in temperate regions. 


Islands: Apparently naturalized in Cabo Rojo, 

Mona Island, San Juan, and Utuado; St. Croix, St. 


Common name: Puerto Rico: Lirio. 


Cabo Rojo: Bo. Boquerón, Proctor 44947 (US). 

Mona Island, Acevedo-Rdgz. & Siaca 4376 (NY, 

SJ, US). San Juan: Bo. Puerta de Tierra, Proctor 

43989 (US). Utuado: N.L. Britton & Cowell 1236 

(US). ST. CROIX: Bassin Yard, Ricksecker 494 

(US). ST. JOHN: Coral Bay Quarter, Road to Ajax 

Peak, Acevedo-Rdgz. et al. 5112 (NY, UPRRP, 

US). ST. THOMAS: Eggers s.n. (US). 


Crinum americanum L. and C. erubescens 

Aiton are commonly cultivated, but the species are 

not known to become naturalized. Crinum 

giganteum Andr., from Africa, was recorded by 

Eggers as cultivated in the Virgin Islands 

according to Britton & P. Wilson (1924). 

Bulbs tunicate. Leaves several, the blades broad, abruptly narrowed into slender petioles. 

Inflorescence scapose, bracteate below the umbellate flowers. Flowers pedicellate; perianth tube 

cylindrical, somewhat curved dilated above, the segments equal, spreading, ovate or oblong; stamens 

inserted at the throat, the filaments appendaged below the middle, the anthers versatile; ovary globose, 3celled, 

with many superposed ovules in each cell, the style filiform, the stigma capitate, 3-lobed. Capsule 

lobed, dehiscent; seeds large. A tropical South American genus of 19 species. 

TYPE: Eucharis candida Planch. & Linden. 

References: Meerow, A. W. 1989. Systematics of the Amazon lilies, Eucharis and Caliphruria 

(Amaryllidaceae). Ann. Missouri Bot. Gard. 76: 137-220. Meerow, A. L. & B. Dehgan. 1984. Reestablishment 

and lectotypification of Eucharis amazonica Linden ex Planch. (Amaryllidaceae). Taxon 

33: 416-422.


1. Eucharis amazonica Linden ex Planch., Fl. 

Serres Jard. Eur. 12: 1216. 1857. Lectotype: 

Peru. Planch., Fl. Serres Jard. Eur. 12: fig. 

103. 1857, designated by Meerow & Dehgan, 

Taxon 33: 421. 1984. 

Eucharis grandiflora sensu authors, non Planchon 

& Linden, 1853. 

Fig. 61. D 

Bulbs globose, to 7 cm in diam. Leaves with 

petioles 15-30 cm long; blades oblong to elliptic or 

oval, 25-35 × 10-15 cm, the base cuneate or 

rounded, the apex abruptly short-acuminate. 

Scape to 50 cm tall. Inflorescence umbellate, with 

4-6 hanging flowers. Pedicels 1-2.5 cm long; 

3. HIPPEASTRUM 

Hippeastrum Herb., Bot. Reg. 7 (App.): 31. 1821, nom. conserv. 

perianth tube 5 cm long, the segments oblongovate, 

3-5 cm long, white with apiculate apex; 

filament flanges connate, free portion lanceolate. 

Capsule 2.5 cm in diam. 

General distribution: Native of Peru, found 

in cultivation in many tropical countries. 

Distribution in Puerto Rico: Naturalized or 

persistent in forest understory. Reported from 

Adjuntas and Arecibo. 

Common names: Puerto Rico: Anunciación, 

Lirio turco. 


Adjuntas: Monte Guilarte State Forest, Proctor 

48711 (US). Arecibo: Río Abajo Forest Reserve, 

Acevedo-Rdgz. 9394 (UPR, US). 

Herbs with tunicate bulbs. Leaves basal, linear or strap-shaped. Inflorescence umbellate on stout 

hollow scape; spathe bracts lanceolate, membranaceous. Flowers few or several, stalked, declinate; 

perianth tube short or long, with scales or corona in throat, the segments spreading, nearly equal; stamens 

6, inserted at the throat, the filaments filiform, the anthers linear, versatile; ovary 3-celled, with many, 

superposed ovules in each cell, the style long, declinate, the stigma capitate or trifid. Capsule globose, 

loculicidally dehiscent; seeds flattened, black. An American genus with 40-60 species indigenous to South 

America, two of which are widely cultivated. 

TYPE: Hippeastrum reginae (L.)Herb. ( ≡ Amaryllis reginae L.), typ. conserv. 

1. Hippeastrum puniceum (Lam.) Kuntze, Revis. 

Gen. Pl. 2: 703. 1891; Amaryllis punicea 

Lam., Encycl. 1: 122. 1783; Amaryllis 

equestris Aiton, Hort. Kew. 1: 417. 1789, 

nom. illeg. Hippeastrum equestre Herb., Bot. 

Reg. 7 (App.): 31. 1821, nom. illeg. 

Lectotype: Surinam. Merian, Metamorph. 

Insect. Surinam. t. 22. 1705, designated by 

R.A. Howard, Fl. Lesser Antill. 3: 476. 1979. 

Bulbose herb; bulbs globose or globoseovoid, 

4-5 cm long; plants commonly stoloniferous. 

Leaves strap-shaped or oblong-lanceolate, 28-65 

× 3-5 cm, narrowed to a blunt apex. Scape 40-60 

cm long, terete, glaucous; spathe bracts lanceolate, 

to 7 cm long. Inflorescence a 2- to 4-flowered 

umbel. Flowers declinate; pedicels 3.5-7 cm long; 

perianth tube to 3.5 cm long, green; tepals to 9 cm 

long, spreading, elliptic-obovate or obovate, 

bright red, red-orange and with a greenish base. 

Capsule rare, to 2 cm in diam. 

General distribution: Apparently native to 

northern South America, cultivated and naturalized 

in southern United States (Louisiana), Central 

America and throughout the West Indies. 


Islands: Reported for Arecibo, Bayamón, Cabo 

Rojo, Fajardo, San Juan, and Yauco; St. Croix, St. 

John, St. Thomas, and Tortola. 

Common names: Puerto Rico: Amapola, Lirio 

rojo. 



10555 (US). Bayamón: Underwood & Griggs 852 

(US). Cabo Rojo: Sintenis 778 (US-2). Fajardo: 

Sintenis 1281 (US-2). San Juan: Río Piedras, 

Heller & Heller 966 (US). Yauco: Sargent 512 

(US). ST. CROIX: Bassin, Ricksecker 286 (US). 

Bassin Yard, Ricksecker 393 (US). ST. THOMAS: 

Eggers 1325 (US).

110 


4. HYMENOCALLIS 

Hymenocallis Salisb., Trans. Hort. Soc. London 1: 338. 1812. 

Tunicate, bulbous plants. Leaves sessile or petioled, linear or lanceolate. Scape solid, angled or 

flattened and ridged. Inflorescence umbellate, subtended by spathe-like, lanceolate, membranaceous 

bracts. Flowers sessile or slightly stalked or becoming so in fruit, each flower with 2 narrow, 

membranaceous bracts; perianth united below into a long, cylindrical tube, the 6 lobes spreading or 

recurved, linear, white; stamens 6, the filaments free above, united below into a long, cylindrical, flaring 

or undulate cup, often with processes between the filaments, the anthers linear, versatile; ovary 3-celled, 

with 2 or more ovules in each locule, the style filiform, long-exserted, the stigma capitate. Capsule fleshy; 

seeds 1 or 2 per locule, large, green and fleshy. An American genus of about 50 species found in warm 

climates. 

LECTOTYPE: Hymenocallis littoralis (Jacq.) Salisb. ( ≡ Pancratium littorale Jacq.), designated by 

Britton & A. Brown, Ill. Fl. N. U.S. ed. 2, 1: 533. 1913. 

References: Sealy, J. R. 1954. Review of the genus Hymenocallis. Kew Bull. 1954: 201-240. Smith, 

G. L. & M. A. Garland. 2003. Nomenclature of Hymenocallis taxa (Amaryllidaceae) in Southeastern 

United States. Taxon 52: 805-817. 

Key to the species of Hymenocallis 

1. Leaves elliptic, clearly petiolate …….......................................................................…… 3. H. speciosa 

1. Leaves oblong, linear, or oblanceolate, long-tapering at base. ….........................................………… 2 

2. Floral tube 4-6.5 cm long; tepals 8-12 cm long ……….................................…… 1. H. caribaea 

2. Floral tube 8-15 cm long; tepals 9-14.5 cm long. ……..................................................……….. 3 

3. Leaves 4-8.5 cm wide …………...................................................................…. 2. H. latifolia 

3. Leaves 2-2.5 cm wide ……...................................................................………. [H. praticola] 

1. Hymenocallis caribaea (L.) Herb., Bot. Reg. 7 

(App.): 44. 1821; Pancratium caribaeum L., 

Sp. Pl. 291. 1753. Lectotype: Jamaica. J. 

Commelijn, Horti med. amstelod. 2: t. 87. 

1701, designated by R.A. Howard, J. Arnold 

Arbor. 60: 295. 1979. 

Pancratium declinatum Jacq., Select. Stirp. Amer. 

Hist. 99. 1763; Hymenocallis declinata 

(Jacq.) Sweet, Hort. Brit. ed. 2, 513. 1830. 

Type: Type unknown, from Martinique and 

Cayenne. 

Fig. 25. C, D 

Herb of sandy coastal areas. Leaves many, 

green, oblanceolate, 30-60 × 5-7.5 cm, tapered to 

an acute or acuminate apex, slightly narrowed at 

the base. Scape 30-60 cm long; bracts ovate to 

lanceolate, 3-6 cm long. Flowers 8-10 or more, 

fragrant; perianth tube 4-6.5 cm long, tepals 9-11 

cm long, white; staminal cup funnel-shaped, 

margins erect 2-3 cm high, the filaments 3-5 cm 

long. Fruit globose, 1.5-2 cm in diam., lobed. 

General distribution: Jamaica, Virgin Islands, 

and the Lesser Antilles. 

Distribution in the Virgin Islands: St. Croix 

and St. John (fide Sealy, 1954) . 

Common names: Virgin Islands: Lady bug, 

White lily, Spider Lily. 


Maho Bay Quarter; trail to Brown Bay, Acevedo- 

Rdgz. et al. 1873 (VINPS, NY, UPR). 

2. Hymenocallis latifolia (Mill.) M. Roem., Fam. 

Nat. Syn. Monogr. 4: 168. 1847; Pancratium 

latifolium Mill., Gard. Dict. ed. 8, no. 8. 1768. 

Type: Jamaica. Sloane IV-112 [as 115] (BM- 

589968). 

Hymenocallis expansa (Sims) Herb., Bot. Reg. 7 

(App.): 44. 1821; Pancratium expansum 

Sims, Bot. Mag. t. 1941. 1819. Type: Country 

unknown. Bot. Mag. t. 1941. 1819. 

Figs. 25. C-D; 61. F 

A large vigorous plant with numerous 

evergreen leaves. Leaves broad oblong-ensiform 

or narrowly and obliquely oblong-elliptic or


Fig. 25. A-B. Crinum zeylanicum. A. Habit and inflorescence. B. Upper portion of stamen showing anther. C-D. Hymenocallis 

latifolia. C. Upper portion of scape with open flowers. D. Infructescence. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. 

New York Bot. Gard. 78.

112 


oblong-oblanceolate, tapering to the acute apex 

and more gradually tapered to the lowest quarter 

which is distinctly narrower, strongly channeled 

and thick, 40-77 cm long, 3.8-7.5 cm wide in upper 

part, 2.8-3 cm wide in lowermost quarter. Scape 

40-60 (-80) cm high; flowers 10-20, sessile. 

Perigone: tube 8-10.6 cm long; segments 9-14.5 

cm long. Staminal cup narrowly funnel-shaped 

with erect margins, (2.5)2.7-3.5 cm high, 

somewhat tube-like in lower third then widened 

upwards, 2.4 cm wide at the mouth, somewhat 

fluted and with a large usually bifid tooth between 

the filaments; 4.5-6 cm long. Ovary with 2, rarely 

3, ovules per carpel. 


the southeastern United States, Cuba, Jamaica, 

Caiman Islands, Hispaniola, Puerto Rico, and 

Martinique. 


Islands: Reported for Cayo Lobos, Cayo Ramos, 

Coamo, Fajardo, Guayama, Juana Díaz, Naguabo, 

Mona Island, San Juan, San Lorenzo, San 

Sebastián, and Vieques; St. Croix, St. John, St. 

Thomas, and Virgin Gorda. 

Common names: Puerto Rico: Lirio blanco, 

Lirio de playa; Lirio; Virgin Islands: Spider Lily. 


Coamo: Coamo Springs, N.L. Britton & Cowell 

1341 (US). Guayama: From Guayama to Aguirre, 

Underwood & Griggs 387 (US). San Juan: 

Santurce, Heller 257 (US-2). Vieques: Isabel 

Segunda to Santa Maria, Shafer 2588 (US-2). ST. 

CROIX: Bassin, Ricksecker 460 (US). ST. JOHN: 

Coral Bay Quarter, along dirt road to Bordeaux 

Mountain, Acevedo-Rdgz. et al. 5130 (NY, US). 

ST. THOMAS: Elizabeth, along road 334, Acevedo- 

Rdgz. 11359 (US-2). VIRGIN GORDA: Gorda Peak 

National Park, Acevedo-Rdgz. & Clubbe 10892 

(US). 

5. ZEPHYRANTHES 

Zephyranthes Herb., Bot. Reg. 7 (App.): 36. 1821, nom. conserv. 

3. Hymenocallis speciosa (L.f. ex Salisb.) Salisb., 

Trans. Hort. Soc. London 1: 340. 1812; 

Pancratium speciosum L.f. ex Salisb., Trans. 

Linn. Soc. London 2: 73, t. 12. 1794. Type: 

Country unknown. Trans. Linn. Soc. London 

2: t. 12. 1794. 

Leaves evergreen, several, rosulate, the 7-9 

new leaves of the current year well-developed at 

flowering time, arising in the middle of the leaves 

of the previous season, distinctly petiolate, very 

variable in size, 38-60(95.5) cm long; blades 

broadly elliptic or broadly oblong-elliptic, shortly 

acute, cuneate at base, 26-46 cm long and 8-13.3 

cm wide but 57-65.5 cm long and 10.5-15.5 cm 

wide in some cultivated plants; petioles broad, 9- 

17 cm long. Scape 30-40 cm long, glaucous, 

ancipitous; flowers 7-12, widely spreading, 

pedicellate; pedicels to 1 cm long. Perigone tube 

(6.5)7-9 cm long; segments (9)9.5-11.5 cm long, 

sometimes 12.5-15 cm in larger cultivated plants. 

Staminal cup funnel-shaped (2.2)2.5-3.5 cm long 

(4-5 cm in larger cultivated plants ), toothed 

between the filaments; filaments almost erect, 3- 

4.5 (-5) cm long. Ovary with 2 ovules in each 

locule. 

General distribution: Native to the West 

Indies. 


Islands: Known only in cultivation in San Juan; 

and from a sterile specimen from St. John (N. L. 

Britton & Shafer 1207, US). 


There is a collection (Proctor 44848 US) from 

San Juan that seems to be referable to 

Hymenocallis praticola Britton & P. Wilson 

except for its wide floral tube. Also, Britton & P. 

Wilson (1924) list H. undulata Herb., from 

Venezuela, as observed in St. Thomas. 

Herbs with tunicate bulbs. Leaves few to several, linear, flat or channeled, glabrous, appearing with 

the flowers or after them. Scape 1-flowered, spathe-like bracts 2, variously united. Flowers with pedicel 

longer or shorter than the spathe; perianth funnelform, erect or ascending, white, pink, red or yellow, the 

6 lobes subequal; stamens 6, equal or nearly so, the filaments glabrous, adnate to the perianth tube, the 

anthers versatile; ovary glabrous, 3-celled with numerous ovules in 2 rows in each locule, the style


filiform, 3-parted or 3-lobed. Capsule 3-celled, loculicidally 3-valved; seeds black, flattened. A genus of 

35 to 40 species, distributed in warm climates of the New World. 

TYPE: Zephyranthes atamasco (L.) Herb. ( ≡ Amaryllis atamasco L.), typ. conserv. 

Key to the species of Zephyranthes 

1. Perianth yellow ………...................................................................................................… 1. Z. citrina 

1. Perianth white, pink, or red. ………...............................................................................................….. 2 

2. Perianth white …………............................................................................….. 4. Z. puertoricensis 

2. Perianth pink or red. ………............................................................................................………. 3 

3. Perianth 6.3-9 cm long ………….......................................................……. 2. Z. grandiflora 

3. Perianth 2.5-4 (-5.5) cm long. …..................................................................……………… 4 

4. Pedicels (2-) 2.5-4.5 cm long ……........................................................…… 5. Z. rosea 

4. Pedicels 0.4-0.8 cm long ………...................................................………. 3. Z. proctorii 

1. Zephyranthes citrina Baker, Bot. Mag. 108: t. 

6605. 1882. Type: Guyana; Demerara. Baker, 

Bot. Mag. 108: t. 6605. 1882. 

Zephyranthes eggersiana Urb., Symb. Antill. 5: 

292. 1907. Syntypes: Cuba. van Hermann 803 

(B?); Haiti. Eggers 2834 (NY!). 

Amaryllis aurea Ruiz & Pav., Fl. Peruv. 3: 56. 

1802, nom. illeg., non. L’Heritier, 1789; 

Zephyranthes aurea Baker, Handb. Amaryll. 

37. 1888, nom. illeg. Type: Peru. Ruiz & 

Pavón, Fl. Peruv. t. 286a. 1802. 

Bulb 2.5 cm in diam., the neck narrow, 

elongate. Leaves 4 or 5, narrowly linear, 15-32 cm 

× 2-3 mm, channeled, apex obtuse. Scape 10-25 

cm long, spathe bracts 2 cm long. Flowers erect, 

with pedicels 2-4 cm long, exceeding the bracts in 

fruit; perianth 3-4 cm long, yellow except for short 

greenish tube; style shorter than the stamens, the 

stigma capitate, 3-lobed. Capsule depressed, 1 cm 

in diam. 


(Florida and Mississippi), Mexico, Central 

America, Cuba, Hispaniola, Grand Cayman, 

Jamaica, Puerto Rico, and northern South 

America. 


San Juan and Vega Alta. 

Common name: Puerto Rico: Duende 

amarillo. 


San Juan: Proctor 44849 (SJ). Vega Alta: Proctor 

46463 (SJ). 

2. Zephyranthes grandiflora Lindl., Bot. Reg. 

11: t. 902. 1825. Type: Mexico. Lindley, Bot. 

Reg. 11: t. 902. 1825. 

Zephyranthes carinata Herb., Bot. Mag. 52: t. 

2594. 1825; Atamosco carinata (Herb.) P. 

Wilson in Britton & P. Wilson, Bot. Porto 

Rico 5: 159. 1924. Type: Mexico. Herbert, 

Bot. Mag. 52: t. 2594. 1825. 

Bulb ovoid, 2-3 cm in diam., the neck short. 

Leaves present at anthesis, 4-6, linear, 15-40 (-55) 

cm × 4-9 mm, subacute at apex, bright green, 

reddish tinged proximally. Scape slender, 10-33 (- 

50) cm long, hollow; spathe bracts 3-5 cm long, 2- 

5 mm wide. Flower pedicels 2-3.5 cm long; 

perianth erect, funnel-shaped, 6.3-9 cm long, 

bright rose-red or rose, the tube 1.3-2 cm long, the 

perianth segments obovate, 5-7.5 × 1.3-2 cm wide; 

stamens included, half the length of the perianth; 

anthers linear; style filiform, clavate distally, the 

stigma 3-lobed, the lobes recurved. Capsule shortpedicelled, 

broadly rounded or 3-lobed, broadly 

rounded; seeds semi-ovate, compressed, black, 

shiny. 


America, Greater Antilles, and northern South 

America. 


Islands: Ciales and Mayagüez; St. Croix. 

Common names: Puerto Rico: Adelfa, 

Duende rosado. 


Ciales: Toro Negro Forest Reserve, Rt. 143, 

between Rts 149 & 564, Axelrod & Wells 2200 

(UPRRP).

114 


Fig. 26. A-D. Zephyranthes proctorii. A. Habit. B. Flower, lateral view. C. Flower, l.s. showing stamens and gynoecium. D. 

Dehisced fruit.(A, D, from Proctor 43387; B,C, from C.M. Taylor 7019).


3. Zephyranthes proctorii Acev.-Rodr. & M. T. 

Strong, sp. nov. Type: Puerto Rico; 

Guayanilla: Guánica State Forest, 0.6 km 

WNW of Punta Vaquero, pond area, 19 Apr 

1987, Proctor 43387 (holotype: US; isotypes: 

NY, SJ!). 

Fig. 26. A-D 

A Zephyranthes rosea Lindley pedicellis 

brevissimis differt. 

Bulb small, 2-2.5 cm in diam., the neck 1-2 

cm long, 5 mm wide. Leaves 3-6, linear-filiform, 

6-20 cm × 1-1.5 mm, flat, obtuse at apex, bright 

green from base to apex. Scape slender, 6-18 cm × 

1-2.3 mm, spathe bracts 1.6-2.3 cm long. Flowers 

erect, with short pedicel 4-8 mm long; perianth 

funnel-shaped, 3-4 × 2-3.5 cm, bright deep pink 

or pinkish purple with yellow throat, greenish at 

base, the tube 5-8 mm long, the perianth segments 

ovate to oval, abruptly narrowed medially to a 

slender limb, the expanded distal portion 7-10 

mm wide; stamens included; anther linearfiliform, 

elongate, 7-8 mm long; style filiform, 

about equaling the stamens, the stigma 3-lobed, 

the lobes slightly reflexed. Capsule broadly 3lobed; 

seeds small, ca. 4 × 2 mm, semi-ovate, 

compressed, black, shiny. 


Rico. 

Distribution in Puerto Rico: In dry forest 

scrub and thickets on limestone. Recorded from 

Guánica, Guayanilla and Yauco. The new species 

is morphologically similar to Z. rosea from which 

it differs by the much shorter pedicels. 

Additional specimens examined: PUERTO 

RICO: Guánica: Reserva Forestal Guánica, 

Ackerman & Montalvo 1506 (UPRRP). Yauco: 

Guánica Reserve, near the end of the road, Taylor 

& Ross 7019 (UPRRP). 

4. Zephyranthes puertoricensis Traub, Pl. Life 

7: 37. 1951. Type: Puerto Rico. Traub 151 

(holotype: MO!). 

Amaryllis tubispatha sensu Ker Gawl., Bot. Mag. 

38: t. 1586. 1813, non L’Héritier, 1789. 

Atamosco tubispatha sensu Britton & P. Wilson, 

Bot. Porto Rico 5: 158. 1923, non (L’Her.) 

M. Gómez, 1914. 

Bulb subglobose, 2-2.5 cm in diam., the neck 

stout. Leaves 3, linear, 12-17 cm × 5-6 mm, apex 

obtuse. Scape 11-18 cm long; spathe bracts 2-3 

cm long, strongly bifid at apex. Flowers 

ascending; perianth white, tinged with green at 

base, the tube 2-4 mm long; tepals 4-5 cm long; 

stamens shorter than the perianth; style deeply 

trifid. Capsule 11 mm in diam. 


(Florida), Panama (?), Jamaica, Puerto Rico, 

Virgin Islands, and the Lesser Antilles. 


Islands: Described from a plant cultivated in 

Maryland (United States) from bulbs collected in 

“northwestern” Puerto Rico. Recorded from 

Fajardo and San Juan; Guana Island and St. 

Thomas. 

Common name: Puerto Rico: Duende blanco. 


Fajardo: Sintenis 1300 (US). San Juan: Bo. Puerta 

de Tierra, Proctor & Rivera 47875 (US). ST. 

THOMAS: Eggers s.n. (US). 

5. Zephyranthes rosea Lindl., Bot. Reg. 10: t. 

821. 1824; Atamosco rosea (Lindl.) Greene, 

Pittonia 3: 188. 1897. Type: Cuba. Bot. Reg. 

10: t. 821. 1824. 

Bulb small, 1.5-2.5 cm in diam., the neck 

short, slender. Leaves 3-6, linear, 10-25 (-30) cm × 

2-6 mm, flat, obtuse at apex, bright green from 

base to apex. Scape slender, 10-30 cm × 1.3-3 mm, 

spathe bracts 1.5-2.5 cm long. Flowers erect, with 

pedicel (2-) 2.5-4.5 cm long; perianth broadly 

funnel-shaped, 2.5-4 (-5.5) × 2.5-3.5 cm, rose-red, 

greenish at base, the tube short 2-3 mm long, the 

perianth segments oval to slightly obovate, (2-) 

2.5-4 × 0.9-1.5 cm; stamens included; anthers 

linear; style filiform, the stigma 3-lobed, the lobes 

slightly reflexed. Capsule rounded or broadly 3lobed; 

seeds small, semi-ovate, compressed, 

black, shiny. 


(Florida), Central America, Cuba, Bahamas, 

Hispaniola, Puerto Rico, Virgin Islands, and the 



Islands: Collected in Bayamón, Guayanilla, 

Manatí, San Juan, and Vega Alta; St. Croix. 

Common name: Puerto Rico: Duende rojo. 


Manatí: Bo. Tierras Nuevas Saliente, Axelrod et 

al. 10965 (UPRRP). San Juan: Santurce, field, 

Guzman 76 (UPRRP). ST. CROIX: Bassin, 

Ricksecker 1 (US).

116 



Amaryllis belladonna L., from South Africa, was cited by Britton & P. Wilson (1924) as planted for 

ornament in Virgin Islands gardens. 

Cyrtanthus uniflorus Ker Gawl., from South Africa, was recorded by Krebs as formerly grown in St. 

Thomas (Britton & P. Wilson, 1924). 

Molineria hortensis Britton, collected and described by Britton (1924) from a plant cultivated in a 

garden near La Muda, Caguas, Puerto Rico, origin unknown. The type is: Britton 7892 (holotype: NY). 

Narcissus tazetta L., native to Europe, was cited by Martorell et al. (1981) as cultivated in Puerto 

Rico. 

Sprekelia formosissima (L.) Herb., of Mexican origin, was cited by Britton & P. Wilson (1924) as 

occasionally grown for ornament in Virgin Islands gardens. 

Family 15. AGAVACEAE Century Plant Family 

Agavaceae Dumort., Anal. Fam. Pl. 57. 1829, nom. conserv. 

by G. R. Proctor & P. Acevedo-Rodríguez 

Chiefly large, more or less succulent, leathery or fibrous rosette herbs to very large rosette shrubs or 

trees with stout woody trunks often with secondary tissue growth. Roots fibrous and sometimes 

rhizomatous; plants sometimes proliferating by means of stolons. Leaves spirally arranged and with 

margins entire, spiny, or toothed, usually broadest near the base and gradually tapered to a sharp apex, the 

vascular bundles in the leaves are accompanied by thick, strong fibers, thus making possible the use of 

various species as textile plants. Inflorescence stalks (scapes) stout, terminal, clothed with few or many 

more or less appressed bracts; inflorescence large, paniculate with more or less cymose branches; in many 

species the plant is monocarpic, producing a giant inflorescence and then dying. Flowers usually bisexual, 

trimerous, hypogynous or epigenous, actinomorphic or slightly zygomorphic, and usually rising from the 

axils of well-developed bracts; tepals (segments) most often white or yellow, and are free or more or less 

fused into a tubular or campanulate perianth, sometimes abruptly widened and urceolate in the outer part. 

Stamens 6, inserted at or near the base of the perianth, the filaments sometimes short and relatively stout 

(Yucca) or else filiform and long-exserted (Agave); anthers introrse, often peltate, opening longitudinally; 

pollen with reticulate exine, the grains dispersed singly or in tetrads. Ovary 3-locular, each cavity with 

several to many ovules; style short to rather long, with punctiform to trilobate stigma. Fruit a dry or berrylike 

capsule containing several to many seeds; seeds often flattened or compressed. An entirely New 

World family of about 8 genera and more than 300 species. Many species have been introduced into all the 

warmer parts of the world, either for ornament or for commercial production of textiles, and some of these 

have become naturalized in various countries. 

The circumscription used for this family follows that of Dahlgren et al. (1985), as substantiated by 

Bogler & Simpson (1996). The broader concepts of R.A. Howard (1979, Flora of the Lesser Antilles) and 

Lott & García-Mendoza (1996, Flora Mesoamericana) are not supported by the recent DNA sequence 

studies of Bogler & Simpson (1996). 

TYPE: Agave L. 

References: Bogler, D. J. & Simpson, B. B. 1996. Phylogeny of Agavaceae based on its rDNA 

sequence variation. Amer. J. Bot. 83: 1225-1235. Dahlgren, R. M. T., H. T. Clifford & P. F. Yeo. 1985. 

The Families of the Monocotyledons, pp. 157-161, Springer-Verlag, Berlin, Heidelberg, New York, 

Tokyo.



1. Flowers erect on cymose or umbellate inflorescence branches; perianth yellow or greenish yellow; 

stamens exserted ………….................................................................................................… 1. Agave 

1. Flowers more or less pendulous, on racemose inflorescence branches or in panicles; perianth white or 

greenish white; stamens included. ………….............................................…………. 2 

2. Plants acaulescent with a large rosette of elongate leaves with toothed or spiny margins 

............................................................................................................................ 2. Furcraea 

2. Plants shrubby with branched woody stems each ending in a tight spiral rosette of entire dark green 

spine-tipped leaves …….....................................................................................………. 3. Yucca 

Agave L., Sp. Pl. 323. 1753. 

1. AGAVE 

Robust perennial (usually monocarpic) rosette plants, usually acaulescent but sometimes producing 

a short sub-woody trunk; roots hard, fibrous. Leaves large, succulent, fibrous, spine-tipped, the margins 

sparsely to densely armed with hard, sharp prickles (rarely smooth and entire). Inflorescence paniculate 

(rarely racemose or spike-like, these not Antillean), often up to 6 m tall or more, with a stout main stalk 

(scape) clothed with more or less appressed spine-tipped bracts; branches usually many, horizontal or 

ascending. Flowers in cymose or umbellate clusters, these often dense; perianth tubules toward the base, 

the segments imbricate in bud. Stamens 6, the filaments long-exserted, the anthers attached at the middle. 

Ovary inferior, 3-locular, the many ovules in 2 rows in each locule; style elongate with 3-lobed stigma. 

Fruit a dry, many-seeded, loculicidally dehiscent capsule; seeds flattened, black (or white if abortive). A 

widely distributed genus of more than 200 species (300 according to some authors), found from southern 

United States, Mexico and Central America, the West Indies (including the Bahamas, Greater Antilles, 

Cayman Islands, and Lesser Antilles), and South America to Peru and Argentina. 

LECTOTYPE: Agave americana L., designated by Britton & P. Wilson, Bot. Porto Rico 5: 156. 1923. 

References: García-Mendoza, A. & F. Chiang. 2003. The confusion of Agave vivipara L. and A. 

angustifolia Haw., two distinct taxa. Brittonia 55: 82-87. Gentry, H. C. 1982. Agave of Continental North 

America. 670 pp. Univ. of Arizona Press. Trelease, W. 1913. Agave in the West Indies. Mem. Natl. Acad. 

Sci. 11: 1-55, t. 1-116. 1913. 

Key to the species of Agave 

1. Leaf-margins straight, the marginal prickles usually not over 5 mm long, with lenticular bases. ….. 2 

2. Flowers bright yellow (native species). …..........................................................................……. 3 

3. Inflorescences ca. 1.2 m tall; leaves less than 1 m long. …..............................…. 2. A. minor 

3. Inflorescences 3-7 m tall; leaves to 2.75 m long. …….....................................................….. 4 

4. Leaves less than 15 cm wide; marginal prickles 1-1.5 mm long (St. Croix) 

…….................................................................................................….. 1. A. eggersiana 

4. Leaves up to 25 cm wide; marginal prickles mostly 2-5 mm long (rarely more) (Puerto 

Rico & Virgin Islands, except St. Croix) …………..........................….. 3. A. missionum 

2. Flowers pale greenish or yellowish green (introduced species, often cultivated, sometimes 

naturalized). …….................................................................................................................….. 5 

5. Leaves dark green, less than 8 cm wide, with many marginal prickles; perianth 2-3 cm long 

……...............................................................................................................…. 5. A. vivipara 

5. Leaves gray-green, glaucous (at least at first), 10-12 cm wide, with smooth margins or with 

few marginal prickles; perianth 4-6 cm long …….......................................….. 4. A. sisalana 

1. Leaf-margins undulate, the marginal prickles up to 10 mm long, borne on dilated bases of leaf-tissue 

.............................................................................................................................................................. 6

118 


6. Leaves whitish gray-green, strongly arching and often recurved; often cultivated but so far 

naturalized or seen flowering in the West Indies ………............................….. [A. beauleriana ] 

6. Leaves gray-green, straight, mostly erect; capsules black, up to 6 cm long ………. [A. americana] 

1. Agave eggersiana Trel., Mem. Natl. Acad. Sci. 

11: 28, t. 31-33. 1913. Type: St. Croix, U.S. 

Virgin Islands. Ricksecker 282 (US!). 

Acaulescent plant. Leaves numerous, forming 

a rosette, narrowly lanceolate, mostly 1.2-2 m 

long, nearly straight and erect, apex with a brown, 

terminal, awl-shaped, spine, 1.5-2 cm long, 

margins with straight or up-curved prickles 1-1.5 

mm long, with broadly lenticular blackish brown 

base. Inflorescence to 5 m tall or more, the scape 

bearing distant deltate bracts, the slightly 

ascending branches densely-flowered. Flowers 

deep yellow, on pedicels 2-2.5 cm long; ovary 2-3 

cm long, nearly equaling the length of the perianth, 

the whole flower (excluding the stamens) thus 5-6 

cm long; filaments 4.5-5 cm long, nearly twice as 

long as the tepals. Fruits unknown; after 

flowering, the panicles produce numerous 

vegetative bulbils by which the species can be 

propagated. 

General distribution: Endemic to St. Croix, 

where it now appears to be extinct in the wild. Now 

cultivated in St. Croix and St. Thomas, but 

relatively few plants exist. 


Christiansted: Rose et al. 3535 (US); District of 

Prince: S side of Alexander Hamilton Airport, 

Proctor et al. 45490 (US-2). 

2. Agave minor Proctor, sp. nov. Type: Puerto 

Rico. Cabo Rojo; Bo. Llanos Costa: Outlying 

s ridge of Sierra Bermeja, between Rancho 

Cassaba and Arroyo Cajul, elev. 100-200 m, 

15 Jun 1991 (fr), Proctor, Judd, and Rivera 

46978 (holotype: SJ, pictures at NY UPRRP, 

US!). 

A Agave missionum Trelease inflorescentia, foliis 

et fructibus minoribus differt. 

Acaulescent, the plants clustered by means of 

short stolons. Leaves ascending to erect in a tight 

rosette, lanceolate or narrowly elliptic, 40-60 × 

4.5-6 cm, the apical spine needle-like, up to 3 cm 

long, the marginal prickles ca. 1 mm long, mostly 

spaced 6-10 mm apart. Inflorescence (1 seen) 1.2 

m tall, the axis ca. 90 cm long, ca. 1.5 cm in diam. 

near the base, tapering to ca. 0.7 cm at base of 

panicle; scape-bracts widely separated, deltateacuminate 

with spinose apex, 3-4.5 cm long. 

Inflorescence branches few, short, ascending, with 

peduncles 1.5-4 cm long, the flowers densely 

clustered on very short branches from the apex. 

Perianth (only dried withered ones seen), yellow, 

ca. 1.5 cm long. Capsules rounded-oblong, ca. 2 

cm long, 1.4-1.6 cm in diam., deeply grooved 

between locules; seeds angular, rugose, ca. 5 mm 

wide. 

Agave minor differs from any other native 

Agave species by its shorter leaves and 

inflorescences. 

General distribution: Known only from a 

population of several individuals, in dry scrubland. 

3. Agave missionum Trel., Mem. Natl. Acad. Sci. 

11: 37. 1913. Type: St. Thomas, U. S. Virgin 

Islands. Trelease 15 (holotype: MO). 

Agave portoricensis Trel., Mem. Natl. Acad. Sci. 

11: 38. 1913. Type: Puerto Rico; between 

Coamo & Aibonito. Trelease 7 (holotype: 

MO?). 

Figs. 27. A-E; 61. E 

Acaulescent, not proliferating by stolons or 

offsets; leaves fleshy, forming a large rosette, 

lanceolate to broadly lanceolate, mostly 0.9-2.75 

m × 13-23 cm, dark glossy green, acuminate to a 

spinose apex, the apical spine awl-shaped, 1.5-2.5 

cm long, brown, smooth, decurrent at base and 

dorsally produced into the green leaf-tissue; 

marginal prickles brown or blackish, 3-5 (-6) mm 

long, straight or variously recurved, mostly spaced 

7-15 mm apart. Inflorescence an elongate stalked 

panicle up to 7 m tall, the scape about equaling the 

flowering part; main-panicle-branches horizontal 

or sometimes slightly ascending, bearing dense 

cymose clusters of erect yellow flowers; pedicels 

1.5-2 cm long. Flowers 4.5-5.5 cm long including 

ovaries ca. 2 cm long. Stamens long-exserted, with 

falcate anthers 1.4-1.7 cm long. Style 4-5 cm long, 

with a 3-lobed stigma. Capsules broadly oblongellipsoid 

or somewhat turbinate, 2.5-3.5 cm long, 

1.8-2.5 cm in diam. Seeds flat, irregularly and 

obtusely triangular, 5-9 mm wide. Many plants


Fig. 27. A-E. Agave missionum. A. Habit. B. Basal rosette of leaves and detail of leaf margin. C. Upper portion of inflorescence. 

D. Flower. E. Dehisced capsule. F-H. Yucca aloifolia. F. Habit. G. Basal portion of leaf, and detail of margin. H. Upper portion 

of inflorescence with open flowers. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78.

120 


produce vegetative bulbils on the inflorescence, 

sometimes mixed with capsules, or sometimes in 

the absence of capsules. 

General distribution: Endemic to Puerto Rico 

and the Virgin Islands, but absent from St. Croix. 


Islands: In dry, rocky, mostly non-calcareous 

habitats from near sea level to ca. 800 m elevation. 

Recorded from Aibonito, Coamo, Culebra, 

Guánica, Sabana Grande, Salinas, and Vieques; 

Anegada, Guana Island, St. John, St. Thomas, 

Tortola, and Virgin Gorda; also present on many 

of the smaller islets. 

Common names: Puerto Rico: Cocuiza, 

Maguey; Virgin Islands: Karata. 

Note: This species is somewhat variable in 

several characters, including size of marginal leafprickles, 

shape of panicle, texture of capsules, and 

presence or absence of bulbils. 


Coamo: vicinity of Coamo Springs, N.L. Britton & 

Brown 5702 (US). Coamo-Aibonito, Stevenson 

6934 (US). Culebra: N.L. Britton & Wheeler 236 

(US). Guánica: Ensenada, Britton et al. 8322 (US); 

Montalva, N.L. Britton et al. 4878 (US). Sabana 

Grande, Sargent 755 (US). Salinas: Bo. Río 

Jueyes, Proctor & Díaz 44753 (US-2); Bo. Lapa, 

Proctor et al. 43725 (US-2). Vieques: Ensenada 

Honda to Puerto Medio, Shafer 3018 (US). ST. 

JOHN: Lameshur: N.L. Britton & Shafer 497 (US). 

TORTOLA: Guana Island, Proctor 43413, 43699 

(US). 

4. Agave sisalana Perrine, House Rep. Document 

564: 87. 1838. Neotype: Mexico; Chiapas. 

Ocosocoautla, Gentry 16434 (US), designated 

by H.S. Gentry, Agaves Cont. N. Amer. 628. 

1982. 

Fig. 61. G 

Acaulescent or sometimes with a short caudex 

up to 1 m tall; plants spreading by numerous 

stolons. Leaves linear-lanceolate, to 1.5 m long, 

10-12 cm wide, the central ones rigidly erect, but 

many of the outer ones more or less reflexed, 

glaucous gray-green at first, becoming green; 

apical spine conic-subulate, 2-2.5 cm long, 

shallowly round-grooved at base; margins usually 

smooth but sometimes with small scattered 

prickles. Flowers in clusters of 2 or 3 on the many 

panicle-branches; perianth 4-6 cm long, yellowish 

green; stamens inserted at the level of perianthfusion, 

the filaments linear, red-spotted, 7.5-8 cm 

long; anthers 2.5-2.7 cm long, brown-spotted. 

Ovary 2-2.5 cm long with style becoming 7-8 cm 

long. Capsules rarely formed, oblong, up to 6 cm 

long, 2-2.5 cm in diam., stipitate and beaked; seeds 

(if any) probably not viable. Vegetative bulbils 

commonly produced below the flowers in the axils 

of bracts. Cytologically this species is a sterile 

pentaploid evidently of hybrid origin. 

General distribution: Probably native in the 

region of Chiapas, Mexico (Gentry, 1982), but 

now grown all over the world in hot dry regions as 

a source of useful textile fibers. It often persists 

indefinitely in abandoned plantations, and easily 

becomes naturalized. 


Islands: No voucher specimens from Puerto have 

been seen except from Mona Island, but this 

species is definitely naturalized in parts of the 

Guánica State Forest, and can be expected to occur 

elsewhere. In the Virgin Islands it was long ago 

recorded from St. Croix as “established” by 

Trelease (1913). 

Common name: Puerto Rico: Sisal. 

Note: In 1982, Gentry (p. 628) neotypified the 

name of this species with one of his own 

specimens from Chiapas, Mexico, without 

mentioning the original material collected by 

Perrine. 


Mona Island, Acevedo-Rdgz. & Siaca 4373 (FTG, 

NY, SJ, US). 

5. Agave vivipara L., Sp. Pl. 323. 1753. 

Lectotype: C. Commelijn, Praeludia Bot. t. 

15. 1703, designated by Wijnands, Bot. 

Commelins. 35. 1983. 

Acaulescent rosette plants sometimes 

developing a short trunk; freely surculose. Leaves 

narrowly linear to lanceolete, 40-75 cm × 3.5-8 

cm, the apical spine 2.4-4 cm long, the upper 

surface flat, the marginal prickles black, 2-5 cm 

long, up-curved, sinuous or recurved from a black 

deltoid base. Inflorescence up to 3 m tall or more, 

the scape-bracts spreading; flowers pale green, the 

tepals shading to light yellow toward the tips, the 

perianth ca. 2 cm long. Capsules subglobose to 

broadly ovoid, 3-5 cm long. Proliferous bulbils 

often produced on the inflorescence-branches 

after flowering.


General distribution: Occurs widely, both 

naturally and under cultivation, from northwestern 

most Mexico to Nicaragua; commonly cultivated 

in the West Indies and probably in most warm 

countries, sometimes escaping and becoming 

naturalized. 


Islands: Frequently cultivated in Puerto Rico, 

especially a yellow-margined variant (A. vivipara 

‘marginata’ (Gentry) P. Forst., Brittonia 44: 74. 

1992). The non-marginate form is recorded from 

“Tortuguero area”; escaped on Guana Island 


Manatí (?): Tortuguero, Liogier et al: 33548, 28 

2. FURCRAEA 

Furcraea Vent., Bull. Sci. Soc. Philom. Paris 1: 65. 1793. 

Oct. 1982 (UPR). 


Agave americana L., a Mexican species 

sometimes cultivated in Puerto Rico and the 

Virgin Islands, apparently is not naturalized. 

Agave beauleriana Jacobi is sometimes cultivated 

in Puerto Rico. In addition to these, Britton and P. 

Wilson (1923) listed the following as cultivated: 

A. fourcroydes Lem., of Yucatan, planted for fiber 

in Puerto Rico and the Virgin Islands; and A. 

evadens Trel., native of Trinidad, A. jacquiniana 

Schult., and A. neglecta Small, endemic to Florida, 

U.S.A., as all cultivated in St. Thomas at 

Louisenhoj. 

Monocarpic, large rosette-forming, mostly acaulescent herbs. Leaves elongate, often longitudinally 

ridged, usually green and of thick texture, spine-tipped and with marginal teeth or sometimes the margins 

smooth. Inflorescence a large open panicle, the axis bearing large triangular bracts. Flowers clustered 

along the branches, more or less pendulous; perianth funnelform, the 6 segments spreading, white or 

greenish white. Stamens shorter than the perianth, borne on the bases of the tepals, the filaments with 

swollen bases. Ovary inferior, 3-locular, with many ovules, the style angulate and thickened below, the 

stigma capitate. Capsules oblong, 3-angled; seeds flat, numerous. Capsules and viable seeds are seldom 

produced, but reproduction is frequently accomplished by means of vegetative bulbils formed on the 

inflorescence. A Neotropical genus of about 20 species. 

Lectotype: Furcraea cubensis (Jacq.) Vent. (≡ Agave cubensis Jacq.), (= Furcraea hexapetala (Jacq.) 

Urb.), designated by Britton, Fl. Bermuda 80. 1918. 

References: Álvarez de Zayas A. 1996. El género Furcraea (Agavaceae) en Cuba. Anales Inst. Biol. 

Univ. Nac. Autón. México, Bot. 67(2): 329-346. Drummond, J.H. 1907. The literature of Furcraea with 

a synopsis of the known species. Rep. (Annual) Missouri Bot. Gard. 18: 25-75. 

Key to the species of Furcraea 

1. Leaves with marginal prickles mostly paired; flowers solitary along main axis of inflorescence; tepals 

cream .........................................................................................................................… 2. F. stratiotes 

1. Leaves with marginal prickles solitary, sometimes few or lacking; flowers clustered along secondary 

inflorescence branches; tepals greenish white …..................................................................………… 2 

2. Leaves narrowly lanceolate or narrowly elliptic, with marginal prickles 0.5-2 cm apart; bulbils 

fusiform; tepals 1.4-1.9 cm long ……....................................................……….. 1. F. antillana 

2. Leaves widely lanceolate or elliptic, with marginal prickles 1.5-4(8) cm apart; bulbils ovoid; 

tepals 2.4-2.7 cm long ………….................................................................…….. 3. F. tuberosa 

1. Furcraea antillana A. Álvarez, Anales Inst. 

Biol. Univ. Nac. Autón. México, Bot. 

67(2): 331. 1996. Type: Cuba. Havana. A. 

Álvarez 63654 (holotype: HAJB). 

Large monocarpic plants. Leaves in a rosette, 

narrowly oblong-lanceolate (60)90-120(200) × 5- 

10 cm, acute or acuminate at apex with a blunt or 

sharp terminal spine, the margins with numerous

122 


curved prickles 2-5(7) mm long, brown to almost 

black, sometimes prickles few and widely 

scattered. Inflorescence 4-6 m tall, with branches 

on the 2 upper thirds. Flowers in small clusters of 

2 or 3, greenish white; pedicels 0.4-1 cm long; 

tepals elliptic, (1.2)1.4-1.9(2.7) cm. Stamens 1.2- 

2.4 cm long. Ovary trigonous, 1.8-2 cm long. 

Capsules oblong, 2.5-5.0 × 1.6-3 cm; seeds 8-12 

mm long. Bulbils fusiform. 

General distribution: Cuba, Hispaniola and 

Puerto Rico. 

Distribution in Puerto Rico: Known from two 

collections, one from Morovis (Sintenis 6854, US) 

the other from Vega Baja (Howard et al. 16988). 

2. Furcraea stratiotes J. B. Petersen, Bot. 

Tidsskr. 37: 306. 1922. Type: Nicaragua 

(from a plant brought into cultivation in the 

Copenhagen Botanical Garden, by Oersted in 

1848). Boye-Petersen s. n. (holotype: C). 

Acaulescent or nearly so; leaves many, linearlanceolate, 

up to 1 m long, and 7 cm wide near-the 

middle, light green, the apex with an awl-shaped 

indurate portion ca. 1 cm long, tipped with a 

mucron, the margins bearing numerous mostly 

twinned prickles, 1-3 mm long, those of a pair 

usually turned away from each other. Inflorescence 

slender, up to 4 m tall or more, bipinnate, the 

ultimate branchlets flexuous or up-curved, often 

bulbiferous, the bulbils to 3.5 cm long, strongly 

flattened or compressed, with 3-5 leaves usually 

showing. Lower scape bracts ca. 6 cm long, 2 cm 

broad, obtuse, the margins minutely denticulate; 

upper bracts smaller, acuminate, entire. Flowers 

solitary, cream-colored; pedicels 0.5-1 cm long; 

tepals ca. 1.4 cm long; filaments 2.5 mm long, ca. 

1 mm thick at base, subulate toward apex, with 

anthers 4 mm long, 1.5 mm thick. Ovary 8 mm 

long, obtusely triangular; style 5 mm long. 

Capsules and seeds unknown. 

General distribution: Endemic to Nicaragua, 

but naturalized in the Virgin Islands. 

Distribution in the Virgin Islands: Naturalized 

on St. Croix (rocky hillsides near Eliza’s Retreat) 

and cultivated in several St. Croix gardens. There 

was a general flowering of the wild population 

during early November, 1997. It is presumed that 

the species may have been introduced from 

Copenhagen, Denmark, after its first known 

flowering (and production of bulbils) in 1921. F. 

stratiotes has not been rediscovered in Nicaragua 

since its first collection by the Danish botanist 

Oersted in 1848, and St. Croix is therefore the only 

known locality where this species occurs in the 

wild. 

3. Furcraea tuberosa (Mill.) W.T. Aiton, Hort. 

Kew. ed. 2, 2: 303. 1811; Agave tuberosa 

Mill., Gard. Dict. ed. 8, Agave no. 4. 1768. 

Type: Hispaniola. Plukenet, Phytogr. t. 258 f. 

1. 1691. 

Furcraea hexapetala sensu Urban, 1903, non 

Jacquin,1760. 

Figs. 28. A-G; 62. A, B 

Large monocarpic plants. Leaves in a rosette, 

broadly oblong-lanceolate up to 1.5 m long or 

more, 13-18 cm broad near the middle, acuminate 

at apex with a blunt or sharp terminal spine, the 

margins usually with numerous straight prickles 2- 

4 mm long, or sometimes prickles few and widely 

scattered. Inflorescence to 7 m tall, the upper half 

loosely paniculate. Flowers solitary or often in 

small clusters of 2 or 3, pale green or greenish 

white, fragrant; pedicels 0.6-1 cm long; tepals 

more or less oblong, 2.4-2.7 × 0.9-1.5 cm. Stamens 

much shorter than the perianth. Ovary linear or 

narrowly oblong-linear, up to 2 cm long, perhaps 

not functional. Capsules and seeds unknown; the 

species reproduces entirely by means of vegetative 

bulbils borne on the inflorescence after flowering. 


Antilles. 


Islands: Apparently native, but also cultivated. 

Recorded from Bayamón, Cidra, Culebra, Fajardo, 

Guayanilla, Maricao, Morovis, Ponce, Rincón, 

Sabana Grande, San Germán, San Juan, San 

Sebastián, Vega Baja, Vieques, and Yauco; St. 

Croix and St. Thomas. 

Note: This species was incorrectly considered 

by R.A. Howard (Fl. Lesser Antill. 3: 499. 1979) to 

be endemic to the Lesser Antilles. The type of the 

species is from the island of Hispaniola and its 

identity applies to plants known from the Greater 

and Lesser Antilles as well. 

Common names: Puerto Rico: Cocuisa, 

Maguey, Maguey criollo. 


Cidra: Bo. Rabanal, roadside cliffs of Cerro Viento 

Caliente, 560 m, Proctor et al. 47640 (SJ, US).


Fig. 28. A-G. Furcraea tuberosa. A. Habit. B. Portion of leaf. C. Inflorescence branch. D. Flower. E. Stamen. F. Vegetative 

bulbil. G. C.s. of fruit showing 3-locular ovary and ovules. (A, Photo; B, from Proctor 43844; C-G, from Proctor 42326).

124 


Culebra: N.L. Britton & Wheeler 242 (US). 

Fajardo: Heller & Heller 817 (US). Guayanilla: 

N.L. Britton & Shafer 1801 (US). Maricao: 

Sargent 417 (US). Morovis: Bo. Frances, Sintenis 

6854 (US). Sabana Grande: Cust s.n. (US). San 

Juan: Gardner s.n. (US-5). Vieques: Shafer 2913 

(US). Yauco: Underwood & Griggs 722 (US). ST. 

CROIX: Bassin, Ricksecker 298 (US). ST. THOMAS: 

Morrow 71 (US). 

Yucca L., Sp. Pl. 319. 1753. 

3. YUCCA 


At least two other species (F. foetida (L.) 

Haw. and F. selloa K. Koch cv. “marginata”) have 

been cultivated in St. Croix and also possibly in St. 

Thomas. Also, Britton & P. Wilson (1923) stated 

that F. elegans Tod. had been planted in St. 

Thomas and F. andina Trel. (native to Peru) and F. 

macrophylla Baker (of the Bahamas) were planted 

at the Agricultural Experimental Station at 

Mayagüez, Puerto Rico. The fate of these plants is 

unknown; most probably they are no longer extant. 

Shrubby or tree-like woody plants with simple or branched stems, the leaves crowded in a tight spiral 

at the ends. Leaves thick, rigid, spinose at the apex, the margins entire, denticulate or filiferous. 

Inflorescence a large terminal panicle; flowers pedicellate, usually pendent or directed somewhat 

downward; perianth-segments (tepals) lance-ovate, distinct, often succulent; stamens 6, with short stout 

filaments. Ovary sessile or short-stalked, 3-locular, with numerous ovules. Fruit a fleshy or chartaceous 

capsule; seeds black, compressed. A New World genus of about 40 species occurring in the southern 

United States, Mexico, and the West Indies. 

LECTOTYPE: Yucca aloifolia L., designated by Britton & Shafer, N. Amer. Trees 151.1908. 

Key to the species of Yucca 

1. Leaves 35-60 cm long, less than 3.5 cm wide near the middle; filaments free, longer than the ovary; ripe 

fruits with purple pulp and without a papery core …………..............................… 1. Y. aloifolia 

1. Leaves to 100 cm long and up to 5 cm wide or more; filaments adnate to and shorter than the ovary; ripe 

fruits with white or yellowish pulp and a papery core …………..............… 2. Y. guatemalensis 

1. Yucca aloifolia L., Sp. Pl. 319. 1753. 

Lectotype: Dillenius, Hort. Eltham. t. 323, 

fig. 416. 1732, designated by Wijnands, Bot. 

Commelins. 140. 1983. 

Figs. 27. F-H; 62. C, D 

Stems erect, woody, simple or branching, 

most often 1-3 m tall, seldom more than 5 m tall. 

Leaves numerous in a tight spiral, dark green, 

rigid, thick, narrowly lanceolate or linearlanceolate, 

30-60 cm long (rarely more), mostly 2- 

3.5 cm wide (rarely wider), dilated at base, the 

apex armed with a sharp, cylindrical, brown or 

blackish spine; margins usually minutely 

denticulate. Inflorescence a dense, erect, terminal 

panicle 20-70 cm long; flowers numerous, 

nodding, campanulate, succulent, creamy-white 

(sometimes purple-tinged toward base), the tepals 

3-5.8 cm long. Stamens hypogynous, free, 

thickened towards apex, the anthers versatile. 

Ovary short-stalked. Fruit an indehiscent fleshy 

capsule, oblong-ellipsoid, mostly 4-6 cm long 

(rarely longer), 1.7-2.5 cm in diam. (rarely to 4 

cm), blackish with dark purple pulp. Seeds shiny 

black, flattened, round or oval, 5-6 × 6-7 mm. 

General distribution: Florida to Louisiana, 

Mexico, Bermuda, Bahamas, Greater Antilles, 

Cayman Islands, and Lesser Antilles. This species 

has often been planted as an ornamental and tends 

to persist after cultivation and become naturalized, 

so that its true natural distribution cannot be 

determined with certainty. 


Islands: Recorded from Carolina and Loíza, where


it grows in sandy coastal thickets. In the Virgin 

Islands, recorded from St. Croix and St. John, but 

probably occurs on other islands. 

Common names: Puerto Rico. Aguja de Adán, 

Bayoneta española, Mata de huevo. 


Carolina: Bo. Torrecilla Baja, E of Punta Vacia 

Talega, Axelrod & Thomas 12689 (UPRRP). ST. 

CROIX: Fosberg 59209 (US). ST. JOHN: Coral Bay 

Quarter; Emmaus along Rt. 107, Acevedo-Rdgz. & 

Reilly 2008 (NY, UPR, UPRRP, US, VINPS). 

2. Yucca guatemalensis Baker, Refug. Bot. 5: t. 

313. 1872; Yucca elephantipes E. Regel ex 

Trel., Rep. (Annual) Missouri Bot. Gard. 13: 

94. 1902, nom. illeg. Type: Guatemala. (based 

on material cultivated in England). Baker, 

Refug. Bot. 5, t. 313. 1872. 

Stems erect, woody, tree-like, often 10 cm 

thick or more, branched, up to 10 m tall or more. 

Leaves numerous in moderately dense spiral 

clusters toward ends of branches (the trunk and 

lower parts of the branches naked with somewhat 

rough bark), narrowly lanceolate, up to 1 m long 

and 4-5 cm wide near the middle, gray-green and 

of hard, rigid texture, tapering toward base and 

long-acuminate at the apex, terminating in a long, 

sharp, needle-like spine, the margins very 

minutely serrulate. Inflorescence a dense, erect, 

terminal panicle usually 65-90 cm long, with 

numerous nodding white or creamy-white flowers 

of succulent texture, on pedicels up to 2.5 cm long; 

corolla campanulate, with narrowly elliptic tepals 

mostly 3-4 cm long. Stamens hypogynous, the 


short, thick filaments closely appressed to the 

lower half of the ovary, the short anthers opening 

introrsely. Ovary oblong, capped by a short style 

and 3 bifid stigmatic lobes ca. 3.5 mm long. 

Capsules oblong-ovoid, up to 9 cm long, 

containing white or yellowish pulp and a papery 

core. 


the lowlands of Guatemala, ascending to medium 

heights in the mountains, and occurring commonly 

in cultivation and naturalized from Mexico to 

Costa Rica, also widely cultivated in the West 

Indies. The flowers are edible and are a common 

article of food in Guatemala and adjacent areas. 

They can be eaten raw, but are usually dipped in 

egg and fried, or included in an omelet, or used as 

a vegetable ingredient of soups and meat stews. 

These culinary uses are generally unknown in the 

West Indies. 


Islands: Frequently planted in Puerto Rico; in St. 

Croix it has become somewhat naturalized in 

roadside thickets toward the western end of the 

island. 

Common name: Puerto Rico: Bayoneta. 


Estate Thomas Exp. Forest, 3 mi W of 

Christiansted, Little 23037 (US). Virgin Island 

Agric. Exp. Station, Little 21521 (US). 


Yucca gloriosa L., a native of the southeastern 

United States, was grown in Puerto Rico gardens 

and is occasionally adventive on roadsides 

(Britton & P. Wilson, 1923). 

Polianthes tuberosa L., a native of Mexico, is cultivated in Puerto Rico (commonly known as 

Azucena) for floral arrangements and in Virgin Islands gardens. 

Family 16. ASPARAGACEAE Asparagus Family 

Asparagaceae Juss., Gen. Pl. 40. 1789, nom. conserv. 

by P. Acevedo-Rodríguez, M.T. Strong & G. R. Proctor 

Shrubs, sub-shrubs, or vines, usually glabrous with woody or often partly herbaceous, evergreen or 

annually withering branches growing from a rhizome; roots often swollen and fusiform; many species

126 


have green photosynthetic ultimate branchlets, and a few have ultimate branchlets transformed into 

flattened leaf-like phylloclades. Leaves, if present are usually reduced and often minute or scale-like. The 

needle-like “leaves” of commonly grown species are actually much-reduced stems. Flowers small, 

solitary or few to many in racemes or umbels; perianth campanulate with 6 free or fused segments, these 

all similar, bisexual or unisexual (the plants dioecious), inconspicuous, white, yellowish or green in color. 

Stamens 6 with free filaments attached at the base of the perianth; anthers dorsifixed, introrse. Ovary 

superior, 3-locular, with 2-12 ovules per locule. Fruit a globose berry, often red or blue; seeds with black 

epidermis and endosperm lacking starch. An Old World family of more than 300 species mostly of arid 

habitats in Africa, the Mediterranean region, and western Asia; most are tropical or subtropical, but a few 

can grow in temperate areas. 

TYPE: Asparagus L. 

1. ASPARAGUS 

Asparagus L., Sp. Pl. 313. 1753. 

Asparagopsis (Kunth) Kunth, Enum. Pl. 5: 76. 1850, non Mont., 1840. Protasparagus Oberm., S. 

African J. Bot. 2: 243. 1983. 

Erect or scandent and twining shrubs or sub-shrubs with tuberous roots. Leaves absent or reduced to 

tiny scales. Photosynthetic function carried out by verticillate or fasciculate needle-shaped or leaf-shaped 

cladodes. Flowers bisexual, actinomorphic, minute, in racemes or dispersed among the phylloclades; 

perianth of 6 free segments; stamens 6; ovary superior, 3-locular, with 4-10 ovules per locule, and three 

stigmata. Fruit a berry; seeds globose, smooth, black. A genus of about 100 species, naturally distributed 

in the S and N of Africa and Asia. 

Protasparagus Oberm., is here considered part of Asparagus as further analysis by Malcomber & 

Sebsebe Demissew (1992) does not support Obermeyer’s generic concept. 

LECTOTYPE: Asparagus officinalis L., designated by Britton and A. Brown, Ill. Fl. N. U.S. ed. 2. 1: 313. 

1913. 

References: Malcomber, S. T. & S. Demissew. 1992. The status of Protasparagus and Myrsiphyllum 

in the Asparagaceae. Kew Bull. 48: 63-79. Straley, G. B. & F. H. Utech. 2002. Fam. 222. Liliaceae: 

Asparagus. Pp. 213-214. In: Flora of North America Editorial Committee (eds.), Fl. North Amer., vol. 26. 

Key to the species of Asparagus 

1. Cladodes (leaf-like stems) flattened, solitary or few in a fascicle; inflorescence an axillary raceme, 5– 

to 9(–17)-flowered; pedicels 5–8 mm long; berry red ………..................................... 1. A. aethiopicus 

1. Cladodes (leaf-like stems) filiform, arranged in hemispherical to spherical fascicles, numerous in each 

fascicle; inflorescence a terminal umbel, 1- to 4-flowered; pedicels 1–3 mm long; berry purplish black 

……..................................................................................................................................... 2. A. setaceus 

1. Asparagus aethiopicus L., Mant. Pl. 63. 1767; 

Protasparagus aethiopicus (L.) Oberm., S. 

African J. Bot. 2: 243. 1983. Neotype: South 

Africa; Cape. (LINN-434.6), designated by 

Obermeyer, S. African J. Bot. 2: 243. 1983. 

Asparagus sprengeri sensu authors, non Regel, 

1890. 

Asparagus densiflorus sensu authors, non (Kunth) 

Jessop, 1966. 

Scandent sub-shrub, scrambling or climbing 

to 3 m; rhizome short and stout with elongate 

fibrous roots that often form tubers. Stems firm, 

woody, with numerous striate-ridged branches; 

cladodes solitary or in fascicles of 2-5 per node, 

linear, flattened, straight or curved, 8–30 × 1.5-2.5 

mm, with a single prominent or sometimes 

inconspicuous vein; leaves scale-like, membranous, 

1–2 mm long, spurred at base and bearing a spine. 

Inflorescences axillary racemes, 5– to 9(–17)flowered; 

bracts linear, 2-5 mm long. Flower 

pedicels 5–8 mm long, articulate at middle;


perianth rotate to campanulate, the segments 

oblong-ovate, white to pale pink, 3–4 × 1.5–2 mm; 

pedicel. Berries red, 5–10 mm diam.; seeds 1. 

General distribution: Native of South Africa; 

widely cultivated elsewhere and sometimes 

persisting or becoming naturalized. 

Distribution in Puerto Rico: Naturalized at 

several localities. 

Common names: Puerto Rico: Cabellera de la 

reina, Canastillo, Espárrago de jardín. 



Utuado, Cerro Roncador, escaped, 25 Aug 1981 

(fr), Liogier et al. 32198 (UPR). 

2. Asparagus setaceus (Kunth) Jessop, Bothalia 

9: 51. 1966; Asparagopsis setaceus Kunth, 

Enum. Pl. 5: 82. 1850; Protasparagus 

setaceus (Kunth) Oberm., S. African J. Bot. 2: 

244. 1983. Lectotype: South Africa; Cape. 

Drege 8584c (KIEL), designated by Jessop, 

Bothalia 9: 51. 1966. 

Fig. 29. A-E 

Twiner, 2-3 m long. Stems cylindrical, tough, 

green, spiny at the base, much-branched, the 

ultimate branches and cladodes close together in 

flat, frond-like whorls. Cladodes filiform, needlelike, 

numerous in close fascicles. Flowers axillary, 

Fig. 29. A-E. Asparagus setaceus. A. Fruiting branch. B. Base of lateral branch. C. Cladodes. D. Flower, side view, with perianth 

removed, tepal with adnate stamen, and side view. E. Fruit, basal view, showing sepals. From Acevedo-Rdgz., P. 2003, Bejucos 

y plantas trepadoras de Puerto Rico e Islas Vírgenes, Smithsonian Institution.

128 


hanging, solitary or up to 4 together; pedicels 2-2.4 

mm long; perianth segments pale green, oblong, 

obtuse, ca. 2.5 mm long. Berries olive green to 

black, globose, ca. 5 mm in diam.. 

General distribution: Native of South Africa; 

widely cultivated elsewhere and sometimes 

persisting or becoming naturalized. 


Islands: A commonly cultivated plant in Puerto 

Rico and the Virgin Islands. Collected in Maricao 

and San Juan; St. Thomas (no record of being 

naturalized). 

Common names: Puerto Rico: Abeto, Ala de 

Family 17. DRACAENACEAE She-dragon Family 

Dracaenaceae Salisb., Gen. Pl. 73. 1866, nom. conserv. 


pájaro, Helecho plumoso. 


Maricao: Hacienda Juanita, Acevedo-Rdgz. et al. 

7067 (US). San Juan: Botanic Garden, Río 

Piedras, Oquendo 155 (US). 


Asparagus officinalis L., the common 

Asparagus vegetable, is sometimes cultivated in 

Puerto Rico. In addition to this taxon, Britton & P. 

Wilson (1923) listed Asparagus plumosus Baker, a 

native of South Africa, as grown in Virgin Islands 

(St. Thomas: Morrow 87, US) and Puerto Rico 

gardens. 

Trees or shrubs with woody stems or else short-stemmed or acaulescent succulent herbs with fleshy 

creeping rhizomes; woody stems usually with secondary thickening. Leaves entire, lanceolate or linearlanceolate 

to ovate and flat, channeled, cylindrical or laterally compressed, sessile, glabrous, fleshy or 

leathery, rigid or flexible, often clustered at growing tips or in rosettes at the ends of branches. 

Inflorescence an axillary raceme or panicle; flowers bisexual, hypogynous, often fragrant; pedicels 

articulate; perianth tubular or funnel-shaped, the lobes 6, narrow, subequal, spreading or reflexed when 

fully expanded. Stamens 6, inserted at base of the lobes, the anthers dorsifixed, versatile, introrse. Ovary 

superior, 3-locular, with a single or many ovules per locule; style slender, terminated by a 3-lobed or 

capitate stigma. Fruit a fleshy berry or sometimes hard and woody, usually red or orange; seeds 1-3, bony, 

globose or elongate. A Family here construed as having 3 genera and ca. 225 species occurring primarily 

in the Old World tropics, subtropics, and islands of the Pacific Ocean. Representatives of all three genera 

have been introduced into Puerto Rico and the Virgin Islands and some of these have escaped and become 

more or less naturalized. Contrary to the treatment of Dahlgren et al. (1985), Cordyline is here placed in 

the Dracaenaceae. This was done primarily for reasons of convenience but also because Cordyline does 

not fit well into the small, heterogeneous taxon Asteliaceae. 

TYPE: Dracaena L. 


1. Herbaceous plants with creeping rhizomes and rather few, leathery-succulent leaves, these upright or 

in a spreading, sessile rosette; inflorescence of racemes …….............................…. 3. Sansevieria 

1. Shrubby plants with woody, simple or branched, upright stems bearing terminal clusters of thin or 

membranous leaves; inflorescence of panicles. ……….................................................................… 2 

2. Leaves petiolate, flowers scattered separately on panicle branches; ovules 4-12 per 

locule........................................................................................................................ 1. Cordyline 

2. Leaves sessile; flowers in dense, umbel-like clusters; ovules 1 per locule ………. 2. Dracaena


1. CORDYLINE 

Cordyline Comm. ex R. Br., Prodr. 280 1810 , nom. conserv. 

Erect shrubs with woody stems, often stoloniferous at base. Leaves crowded near stem apex, petiolate 

or subsessile. Inflorescence paniculate, with small, scarious bracts; flowers short-pedicellate or 

subsessile, the perianth tubular or narrowly campanulate, the tube short, the lobes oblong, erect or 

spreading; stamens 6, inserted at base of perianth-lobes, the filaments filiform or flattened; ovary sessile, 

3-locular, with 4-12 ovules per locule; style filiform, the stigma capitate or 3-lobed. Fruit a globose, fleshy, 

3-locular, indehiscent berry; seeds numerous, flattened, black and lustrous. A genus of 20 species widely 

distributed in tropical and warm temperate regions, including Indomalaysia, Polynesia, New Zealand, 

Australia, Africa, and Brazil. A single introduced species occurs in Puerto Rico. 

TYPE: Cordyline cannifolia R. Br., typ. conserv. 

1. Cordyline fruticosa (L.) A. Chev., Jard. Bot. 

Saigon 66. 1919; Convallaria fruticosa L., 

Herb. Amb. 16. 1754; Taetsia fruticosa (L.) 

Merr., Interpr. Herb. Amboin. 137. 1917; 

Asparagus terminalis L., Sp. Pl. ed. 2, 450. 

1762, non. Illeg.; Dracaena terminalis L., 

Syst. Nat. ed. 12, 246. 1767, non. Illeg.; 

Cordyline terminalis Kunth, Abh. Königl. 

Acad. Wiss. Berlin 30. 1820, nom. illeg. 

Holotype: Rumphius, Herb. Amboin. 4: t. 34. 

1743. 

Slender unbranched or few-branched shrub 1- 

3 m tall, ringed with scars from fallen leaves. 

Leaves clustered at the apex of stems; petioles 15- 

30 cm long; blades elliptic to linear-lanceolate, 30- 

70 × 10-15 cm, the apex acute to attenuate, the base 

cuneate, the tissue green to deep purple or red or 

2. DRACAENA 

Dracaena L., Syst. Nat. ed. l2, 229, 246. 1767. 

often green with red margins. Panicles terminal, 

densely-flowered; perianth white, purple, or red, 

10-12 mm long, the lobes linear-oblong. Berries 

red or purple. 

General distribution: Native of Malaya, but 

very widely planted in tropical countries. In 

addition to its ornamental qualities, the red-leaved 

forms are often planted as boundary markers or in 

hedgerows, where they tend to persist and become 

naturalized. 


Islands: Cultivated in both, Puerto Rico and the 

Virgin Islands. Recorded from Arecibo (Río 

Abajo State Forest) and San Juan. 

Common name: Puerto Rico: Bayoneta. 



Trees or shrubs with erect woody stems sometimes enlarging to form thick trunks. Leaves entire, 

leathery, closely parallel-veined but with numerous minute cross-veins, crowded along branches or 

clustered at the ends of branches. Inflorescence paniculate; flowers closely aggregated in pseudo-umbels 

at nodes on the branches of the panicle, accompanied by small, scarious bracts; perianth funnelform with 

a short tube and narrow subequal lobes. Stamens 6, borne in the perianth-tube at base of lobes, with terete 

or flattened filaments, Ovary sessile, 3-locular; style filiform with a capitate stigma. Fruit berry-like, 

usually 1-seeded. Mostly an Old World genus of about 150 (Howard, 1979) or 80 (Dahlgren et al. 1985) 

species distributed from Micronesia and subtropical Africa through Asia to northern Australia. A disjunct 

native species grows in eastern Cuba, and another in Central America. A single species has been 

introduced into Puerto Rico; this is commonly cultivated for ornament and sometimes persists or becomes 

naturalized. 

LECTOTYPE: Dracaena draco (L.) L. (≡ Asparagus draco L.), designated by Rafinesque, Fl. Tellur. 4: 

17. 1838.

130 


1. Dracaena fragrans (L.) Ker Gawl., Bot. Mag. 

27: t. 1081. 1808; Aletris fragrans L., Sp. Pl. 

ed. 2, 456. 1762; Pleomele fragrans (L.) 

Salisb., Prodr. Stirp. Chap. Allerton 245. 

1796. Lectotype: Africa. J. Commelijn, Horti 

med. amstelod. 2: 7, t.4., f.2, 1701, designated 

by Wijnands, Bot. Commelins 1: 130. 1983. 

Shrub becoming arborescent to 15 m tall, the 

older woody stems to 30 cm or more in diam., the 

younger stems conspicuously ringed with leaf 

scars. Leaves narrowly elliptic-oblanceolate, 

usually widest above the middle, (12-) 20-125 (- 

150) × (1-) 2-12 cm, rounded or acute at the apex 

with a subulate mucron, narrowly cuneate or 

narrowed slightly to long-decurrent at the base, 

bright green and often variegated above, paler 

green beneath. Inflorescence a terminal panicle 

with few, widely divaricate branches, (15-) 20-100 

(-160) cm long, the main axis often zigzag; bracts 

deciduous, concave, long-cuspidate; flowers 

arranged in pedunculate or sessile glomerules of 

10 or more flowers, each subtended by a triangular 

whitish scarious bract, (15-) 17-22 (-25) mm long, 

yellowish or white, often with a pink stripe, 

intensely fragrant; perianth-lobes (7) 9-12 × 2-3 

mm; stamens inserted at the throat with inflated 

filaments, the anthers 2-2.5 mm long; ovary 

cylindrical to bottle-shaped; style terete, the 

stigma 3-lobed. Fruit red-orange, spherical to 

depressed-globose, 13-20 mm in diam.; seeds 

rounded-rectangular, 6-14 × 5-9 mm. A description 

by Bos et al., 1992, Edinb. J. Bot. 49: 319, was used 

in preparing this description. 

3. SANSEVIERIA 

Sansevieria Thunb., Prodr. Pl. Cap. [xii] 65. 1794, nom. conserv. 

General distribution: Native of Africa; 

widely cultivated in the Neotropics. 

Distribution in Puerto Rico: A species is 

propagated by lopping off the upper part of the 

stems and planting these canes in the ground, 

where they soon produce roots; meanwhile the old 

cane-bases sprout new branches. Recorded from 

Adjuntas, Aibonito, Arecibo, Lares, Río Grande, 

and Utuado, in roadside thickets at elevations of 

200-600 m. 

Common names: Puerto Rico: Drecina, 

Cocomacaco, and Dracena. 

Note: The name Aloe fragrantissima Jacq. 

applies to this taxon, and has priority by three 

months over the name Aletris fragrans L. 

(Dracaena fragrans (L.) Ker Gawl.). Given that 

the Jacquin’s name has hardly been in use, it would 

be appropriate to propose Aletris fragrans L. for 

conservation. The authors are using the Linnean 

name in anticipation of such conservation. 


Adjuntas: Bo. Limaní, S off Rt. 525, on farm of 

Estación Agricola, Axelrod et al. 11365 (UPRRP). 


10758 (US). 


Dracaena draco (L.) L., of the Canary 

Islands, was cultivated at the St. Croix 

Agricultural Experiment Station (Britton & P. 

Wilson, 1923). Dracaena goldieana Bullen ex 

Mast. & T. Moore and D. surculosa Lindl. were 

cited by Martorell et al. (1981) as cultivated in 

Puerto Rico. 

Acaulescent or short-stemmed non-woody plants with creeping rhizomes. Leaves usually rather few, 

often in a rosette, fleshy or coriaceous, terete or flat, spreading or rigid and erect, often variegated. 

Inflorescence terminal, pedunculate, racemose; pedicels articulated, solitary or fascicled. Perianth tubular 

at base, the lobes spreading or recurved, cream-white or greenish white. Stamens 6, exserted, the filaments 

filiform; anthers oblong, bifid at base, 2-lobed at apex ; style long and slender or filiform, about equaling 

the stamens, the stigma capitate. Fruit a 1- to 3-seeded berry, globose to oblong-ellipsoid, with thin 

membranous pericarp; seeds globose with a fleshy testa. A genus of 60 species found in tropical and 

southern Africa, Madagascar, and Arabia; also, many hybrids and horticultural variants exist. The 

classification of these plants is often very difficult. 

TYPE: Sansevieria hyacinthoides (L.) Druce (≡ Aloe hyacinthoides L.), typ. conserv. 

References: Brown, N.E. 1915. Sansevieria. Kew Bull. 1915. 185-261. Wijnands, D.O. 1973. 

Typification and nomenclature of two species of Sansevieria (Agavaceae). Taxon 22: 158-159.


Key to the species of Sansevieria 

1. Leaves cylindrical ……............................................................................................…. 1. S. cylindrica 

1. Leaves flat or half-cylindrical. ............................................................................................................ 2 

2. Leaves half-cylindrical (concave on one side)….....................................……….. 3. S. pearsonii 

2. Leaves flat. ................................................................................................................................. 3 

3. Margins of leaves with a reddish line ………...............................……..2. S. hyacinthoides 

3. Margins of leaves green or white, lacking a reddish line ………………… 4. S. trifasciata 

1. Sansevieria cylindrica Bojer ex Hook., Bot. 

Mag. 85: t. 5093. 1859; Cordyline cylindrica 

(Bojer ex Hook.) Britton in Britton & Wilson, 

Bot. Porto Rico 5: 150. 1923. Type: 

Mauritius. Bot. Mag. 85: t. 5093. 1859. 

Acaulescent plants with stout rhizomes. 

Leaves 3 or 4 together, 2-ranked, stiffly erect, 35- 

100 cm tall or more, cylindrical or slightly 

compressed, mostly 1-2 cm thick, tapering upward 

gradually to a sharp, rigid point. Inflorescence to 1 

m tall; flowers 5 or 6 in a cluster, the perianth white 

or tinted slightly pink, the lobes 2-3 cm long, 

linear, revolute at anthesis. Ripe fruits globose, 7- 

8 mm diam, orange. 

General distribution: Originally endemic to 

Mauritius, now cultivated in most warm countries, 

often persisting or becoming naturalized. 


Islands: No voucher specimens seen from Puerto 

Rico; sight record from Guánica; also seen in St. 

Croix and recorded by Britton & P. Wilson (1923) 

as having been planted in St. Thomas. 

Common name: Puerto Rico: Sansiviera 

redonda. 


Eliza’s Retreat road, SE of Christiansted, along 

road, Proctor & O’Reilly 45235 (US). 

2. Sansevieria hyacinthoides (L.) Druce, Bot. 

Exch. Club Soc. Brit. Isles 3: 423. 1914; Aloe 

hyacinthoides ß guineensis L., Sp. Pl. 321. 

1753; Sansevieria guineensis (L.) Willd., Sp. 

Pl. 2: 159. 1799; Cordyline guineensis (L.) 

Britton, Brooklyn Bot. Gard. Mem;. 1: 35. 

1918; Sansevieria thyrsiflora Petagna, Inst. 

Bot. 3: 643. 1787, nom. illeg. ; Sansevieria 

thyrsiflora Thunb., Prodr. Fl. Cap. 65. 1794, 

nom. illeg. Lectotype: C. Commelijn, 

Praeludia Bot. t. 33. 1703, designated by 

Stearn, Cat. Bot. Books Miller Hunt liii. 1961. 

Sansevieria metallica Gérôme & Labroy, Bull. 

Mus. Hist. Nat. (Paris) 9: 170, 173, fig. 2. 

1903. 

Figs. 30. A-C; 62. E 

Acaulescent plants with stout creeping 

rhizomes. Leaves 2-4 together in a tight cluster, 

stiffly erect, lanceolate or narrowly elliptic, 30- 

100 × 3-9 cm, the apex acute or obtuse, the tissue 

leathery and dull green but mottled transversely 

with numerous more or less obscure pale green 

bands, the margins with a fine reddish line. 

Inflorescence pedunculate, 30-70 cm long, 

bracteate; pedicels to 5 mm long or more; flowers 

with narrow revolute greenish white perianthlobes 

to 1.5 cm long. Fruits globose, shortstipitate, 

red-orange, 8 mm diam.; seeds 7-8 mm 

diam. 

General distribution: Native of South Africa, 

now very widely cultivated, commonly persisting 

and becoming naturalized, weedy, or rampant. 


Islands: Recorded from Arecibo, Barranquitas, 

Ciales, Fajardo, Guánica, Guayanilla, Mayagüez, 

Rincón, San Juan, and Vieques; undoubtedly 

occurs unrecorded in many other municipalities; 

St. Croix, St. Thomas, and St. John, probably also 

in Tortola. This species has several horticultural 

varieties. 

Common names: Puerto Rico: Cocuisa, 

Chucho, Lengua de chucho, Lengua de vaca. 

Note: This species was incorrectly treated in 

Flora of St. John, U.S. Virgin Islands (Acevedo- 

Rdgz., 1996) as Sansevieria trifasciata a species 

not found there. 


Ciales: Camino de la Ceiba, towards Quebrada del 

Pozo Azul, Acevedo-Rdgz. & Vicens 11895 (US). 

Fajardo: “Seven Seas” beach, edge of coastal 

scrub, Acevedo-Rdgz. et al. 14455 (US). 

Guayanilla: Bo. Boca: Punta Verraco, Proctor & 

Haneke 44523 (US). Mayagüez: Landreau s.n.

132 


(US). San Juan: Underwood & Griggs 917 (US). 

ST. CROIX: Bassin, Ricksecker 341 (US). Salt River 

near Morningstar, Fosberg 54061 (US). ST. JOHN: 

Reef Bay Quarter, Bordeaux Mountain, Acevedo- 

Rdgz. & Siaca 3859 (US); Coral Bay Quarter, 

along Bordeaux Road, Acevedo-Rdgz. et al. 5129 

(US). 

3. Sansevieria cf. pearsonii N. E. Br., Bull. Misc. 

Inform. Kew 1911: 97. 1911. Type: Angola. 

Pearson 2073 (K). 

Acaulescent, with a stout creeping rhizome. 

Adult leaves 3-5 to a growth, stiff, erect but 

slightly divergent, the longest to 75 cm long or 

more, 1.5-3 cm thick, cylindrical but with a 

concave channel on the inner face from the base 

nearly to the apex, tapering gradually to the 

attenuate, sharp-pointed tip, the outer surface 

smooth except for a few fine longitudinal grooves, 

bright to dark green with faint lighter horizontal 

bands. Flowers not described. Inflorescence ca. 60 

cm long. Fruits globose, orange, 7-9 mm diam. 

General distribution: Native to Angola. 

Distribution in the Virgin Islands: Known 

only along the Eliza’s Retreat road in St. Croix, 

where it is abundantly naturalized in roadside 

thickets. 

Note: The identity of our St. Croix plants may 

be subject to correction. 

Fig. 30. A-C. Sansevieria hyacinthoides. A. Habit. B. Flower, lateral view, l.s. basal portion showing pistil, and top view. C. 

Anther, frontal and dorsal views. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78.


4. Sansevieria trifasciata Prain, Bengal Pl. 2: 

1054. 1903. Type: India. a cultivated plant, 

Unknown. 

Acaulescent plants with stout creeping 

rhizomes. Leaves 1 or 2 together, linearoblanceolate, 

stiffly erect, 30-100 × 3 cm, the apex 

tapering to a stiff green point, the blades 

transversely banded with contrasting green and 

whitish zones, the margins green. Inflorescence 

pedunculate, 30-75 cm long. Flowers solitary or in 

clusters of 2 or 3, bracteate, pedicels to 5 mm long; 

perianth tube 1 cm long or less, the linear lobes to 

2 cm long, white or greenish white. Fruits 

subglobose to oblong-ellipsoid, 7-9 × 5-8 mm, 

bright orange. 

General distribution: Native of Africa; 

Family 18. ASPHODELACEAE Aloe Family 

Asphodelaceae Juss., Gen. Pl. 51. 1789. 


Perennial often succulent or leathery herbs with rhizomatous roots, or sometimes trees with woody 

trunks. Leaves primarily basal, spirally arranged or sometimes 2-ranked, sheathing at base. Leaves more 

or less fleshy, often internally gelatinous, flat or cylindrical, subulate to linear-lanceolate or elliptic, the 

apex often spine-tipped, the margins smooth, toothed or serrate. Inflorescence terminal or axillary, of 

simple or branched spikes or racemes; scapes leafless or beset with small bracteal leaves. Flowers bisexual 

with 6 perianth segments. Stamens 6, with free filaments, the anthers dorsifixed, introrse, longitudinally 

dehiscent. Ovary 3-locular, with 2 to many ovules in each locule, the placentation axile; style simple, with 

small stigma. Fruit a loculicidal capsule; seeds ovoid or elongate, usually with an aril. A family of about 

17 genera and 814 species, widely distributed in tropical and subtropical regions of the Old World, 

especially common in Africa. In older classifications these plants were considered part of the Liliaceae. 

A single genus, Aloe, has been introduced into Puerto Rico and the Virgin Islands. 

TYPE: Asphodelus L. 

Aloe L., Sp. Pl. 319. 1753. 

1. ALOE 

widely cultivated and often persisting, but less 

common than S. hyacinthoides. 


Islands: Recorded from Ceiba, Dorado, San Juan, 

and Vieques; in the Virgin Islands there are no 

authentic records of naturalized populations, but it 

occurs in cultivation in St. Croix and St. Thomas. 

Note: This species has sometimes been 

confused with S. hyacinthoides, which it 

resembles except for its slightly narrower, more 

brightly green-and-whitish leaves lacking a fine 

reddish line along the margins. According to 

Howard (1979), Jacquin, Hort. Vind, t. 308. 1762 

is a good representation of this species. 



Vieques: Las Marias, Shafer 2705 (US). 

Succulent herbs with basal leaves, or else with a woody caudex more or less well-developed (a few 

species become large branching trees). Leaves in a rosette or rarely distichous, fleshy, with toothed 

margins, containing bitter sap which turns black on exposure or drying. Inflorescence axillary, racemose 

or paniculate, from a naked scapose base; flowers usually nodding; perianth a cylindrical tube, the lobes 

coherent except at the spreading or recurved tips, often brightly colored. Ovary sessile, ovoid to oblong, 

usually 3-angled, the style filiform with a small capitate stigma. Fruit a coriaceous, dehiscent capsule; 

seeds numerous, flattened, black. A large genus of approximately 330 species, the majority occurring in

134 


tropical and southern Africa and Madagascar. 

Many species are cultivated as ornamentals; a 

single one has escaped and become naturalized in 

the West Indies, including Puerto Rico and the 

Virgin Islands. 

Lectotype: Aloe perfoliata L., designated by 

Britton & Millspaugh, Bahamas Fl. 69. 1920. 

1. Aloe vera (L.) Burm. f., F1. Indica 83. 1768; 

A1oe perfoliata var. vera L., Sp. Pl. 320. 

1753. Lectotype: Rheede, Hort. Malab. 11: 

t.3, 1692, designated by Wijnands, Bot. 

Commelins 127. 1983. 

A1oe barbadensis Mill., Gard. Dict. ed. 8, Aloe no 

2. 1768; Aloe vulgaris Lam., Encycl. 1: 86. 

1783, nom. illeg . Type: Unknown, said to be 

from the “American Islands” [West Indies]. 

Figs. 31. A-D; 62. F 

Acaulescent or with a very short, erect, woody 

caudex spreading by stolons to form large 

colonies. Leaves ascending to erect, in a rosette, 

sessile, narrowly deltate, 30-50 × 5-6 cm, longacuminate 

at the apex, the margins with remote 

teeth, the tissue glaucous, light green. Inflorescence 

a dense raceme, scapose at base; flowers yellow, 

with very short pedicels; perianth to 2.5 cm long, 

equaled or slightly exceeded by the stamens; style 

exserted. Fruits apparently not described, seldom 

produced. 

General distribution: Thought to be native to 

the Mediterranean region, but now widely 

cultivated and extensively naturalized. This 

species has important medicinal and cosmetic 

properties. 


Islands: Widely cultivated and naturalized; 

recorded from Guánica, Mona Island, Salinas, and 

Fig. 31. A-D. Aloe vera. A. Habit. B. Inflorescence. C. Flower. D. Stamen and pistil. From Acevedo-Rdgz., P. 1996, Flora of St. 

John, Mem. New York Bot. Gard. 78.


Vieques; it undoubtedly grows in many more sites, 

especially in the dry southern districts; St. Croix, 

St. Thomas, St. John, Tortola, and Virgin Gorda. 

Common names: Puerto Rico: Sábila, Zábila; 

Virgin Islands: Aloe. 

Note: The type of Aloe vera pertains to the 

original publication by Rheede, since the facsimile 

edition has instead a wrong illustration. The 

illustration presented there is a duplicate of t. 35, 

where the number 5 has faded off of the plate 

number 35. 


Salinas: Ensenada, Liogier & Martorell 31602 

(US). Vieques: Brigadier Point to Puerto Negro, 


Shafer 2922 (US). ST. CROIX: South Shore, 

Ricksecker 309 (US). ST. JOHN: East End Quarter; 

Center Line Road, Acevedo-Rdgz. & Siaca 4223 

(MO, NY, UPR, US). ST. THOMAS: Morrow 131 

(US). 


The below listed taxa of Aloe, mostly from 

southern Africa, were listed by Britton & P. 

Wilson (1923) as cultivated on St. Thomas: Aloe 

arborescens Mill., A. ciliaris Haw., A. ferox Mill., 

A. grandidentata Salm-Dyck, A. greenii Baker, 

and A. obscura Mill. 

Haworthia tessellata Haw., from Africa, was cultivated in St. Thomas at Louisenhoj (Britton & P. 

Wilson, 1923). 

Family 19. ARECACEAE (PALMAE) Palm Family 

Arecaceae Schultz, Nat. Syst. Pflanzenr. 317. 1832; nom alt. Palmae Juss., Gen. Pl. 37. 1789, nom. 

conserv. 


Trees, shrubs, or vines with thick or fine fibrous roots, perennial or sometimes monocarpic. Stems 

slender to massive, solitary, clumped, or stoloniferous, rarely branched, ringed with leaf-scars and smooth 

internodes, or rough with persistent petiole-bases or sometimes beset with long sharp spines. Leaves 

spirally arranged, with sheathing, tubular, armed, or unarmed, deciduous or persistent petioles; blades 

simple and linear-lanceolate, or pinnately or rarely bipinnately compound, or palmately or costal 

palmately divided; leaflets and segments linear-elliptic, falcate, or fan-shaped, the tissue glabrous, scaly, 

hairy, prickly, or waxy at least during development; stipules absent, but petiolar sheath sometimes 

prolonged as a ligule at base of blade. Flowers 3-merous, usually actinomorphic and small, unisexual 

(plant monoecious, dioecious, or polygamous), or less often bisexual, in axillary (or less often terminal) 

spikes, racemes, or panicles, the inflorescences subtended by a prophyll or spathe, in position infra-, interor 

suprapetiolar, and sessile or pedunculate. Individual flowers sessile or pedicellate, solitary or clustered 

in triads of 2 staminate on either side of 1 pistillate flower. Perianth tepals biseriate (or in Thrinax and 

allies, a single usually irregularly 6-lobed series), persistent in fruit. Stamens 3 to 200 or more; filaments 

free, connate, or epipetalous; anthers 2- (rarely 1-) locular, opening by a longitudinal slit. Ovary superior, 

of 3 (or rarely more), 1-ovulate carpels (1-carpellate in Thrinax and allies); carpels free or variously 

united: styles free or fused, or stigmas sessile. Fruits small to very large, 1- to 3-seeded (seeds rarely more), 

dry and fibrous or berry-like with smooth, scaly, prickly, hairy, or warty exocarp, fibrous or fleshy 

mesocarp, and papery to bony endocarp. Seeds with solid, hollow, perforated, homogeneous or ruminate 

endosperm, and a subapical, lateral, or basal, embryo. A large, diverse, economically important family of 

212 genera and more than 2600 species, distributed worldwide in tropical and subtropical regions, also a 

few extending into warm-temperate areas. The flora area has 11 genera, each genus represented by a single

136 


species. This does not take into account numerous species from many parts of the world, introduced 

primarily for ornament. Apparently none of these has escaped or become naturalized, and therefore 

inclusion in this book would not be appropriate. They deserve a separated illustrated publication. 

TYPE: Areca L. 

References: Cook, O. F. 1901. A synopsis of the palms of Puerto Rico. Bull. Torrey Bot. Club 28: 

525-560. Moore, H. E. & J. Dransfield. 1979. The typification of Linnean Palms. Taxon 28: 59-70. Read, 

R. W. 1979. Palmae. 320-368. In: R. A. Howard (ed.), Fl. Lesser Antill. 3: 320-388. Uhl, N. W. & J. 

Dransfield. 1987. Genera Palmarum. Allen Press, Lawrence, Kansas. 


1. Leaves palmate (with or without a central rib). …......................................................................…….. 2 

2. Leaves costa-palmate, the mid rib continuous with the petiole, extending through the length of the 

blade and strongly recurved; inflorescence very large and much branched ……………. 10. Sabal 

2. Leaves palmate, without a prominent mid rib (a short hastula present); inflorescence slender, erect 

or arching, with short secondary branches …………............................................................….. 3 

3. Leaf sheath split below the insertion of the petiole; ripe fruits white; seeds smooth, semiperforated 

on one side …………………............................................................ 11. Thrinax 

3. Leaf-sheath not split below the insertion of the petiole, the tubular portion net-like and 

persistent; ripe fruits black or purple-black; seeds with grooved surface, separating into 

several divisions when dry ………....................................................……. 4. Coccothrinax 

1. Leaves pinnate. …................................................................................................................………….. 4 

4. Spines present on leaves, petioles, spathes, or trunk, or on all of these. ………...................…. 5 

5. Leaf-segments with apices broad and erose; ripe fruits red or scarlet …………… 2. Aiphanes 

5. Leaf-segments with apices narrow and sharply pointed, or sometimes bifid; ripe fruits 

yellowish green or pale orange, not red or scarlet ………......................……… 1. Acrocomia 

4. Spines absent, plants unarmed. ……...........................................................................………… 6 

6. Crownshaft present; inflorescences infrapetiolar, attached at base of crownshaft. ……….. 7 

7. Crownshaft split opposite the petioles; mature trunks usually less than 20 cm in diam.; 

primary bracts not filled with loose scales: seedling leaves bifid ………… 7. Prestoea 

7. Crownshaft not split; mature trunks more than 40 cm in diam.; primary bracts filled with 

loose scales when young; seedling leaves lanceolate, not bifid ………… 9. Roystonea 

6. Crownshaft absent; inflorescences interpetiolar. ……………......................................……. 8 

8. Leaf-segments with several prominent longitudinal nerves; flowers sunk in pits; petals 

united, falling like a cap …………..............................................…… 3. Calyptronoma 

8. Leaf-segments with a single prominent longitudinal nerve; flowers not sunk in pits; 

petals free, not forming a cap. …………............................................................…….. 9 

9. Trunk smooth, with very faint annular rings, strongly tapering from a swollen base 

to a slender apex 5-9 cm in diam.; leaves less than 1.8 m long ………….. 6. Gaussia 

9. Trunk ringed with more or less conspicuous annular leaf-scars, and more than 15 cm 

in diam. at apex; leaves more than 2 m long. ………................................……… 10 

10. Small trees with straight trunks not more than 8 m tall; leaves 1.5-3 m long; 

flowers all alike, bisexual; ripe fruits red, up to 1.7 cm in diam. 

……..........................................................................………. 8. Pseudophoenix 

10. Tree up to 30 m tall, the trunk often curved or bent; leaves 3-6 m long; flowers 

unisexual, the inflorescences monoecious, fruits more than 20 cm long 

…...........................................................................................……….. 5. Cocos


1. ACROCOMIA 

Acrocomia Mart., Hist. Nat. Palm. 2: 66. 1824. 

Large solitary monoecious palms armed with sharp spines; trunk columnar, often variously swollen, 

fusiform or ventricose; usually heavily armed at first with annular rows of spines, these commonly 

deciduous with age. Leaves pinnate; sheaths spiny; petioles very short, hardly evident; rachis armed with 

large flat spines; segments numerous, narrowly lanceolate, acuminate or bifid, inserted singly or in 

clusters in more than one plane except toward apex of leaf, the segments spreading or arching at different 

angles, usually glabrous and glossy green on upper side, often glaucous-green and pubescent or pilose on 

the under side. Inflorescences paniculate, interpetiolar, once-branched, the peduncle stout, arching, 

usually densely scaly, and spiny, the first and second nodes with large and very unequal primary bracts, 

the first bract much shorter than the second, quickly withering, the second bract woody and scurfy or 

brown-tomentose, variously armed, crooked basally, and pitted toward the apex. Flowers unisexual; 

staminate flowers much smaller than the pistillate, inserted in pits and mostly crowded on the apical half 

of the inflorescence branches, the much larger and fewer pistillate flowers produced only along the 

flexuous basal portion of the same branch, sometimes accompanied by solitary staminate flowers or these 

close on either side of a pistillate flower, forming a triad. Sepals of staminate flowers free, much shorter 

than the petals; petals connate basally, valvate apically; stamens 6, filaments slender, inflexed in bud, 

basally adnate to the petals; pistillode prominent, tufted. Pistillate flowers with free, imbricate sepals; 

petals free, imbricate except toward apex; staminodes represented by a broad, minutely denticulate 

cupule; ovary 3-pistillate, variously scaly or pubescent; stigmas sessile, recurved, the stigmatic remains 

apical and conspicuous on fruit. Fruits depressed-globose, 1-seeded, olive-green or yellowish; exocarp 

chartaceous; mesocarp fibrous and mucilaginous; endocarp bony, 3-porate; endosperm homogeneous, 

oily, edible. A widespread Neotropical genus with ca. 30 taxa. However, Henderson (1994: The Palms of 

the Amazon, Oxford University Press, New York) lumped all the recognized taxa into just two species, A. 

aculeata (Jacq.) Lodd. and A. hassleri (Barbosa Rodrigues) W. J. Hahn. Puerto Rico has a single species, 

here considered endemic. 

TYPE: Acrocomia aculeata (Jacq.) Lodd. (≡ Cocos aculeata Jacq.) . 

1. Acrocomia media O. F. Cook, Bull. Torrey 

Bot. Club 28: 566. 1901. Type: Puerto Rico; 

Ponce. Cook s. n. (US). 

Acrocomia sclerocarpa sensu Bello, Anales Soc. 

Esp. Hist. Nat. 12: 114. 1883, non Martius, 

1824. 

Acrocomia aculeata sensu Britton & P. Wilson, 

Bot. Porto Rico 5: 115. 1923, non Cocos 

aculeata Jacquin, 1763. 

Stout solitary palm, the trunk cylindrical or 

slightly fusiform, becoming 8-10(15) m tall, 20-30 

cm in diam., densely armed with internodal rings 

of long black spines, these eventually falling, 

leaving a naked gray trunk smooth except for leaf 

scars. Leaves numerous, the short petioles densely 

spiny the blades 3-4 m long with 45-60 

multifarious linear segments, the longer ones 55- 

70 × 0.6-2 cm, acuminate or attenuate at apex. 

Inner spathe (second primary bract) up to 60 cm 

long; densely brown-pubescent and somewhat 

spiny on outer surface, fusiform at first, becoming 

hood-shaped after opening. Panicles up to 1.5 m 

long, with short thick densely spiny peduncles and 

numerous slender branches 10-25 cm long, the 

upper ones wholly staminate, the lower ones 

staminate on apical two-thirds, the basal flexuous 

portion bearing 3-5 distant pistillate flowers. 

Staminate flowers densely borne in individual pits 

in the rhachilla, each flower ca. 8 mm long, pale 

yellow at anthesis; pistillate flowers sub-globose, 

ca. 10 mm long. Fruits depressed-globose, 3.5-4.5 

cm in diam., smooth, dull yellowish at maturity. 


and the Virgin Islands. 


Islands: Occurs in moderately dry to moist 

habitats at mostly elevations below 200 m. Often 

transplanted as a street-side ornamental. Recorded 

from Bayamón, Carolina, Humacao, Manatí, 

Peñuelas, Ponce, Sabana Grande, Salinas, San 

Juan, Toa Baja, Utuado, and Vega Baja; St. Croix 

and St. Thomas. Probably distributed more widely 

than records indicate.

138 


Common names: Puerto Rico: Coroso, Palma 

de coroso. 

Note: Acrocomia media was treated as a 

synonym of A. aculeata by Britton & P. Wilson 

(1923). Unfortunately, there is no recent 

monograph of the genus except Bailey’s 

preliminary treatment of 1941, so the relationships 

among the Greater and Lesser Antilles populations 

cannot be clearly assessed. Acrocomia media is 

probably closest morphologically as well as 

geographically to A. aculeata of the Lesser 

Antilles; according to available information it 

differs from the latter species in its longer leaves 

with narrower segments, its much shorter spathes, 

and in its slightly larger flowers. Other 

differences, such as the length of spines and 

character of indument, also exist. It should be 

noted that these comparisons are based on 

measurements of very few specimens, so that 

currently the limits of variation are unknown. It is 

an unfortunate fact that large, spiny palms do not 

invite close examination. 


Bayamón: Mogotes near Bayamón, Read 2061 

(US-4). Manatí: Bo. Tierras Nuevas Saliente, 

Proctor & Thomas 42235a (US). Peñuelas: 

between Peñuelas and Tallaboa Alta, Sintenis 

4935 (US). Sabana Grande: Sintenis 3972 (US). 


Utuado: Sintenis 6483 (US). ST. THOMAS: St. Peter, 

E.G. Britton & Marble 1210 (US). 

2. AIPHANES 

Aiphanes Willd., Samml. Deutch. Abh. Königl. Akad. Wiss. Berlin 1803: 250. 1806. 

Small to medium, solitary monoecious understory palms; trunks with internodal spines, these 

eventually deciduous. Leaves pinnate, the sheaths soon splitting into shreds, usually densely spiny; petiole 

also usually densely spiny; rachis convex beneath, ridged on upper side, often variously armed with 

spines, glabrescent to densely matted-lepidote; segments numerous, in 1 or 2 planes, usually broadest 

toward outer end, the apices erose-truncate, dark green, armed or not on mature plants, but usually densely 

prickly on one or both sides of leaves on juvenile plants. Inflorescences interpetiolar, usually oncebranched; 

peduncles usually densely armed with long black spines, elongate, arching or pendent, the first 

and second nodes producing strongly unequal primary bracts, the first bract relatively smaller and 

unarmed, the second bract narrow and often densely spiny; rachis often spiny toward base and more or less 

lepidote with tiny appressed scales; branches sinuous, spreading. Flowers usually in triads of one pistillate 

and two staminate, the entire triad often sunk in a depression on the branch; paired or solitary staminate 

flowers may also occur; flower clusters usually subtended by 2 or 3 minute bracts in addition to the pit 

bract. Staminate flowers often stipitate, usually also pedicellate; sepals connate at base; petals mostly free 

and valvate; stamens 6, filaments straight, adnate to the petals basally; anthers versatile, sagittate basally, 

retuse at apex; pistillode conical to globular and trifid. Pistillate flowers sessile with free, basally imbricate 

sepals; petals about half-connate; staminodial cup usually adnate to the petals; ovary globose to pyriform, 

3-pistillate; stigmas very short; stigmas 3. Fruit globose, 1-seeded, red or scarlet at maturity; mesocarp 

yellow; endocarp bony, variously sculptured, with 3 pores usually equatorial in position. Seeds irregular 

in form, the embryo opposite one of the endocarp pores; endosperm homogeneous. Seedling leaves more 

or less bifid, conspicuously spiny. A Neotropical genus of about 30 species (Read, 1979) or 22 species 

(Borchsenius & Bernal, 1996), best represented in Colombia and Ecuador. A single species is found in 

Puerto Rico, known elsewhere only in the Dominican Republic. Lesser Antillean populations combined 

with the Puerto Rican taxon by Borchsenius & Bernal represent several different species in the present 

writer’s opinion. 

TYPE: Aiphanes aculeata Willd. 

Reference: Borchsenius, F. & R. Bernal. 1996 Aiphanes (Palmae). Fl. Neotrop. Monogr. 70: 1-95. 

1. Aiphanes acanthophylla (Mart.) Burret, 

Notizbl. Bot. Gart. Berlin-Dahlem 11: 558. 

1932; Bactris acanthophylla Mart., Palm. 

Orbig. 70. 1844. Martinezia acanthophylla 

(Mart.) Becc. in Urban, Symb. Antill. 8: 79.


1920. Type: Puerto Rico. Wydler 192 

(holotype: FI, photo at AAU; isotype: G). 

Curima colophylla O. F. Cook, Bull. Torrey Bot. 

Club 28: 561. 1901. Type: Puerto Rico. 

Underwood & Griggs 878 (US-3 sheets!). 

Bactris pavoniana sensu Bello, Anales Soc. Esp. 

Hist. Nat. 12: 114. 1883, non Martius, 1824. 

Solitary palm with relatively slender trunk, to 

18 m tall and 20 cm in diam., armed with numerous 

blackish spines 4-7 cm long especially when 

young, these eventually deciduous. Leaves dark 

green, up to 2.5 m long, pinnatisect, the majority of 

the numerous segments linear or nearly so, in one 

plane, 3-6 cm broad, at apex obliquely erosetruncate; 

terminal leaf-segments broad, pinnately 

veined, deeply notched and irregularly shortlacerate. 

All parts of leaf variably armed with 

sharp blackish spines, or mature leaves spineless. 

Inner spathe (second primary bract) up to 1 m long, 

densely spiny; inflorescence usually more than 60 

cm long; flowers greenish white, ca. 3 mm long. 

Fruits globose, bright red, ca. 1.5 cm in diam. 

General distribution: Confined to Puerto 

Rico and the Dominican Republic. 

Distribution in Puerto Rico: Occurs as an 

understory tree in forest over limestone (rarely 

serpentine) at low to middle elevations (50-400 

m), uncommon or locally frequent. Recorded from 

Arecibo, Bayamón, Camuy, Cidra, Corozal, 

3. CALYPTRONOMA 

Calyptronoma Griseb., Fl. Brit. W. I. 518. 1864. 

Dorado, Fajardo, Juncos, Maricao, Quebradillas, 

San Juan, San Lorenzo, Toa Alta, Toa Baja, 

Utuado, and Vega Baja. 

Common name: Puerto Rico: Palma de coyor. 

Note: Borchsenius & Bernal (1996) consider 

the correct name for this species to be Aiphanes 

minima (Gaertn.) Burret. However, the typification 

of this name is based on a single endocarp (fruit) 

from an unknown locality in the seed collection of 

Gaertner, collected more than 200 years ago. If the 

epithet “minima” is applied in a broad sense (as did 

Borchsenius & Bernal) to the Antillean populations 

which may represent several species (as the writer 

believes), then “minima” will have to be epi- or 

neotypified so that confusion will be avoided. 


Bayamón: Candelaria, near Bayamón, N.L. 

Britton et al. 2857 (US). Cidra: Pueblo Viejo, 

Stevenson 2105 (US). Corozal: N.L. Britton & 

E.G. Britton 7829 (US). Dorado: on the grounds of 

the Dorado Beach Country Club, Read & 

Woodbury 2055a (US-4). Fajardo: Río Arriba, 

N.L. Britton & Shafer 1701 (US). Juncos: Sintenis 

2500 (US). Maricao: Sintenis 484 (US). San 

Lorenzo: Monte Gregorio, Sintenis 2628 (US). 

Toa Alta: N.L. Britton & E.G. Britton 7854 (US). 

Toa Baja: Bo. Candelaria, wooded mogotes 0.4- 

0.7 km WSW of Pico Nevárez, Proctor & Rivera 

45638 (US-4). Utuado: E.G. Britton & Marble 396 

(US-2). Vega Baja: Goll 1044 (US). 

Solitary, unarmed, monoecious palms, the trunk gray-brown and marked with prominent leaf-scars. 

Leaves pinnatisect; crownshaft lacking; petioles short, less than 50 cm long, clasping at base and with 

fibrous margins; segments linear-lanceolate, borne in one plane, the apices acute; minute chaffy scales 

borne abaxially on major veins. Inflorescences interfoliar, solitary in the axil of each leaf, with 3 (rarely 

4) orders of branching; peduncular bract erect, coriaceous, splitting longitudinally on the abaxial side, 

attached near base of peduncle. Panicle branches borne in clusters on short stalks basally, but solitary 

distally, each subtended by a short bract and covered when young by arachnoid tomentum. Flowers borne 

in triads sunk in pits closed at first by pit bracts. Staminate flowers sessile; sepals 3, imbricate, unequal, 

linear-elliptic, keeled, with denticulate, hyaline margins; petals 3, basally adnate, to the staminal column 

and more or less connate, valvate or joined and calyptrate at apex; staminal tube funnel-form, fleshy, 

white, bearing 6 short, narrowly triangular filaments at its apex; anthers 6, sagittate, with introrse 

dehiscence; pistillode minute. Pistillate flowers sessile; sepals 3, linear-elliptic; petals 3, membranous, 

connate, opening by circumcissile dehiscence, falling together like a cap; staminode tubular-cupulate, 

inflated at apex; ovary superior, of 3 fused carpels; style slender, apically attached to the 3-lobed ovary. 

Fruit an obovoid drupe with a smooth epicarp, fleshy mesocarp and crustaceous endocarp; seed spheroid, 

shiny brown, with conspicuous encircling unbranched raphe and obscure hilum; embryo basal; endosperm

140 


homogeneous. A Greater Antillean genus of 3 species, one of them occurring in Puerto Rico. 

TYPE: Calyptronoma occidentalis (Sw.) H. E. Moore. (≡ Elaeis occidentalis Sw.). 

Reference: Zona, S. 1995. A revision of Calyptronoma (Arecaceae). Principes 39: 140-151. 

1. Calyptronoma rivalis (O. F. Cook) L. H. 

Bailey, Gentes Herb. 4: 171. 1938; Cocops 

rivalis O. F. Cook, Bull. Torrey Bot. Club 28: 

568. 1901; Calyptrogyne rivalis (O. F. Cook) 

León, Contr. Ocas. Mus. Hist. Nat. Colegio 

“De La Salle” 3: 12. 1944. Lectotype: Puerto 

Rico. Underwood & Griggs 89 (US!), 

designated by Zona, Principes 39: 149. 1995. 

Calyptronoma quisqueyana L. H. Bailey, Gentes 

Herb. 4: 168. 1938; Calyptrogyne quisqueyana 

(L. H. Bailey) León, Contr. Ocas. Mus. Hist. 

Nat. Colegio “De La Salle” 3: 12. 1944. 

Lectotype: Haiti. Bailey 729 (BH), designated 

by Zona, Principes 39: 149. 1995. 

Calyptrogyne occidentalis sensu Britton & P. 


(Swartz) M. Gómez, 1893. 

Trunk to 15 m tall, 18-25 cm in diam.; leaves 

3.1-5 m long, the segments 54-× 3-6 cm, glabrous 

on the lower side between the veins. Peduncular 

bract 66-77 cm long, ca. 8.5 cm broad. Peduncle 

39-77 cm long, 1.3-3 cm thick; rhachilla borne in 

clusters with 6(7) rows of floral pits, the proximal 

rachillae borne in clusters of 3-5 (-7) on stalks 1.4- 

4. COCCOTHRINAX 

Coccothrinax Sarg., Bot. Gaz. 27: 87. 1899. 

5.2 cm long; clustered rachillae borne for ½ to 2 /3 

the length of the rachis; floral pits 2.4-3.5. 

Staminate flowers 4.8-5.1 cm long; sepals 3-3.6 

mm long; petals completely connate; staminal 

tube 4.1-4.5 mm long; filaments 0.8-1.2 mm long; 

anthers 1.6-1.7 mm long. Pistillate flowers not yet 

described. Fruits 4.6-7 × 3.6-4.6 mm in diam.; seed 

3.8-4.9 × 2.9-3.5 mm in diam. 

General distribution: Puerto Rico and 

Hispaniola. 


Camuy (along the Río Camuy), Quebradillas 

(along the Río Guajataca), and San Sebastián 

(along stream below Collazo Falls). The largest 

number of wild trees occurs along the Río Camuy. 

This species grows naturally only along the banks 

of streams and rivers at low elevations (35-150 m). 

However, it has been propagated extensively by 

seeds and has been planted rather widely. 

Common name: Puerto Rico: Manaca. 


Camuy: Sintenis 6061 (US). Lares to San 

Sebastián, N.L. Britton & Hess 2785 (US-4); 

Liogier 28727 (US-2). 

Small to medium solitary (rarely clustered), mostly unarmed palms with slender columnar trunks 

these nearly smooth or obscurely ringed with leaf-scars; base of trunk often arising from a mass of exposed 

roots. Leaves palmate with fibrous tubular sheaths at base, these unarmed or sometimes spiny, becoming 

loosely mat-like to very hard like grillwork; petioles flat to convex on upper side and convex beneath, 

terminating at the blade in a hastula, solid throughout and not splitting at base within the sheath; hastula 

conspicuous on upper surface, beneath marked by a small ridge, flap, or projection. Leaf blade entirely 

flabellate, lacking a central costa, pleated during development; upper surface glabrous, the lower surface 

variously lepidote with minute appressed scales; segments variously fused toward base, the whole blade 

remaining somewhat pleated at maturity; segment apices bifid. Inflorescences interfoliar, elongate, erect 

to arching, mostly shorter than the leaves, with a few to many pendulous primary branches, these mostly 

bearing simple secondary branches; lowermost primary bract bicarinate, the others tubular with an oblique 

aperture; all primary bracts appressed-lepidote, each enclosing the base of a primary branch (rhachilla), 

each ultimate branch subtended by a narrow triangular bract. Flowers bisexual, mostly protandrous, 

solitary on short or prominent bracteolate pedicels; perianth a 6-lobed or -dentate cupule. Stamens usually 

9-12 with straight slender filaments, these free to slightly connate at base; anthers basifixed, with oblong 

to linear locules dehiscing by lateral slits; ovary unilocular with a single ovule; style very short, flaring 

upwardly to a laterally-compressed stigma. Fruit small, depressed-globose, black or dark purple, smooth


except for apical stigma remnant and adherent base of perianth cupule; mesocarp thin, juicy; endocarp 

thin, membranous; seed globose, deeply grooved or brain-like; endosperm more or less ruminate by 

deeply folded or irregular lobes. A circum-Caribbean genus of 20 or more species, occurring on most of 

the islands and adjacent continental areas. One species is found in Puerto Rico and the Virgin Islands. 

TYPE: Coccothrinax jucunda Sarg. 

1. Coccothrinax alta (O. F. Cook) Becc., Webbia 

2: 331. 1907; Thrincoma alta O. F. Cook, 

Bull. Torrey Bot. Club 28: 540. 1901. Type: 

Puerto Rico. Underwood & Griggs 848 


Thringis latifrons O. F. Cook, Bull. Torrey Bot. 

Club 28: 545. 1901; Coccothrinax latifrons 

(O. F. Cook) Becc., Webbia 2: 326. 1907. 

Type. Puerto Rico; Coamo. Sintenis 3278 


Thringis laxa O. F. Cook, Bull. Torrey Bot. Club 

28: 545. 1901. Type. Puerto Rico; Vega Baja. 

Cook & Collins 1041 (US!). 

Coccothrinax eggersiana Becc., Webbia 2: 321. 

1907. Type: St. John; U.S. Virgin Is. Eggers 

3117 (holotype: B; isotype: C). 

Coccothrinax eggersiana var. sanctae-crucis 

Becc., Webbia 2: 323. 1907. Type: St. Croix; 

U.S. Virgin Is. Benzon 75-1759 (holotype: C). 

Coccothrinax sanctae-thomae Becc., Webbia 2: 

303. 1907. Type: St. Thomas; U.S. Virgin Is. 

Børgesen s.n. (holotype: C). 

Thrinax argentea sensu Eggers, Fl. St. Croix 100. 

1879, non Loddiges, 1830. 

Coccothrinax argentea sensu Britton & P. Wilson, 

Bot. Porto Rico 5: 117. 1923, non (Loddiges) 

Sargent, 1902. 

Fig. 32. A-D 

Erect solitary palm 2-6 (-11) m tall; trunk 

cylindrical, (5-) 8-12 cm in diam., slightly tapering 

toward apex. Leaves with orbicular blades up to 75 

cm long, silvery-lepidote beneath, cleft beyond the 

middle into numerous narrowly lanceolate, 

acuminate segments; petioles about as long as the 

blades, the base expanded in a persistent, tubular, 

fibrous, sheath surrounding top of trunk below the 

growing point. Inflorescence erect or ascending, 

paniculate, solitary in leaf axils, with several 

pendulous primary branches, the rachillae glabrous. 

Flowers light yellow, on pedicels 1-3 mm long; 

perianth 2.5-3 mm wide; stamens usually 9. Fruits 

globose to depressed-globose, 5-6 mm in diam., 

purple-black when ripe. 




Islands: This species is found at low to lower 

middle elevations (near sea level to 350 m), chiefly 

along the northern side of Puerto Rico, where it is 

confined to a limestone substrate, and in the 

various islands east of Puerto Rico, where it 

chiefly grows over igneous rocks. It probably 

grows on many of the smaller Virgin Islands not 

listed here. Recorded from Arecibo, Barceloneta, 

Coamo, Culebra, Dorado, Florida, Isabela, 

Manatí, Quebradillas, Toa Baja, Utuado, Vega 

Alta, Vega Baja, and Vieques; Guana Island, St. 

Croix (very rare), St. John, St. Thomas, Tortola, 


Common names: Puerto Rico: Palma plateada, 

Palma de abanico. 

Note: It has been suggested (Read, 1979, p. 

329) that Coccothrinax alta may not be 

distinguishable from C. barbadensis (Lodd. ex 

Mart.) Becc. of the Lesser Antilles. However, C. 

alta is here maintained as a separate species 

because of its (on the average) shorter, more 

slender trunk, fewer stamens, and much smaller 

fruits (5-6 mm diam. vs. 7-12 mm diam.). 


Arecibo: Bo. Río Arriba, Axelrod 9404 (US). 

Coamo: On Monte Calabaza near Coamo, Sintenis 

3278 (US). Culebra: N.L. Britton & Wheeler 52 

(US-2). Vieques: Little & Woodbury 23733 (US). 

Florida: Bo. Florida Adentro, 0.5 km along dirt 

road E of old quarry at Rt. 140, km 50.75, Axelrod 

et al. 4668 (US-2). Vieques: Isabel Segunda to 

Santa Maria, Shafer 2610 (US-2). ST. CROIX: 

Cotton Valley, Little 16406 (US). ST. JOHN: Coral 

Bay Quarter; along dirt road to Bordeaux 

Mountain, Acevedo-Rdgz. et al. 5089 (NY, 

UPRRP, US-3). ST. THOMAS: Flag Hill, Shafer & 

Fitch 1470 (US-2); St. Peter, E.G. Britton & 

Marble 1209 (US); Water Island, E.G. Britton & 

Marble 1209 (US-2). TORTOLA: Guana Island, 

Palm Ghut, SE of North Beach, Proctor 41996 

(US-2). VIRGIN GORDA: Little & Woodbury 23847 

(US-2).

142 

Cocos L., Sp. Pl. 1188. 1753. 


5. COCOS 

A monospecific genus believed to have originated somewhere in the region of Malaya or the 

Philippines Islands, now planted and spontaneous or naturalized in all tropical countries and some 

subtropical areas, especially along sandy seacoasts, characterized by the following species. 

TYPE: Cocos nucifera L. 

1. Cocos nucifera L., Sp. Pl. 1188. 1753. 

Lectotype: Malabar Is. Rheede, Hort. Malab. 

1: t. 1-4, 1678, designated by H.E. Moore & 

Dransfield, Taxon 28: 64. 1979. 

Medium to large, solitary, unarmed 

monoecious palm with arching pinnate leaves, 18- 

30 m tall, the often slanted or curved trunk 40-60 

cm in diam., nearly smooth but ringed with broad 

leaf-scars and usually somewhat enlarged at base. 

Leaves arching, the petioles with fibrous netlike 

sheath that pulls apart from the petiole base as the 

leaf matures; petioles heavy, very short; blades 3- 

6 m long, 1 m or more broad, with 75-90 narrowly 

lanceolate segments on each side, these 50-70 cm 

long, 3.5-5 cm broad, all inserted regularly in one 

plane on the rachis. Expanded portion of inner 

inflorescence bract 90-120 cm long and 15-20 cm 

broad near the middle, somewhat glaucous. 

Inflorescence interfoliar, long-pedunculate, 

protandrous, mostly 1-2 m long, once-branched; 

primary bracts 2, persistent, very unequal in size, 

the outer bract bicarinate, truncate, the inner bract 

much larger, fusiform in bud, plicate-grooved, 

splitting abaxially; rachis and rachillae glabrescent; 

rachillae numerous (more than 100), each 30-40 

cm long. Flowers unisexual, both sexes borne on 

the same inflorescence (rarely an entire 

inflorescence staminate), in 1-2 triads of two 

staminate and one pistillate flowers at base of each 

rhachilla, with single or paired staminate flowers 

above; pistillate flowers much larger than the 

staminate ones. Staminate flowers narrowly 

ovoid, somewhat asymmetrical, 1-1.5 cm long, 

sessile or pedicellate, the petals valvate and much 

longer than the sepals; sepals 2 mm long, free, 

imbricate at base; stamens 6; pistillode prominent, 

trifid. Pistillate flowers globose-conic, 2.5 cm 

long; sepals and petals similar, strongly imbricate; 

ovary 3-pistillate, with only a single carpel 

normally fertile, entire, rugose toward apex; 

stigmas 3, sessile. Fruits ovoid or ellipsoid, bluntly 

3-angled, 20-30 cm long; exocarp smooth, green to 

yellow or orange; mesocarp thick and densely 

fibrous, 2-4 cm thick; endocarp bony and hard, 

with 3 large opercula toward one end; normally 

only one operculum covers an embryo; seed large, 

hollow, initially containing liquid endosperm that 

eventually dries and solidifies; seedling leaf bifid. 

General distribution: Throughout the tropics, 

especially at low elevations, and also widespread 

in subtropical areas. 


Islands: Common in nearly all municipalities and 

inhabited islands at low to middle elevations (sea 

level to ca. 500 m), mostly planted but frequently 

self-propagating. This is one of the world’s most 

important economic trees; almost every part of the 

tree has some use. Further, the graceful and 

dramatic appearance of the coconut palm 

epitomizes and symbolizes the tropics. 

Common names: Puerto Rico: Coco, Cocotero, 

Palma de cocos, Coconut. 


Cabo Rojo: Sintenis 749 (US). Cataño: Goll 967 

(US). Peñuelas: Sintenis 4940 (US). San Juan: Río 

Piedras, Stevenson 5605 (US). Yabucoa: Sintenis 

4941 (US). ST. CROIX: Ricksecker 303 (US). 

6. GAUSSIA 

Gaussia H. Wendl., Nachr. Königl. Ges. Wiss. Georg-Augusts-Univ. 1865: 327. 1865. 

Solitary, unarmed, monoecious palms, with tall, slender, flexible, tapering trunks enlarged at base and 

subtended by many tough prop-roots bearing small spine-like rootlets; trunks straight or sometimes


leaning. Leaves pinnate, the petioles bordered by a wide basal sheath, this split at maturity and not forming 

a crown-shaft. Leaf-segments inserted on rachis in a single plane, the rachis ridged on the upper side. 

Inflorescences interfoliar, protandrous, not long-persisting, paniculate; peduncular bracts 4-7, tubular 

with oblique open apices, the uppermost reaching base of first flowering branch; rachis about as long as 

the peduncle, primary branches numerous, crowded, spirally attached to rachis; rachillae slender, 

glabrous, lacking bracts, bearing spirally arranged rows of flowers, the lowest pistillate, the distal 2-7 

staminate, the most distal one opening first. Staminate flowers larger than the pistillate ones; stamens 6; 

filaments subulate, the anthers sagittate or bifid; pistillode prominent, columnar. Pistillate flowers ovoid 

with rounded sepals and narrow petals; staminodes minute, tooth-like; ovary angled-ovoid with 3 

recurved stigmas, 3-locular with a single ovule in each locule. Fruits ellipsoid or globose to kidneyshaped, 

red or orange when ripe, fleshy with smooth exocarp; seed not adherent to endocarp; endosperm 

homogeneous; embryo lateral. A small genus of 5 species, one endemic to Mexico, another found in 

Mexico, Belize, and Guatemala, two endemic to Cuba, and one species shared by Puerto Rico and the 

Dominican Republic. 

TYPE: Gaussia princeps H. Wendl. 

1. Gaussia attenuata (O. F. Cook) Becc., Pomona 

Coll. J. Econ. Bot. 2: 275. 1912; Aeria 

attenuata O. F. Cook, Bull. Torrey Bot. Club 

28: 548. 1901. Type: Puerto Rico; Vega Baja. 

G. F. Goll 1040 (holotype: US!). 

Gaussia portoricensis H. Wendl. ex Kerch., 

Palmiers 245. 1878, nom. nud. 

Palm 8-12 (30) m tall; trunk basally enlarged 

to 25 cm in diam., tapering upwardly to a 

minimum diam. of (5-) 7-10 cm just below the 

leaves. Leaves mostly 5-7, erect or spreading; 

petiole sheath green, 20-30 cm long; leaves 

(including petiole and sheath) 1-1.8 m long, with 

angular rachis; leaf-segments numerous, the 

longest ones 20-30 cm long, 3 cm broad, the more 

distal ones much shorter, all acute to acuminate at 

apex. Inflorescences much shorter than the leaves; 

panicle branches numerous and densely arranged, 

simple or forked, up to 25 cm long. Flowers 

sessile, orange-yellow, the staminate ones ca. 3 

mm long, the pistillate ca. 2 mm long. Fruits 

obovoid, orange-red, 8-10 mm in diam. 

General distribution: Puerto Rico and 

Dominican Republic. 

Distribution in Puerto Rico: Confined to 

rocky limestone hills and outcrops at low to lower 

middle elevations (5-250 m). Recorded from 

Arecibo, Dorado, Isabela, Juana Díaz, Manatí, 

Quebradillas, San Germán, Toa Baja, Vega Alta, 


Common names: Puerto Rico: Llume, Palma 

de lluvia. 


Juana Díaz: Peña de Las Cuevas, near Juana Díaz, 

E.G. Britton & Marble 2298 (US). Toa Baja: Little 

16304 (US-2). Vega Baja: N.L. Britton & Cowell 

1421 (US-2). 

7. PRESTOEA 

Prestoea J. D. Hook. in Bentham & J. D. Hooker, Gen. Pl. 3: 889. 1883, nom. conserv. 

Small to medium, usually solitary, unarmed, monoecious palms, with trunks lightly banded by leafscars. 

Leaves spreading or arching, pinnate, the sheaths tubular, splitting opposite the petiole, forming 

a short, partially open crownshaft; petioles elongate; leaf-segments regularly inserted on the rachis in a 

single plane. Inflorescences sometimes interfoliar in bud but becoming intrafoliar at anthesis and in 

fruit, paniculate with simple branches; peduncles elongate, terete or nearly so, the first and second nodes 

bearing large, markedly unequal, deciduous primary bracts the first much shorter than the second; 

rachillae glabrous to puberulous or lepidote, stiffly ascending or spreading in all directions from a terete 

rachis. Flowers borne superficially in triads of a single pistillate flower subtended by two small 

bracteoles, flanked by two staminate flowers. Staminate flowers often stipitate, with sepals imbricate at 

base and petals valvate; stamens 6; pistillode prominent. Pistillate flowers with free, imbricate sepals 

and petals; staminodes present; ovary slightly stipitate. Fruits globose, 1-seeded, black or deep purple

144 


when ripe; seeds with ruminate endosperm. Seedling leaves bifid. A Neotropical genus of 28 species 

(Read, 1979) or 10 species (Henderson & Galeano, 1996). A single species occurs in Puerto Rico. 

TYPE: Prestoea pubigera (Griseb. & H. Wendl.) J. D. Hook. ( Hyospathe pubigera Griseb. & H. 

Wendl.). 

Reference: Henderson, A. & G. Galeano. 1996. Euterpe, Prestoea, and Neonicholsonia (Palmae). Fl. 

Neotrop. Monogr. 72: 1-89. 

1. Prestoea montana (Graham) G. Nicholson, Ill. 

Dict. Gard. 3: 216. 1886; Euterpe montana 

Graham, Bot. Mag. 67: t. 3874. 1841; 

Prestoea acuminata var. montana (Graham) 

An. Hend. & Galeano, Fl. Neotrop. Monogr. 

72: 53. 1996. Lectotype: Grenada. Bot. Mag. 

t. 3874, 1841, designated by An. Hend. & 

Galeano, Fl. Neotrop. Monogr. 72: 53. 1996. 

Acrista monticola O. F. Cook, Bull. Torrey Bot. 

Club 28: 557. 1901. Type: Puerto Rico; 

Luquillo Mts. Sintenis 1525 (holotype: US!). 

Euterpe oleracea sensu Bold., Fl. Ned. W. Ind. Eil. 

139. 1913, non Martius, 1824. 

Oreodoxa oleracea sensu Bello, Anales Soc. Esp. 

Hist. Nat. 12: 114. 1883, non Martius, 1824. 

Euterpe globosa sensu authors, incl. Britton & P. 


Gaertner, 1788. 

Small to medium-sized palm, up to ca. 15 m 

tall; trunk 12.5-20 cm in diam., prominently ringed 

by leaf scars. Leaves more or less arcuate; blades 

up to 3.5 m long with numerous flat dark glossy 

green segments; sheaths 30-90 cm long, forming a 

short, poorly defined crownshaft which 

approximately equals the mature inflorescence 

bracts in length, splitting opposite the petiole, 

fibrous, and densely pale-lepidote distally, 

becoming glabrescent with age; petioles 50-75 cm 

long, densely lepidote abaxially, the crowded 

appressed scales fimbriate-stellate, mostly falling 

off with age; leaf-segments chiefly 30-50 on each 

side of the rachis, usually 40-70 cm long (or 

slightly shorter or longer and mostly 3-4.5 cm 

broad near the middle), the upper (adaxial) surface 

usually glabrous but the mid rib of with a 

continuous line of persistent, overlapping, darkmembranous, 

peltate scales, or similar scales 

present but widely separated. Inflorescences 2several, 

infrafoliar, not erect in bud, in length 

about equaling the crownshaft; peduncles 10-20 

cm long, densely appressed-lepidote with pale to 

dark stellate scales; primary bracts leathery or subwoody 

in texture, usually inserted 3-5.5 cm apart 

on the peduncle, the first (outer) bract more or less 

glabrescent, 15-45 cm long, about half the length 

of the second (inner) bract, the latter densely 

matted brown-lepidote; rachis ± terete, glabrous or 

nearly so, bearing 25-50 divaricate branches, these 

chiefly 10-50 cm long, white or lavender at 

anthesis, often turning red in fruit, glabrous or 

nearly so, dilated basally to a broad insertion. 

Flowers white to lavender, pink, or magenta, 

subtended by 2 or 3 small bracteoles, in triads 

(usually 2 staminate with 1 pistillate). Staminate 

flowers pedicellate; sepals glabrous, unequal, 1.2- 

2 mm long., imbricate; stamens white, the anthers 

1.8-2.5 mm long; pistillode prominent, variously 

configured. Pistillate flowers sunken in the 

rachillae; sepals ca. 2.5 mm long, glabrous, 

subequal, rounded and concave, minutely ciliate 

or serrulate; petals 3-4 mm long, glabrous, broadly 

imbricate, deeply concave, opening only enough 

to expose the receptive stigmas; staminodia 6; 

ovary with very short style and 3 stigmas that 

spread apart when receptive. Fruits sunken in the 

rachillae, 10-12 mm in diam. with conspicuous 

subapical stigmatic remains, and basally a 

persistent perianth; seeds 9-10 mm in diam. with a 

conspicuous adaxial raphe; endosperm ruminate. 

Seedling leaf bifid. 

General distribution: Throughout the moist to 

wet mountains of the Greater and Lesser Antilles, 

but absent from Jamaica and the Virgin Islands. 

Distribution in Puerto Rico: Frequent to 

abundant in moist or wet montane (rarely 

submontane) forest. Recorded from Adjuntas, 

Aguas Buenas, Arecibo, Barranquitas, Bayamón, 

Cayey, Jayuya, Maricao, Naguabo, Orocovis, 

Patillas, Ponce, Río Grande, San Lorenzo, Utuado, 

and Yauco. 

Common name: Puerto Rico: Palma de 

Sierra. 

Note: Prestoea montana was reduced to a 

variety of the South American Prestoea acuminata 

(Willd.) H. E. Moore by Henderson & Galeano 

(1996). However, the present writer prefers to 

maintain the disjunct Antillean populations at the


species level as they are easily distinguished from 

the Central and South American relatives. This 

Antillean palm has been the subject of numerous 

ecological studies, and recognizing it at the species 

level allows us to both retain the commonly used 

name Prestoea montana and to maintain 

nomenclatural consistency throughout the 

ecological and floristic literature. Prestoea 

montana can be distinguished from its Central and 

South American relatives by its usually glabrous 

and ± terete rachillae, and by its flowers and fruits 

being superficial on the rachillae [vs. rachillae 

8. PSEUDOPHOENIX 

Pseudophoenix H. Wendl. ex Sarg., Bot. Gaz. 11: 314. 1886. 

pubescent (covered with crustose or granular hairs 

or with long, flexuous hairs) and angular, and 

flowers and fruits somewhat sunken in the 

rachillae]. 


Adjuntas: Alto de la Bandera, near Adjuntas, N.L. 

Britton & Shafer 2122 (US). Cayey: 13 km N of 

Cayey, Underwood & Griggs 340 (US-3). 

Maricao: Maricao to Monte Alegrillo, N.L. Britton 

et al. 2624 (US). Naguabo: Sierra de Naguabo, 

Stevenson 5268 (US). Río Grande: Sierra de 

Luquillo, Sintenis 1525 (NY, US). 

Medium to large, unarmed, solitary palms; trunk somewhat waxy, distinctly bended with leaf-scars. 

Leaves pinnate, the sheaths waxy, glaucous, gray-green, splitting opposite the petiole, abscising and 

shedding cleanly; petioles deeply concave above, convex beneath, glabrous except for small rusty-brown 

scales along the margins. Leaf-segments inserted along the rachis in groups of 2-6, those of the lower half 

of the leaf a various angles, those of the distal half more regularly spaced and in one plane; lowest several 

segments much reduced and more crowded; each segment with a pulvinus at the acute angle of insertion 

at the rachis; upper (adaxial) surfaces dark green, often waxy-glaucous, with 1-5 prominent veins; lower 

(abaxial) surfaces lighter in color, with many minute corrugations. Inflorescences interfoliar, 3 or 4 times 

branched, arching among the leaves or pendulous; peduncles strongly flattened, the first and second nodes 

with large, leathery persistent primary bracts, these splitting irregularly, their margins usually bearing 

large, tufted brown scales; first bract normally longer than and completely enclosing the second; the third 

node frequently with a short, collar-like bract encircling the peduncle. Flowers bisexual, the buds stalked 

(but not pedicellate), ovoid, green, usually glaucous, each subtended by a minute, acuminate bract; each 

pseudo-pedicel a constricted elongation of the receptacle. Calyx lobes triangular with rounded, apiculate 

angles; petals 3, ovate, thick, fleshy, concave, valvate, much longer than the calyx; stamens 6, in two 

whorls, the outer whorl more or less adnate to the petals; filaments thin, dilated at base and often connate, 

forming a short cupule; anthers sagittate, 2-locular, opening by longitudinal slits. Pistil conic, with 3 

glands located basally opposite the outer whorl of staminal filaments; stigmas 3, sessile, recurving only 

for a short period each day, then remaining recurved after fertilization; ovary 3-locular. Fruits globose or 

with 2 or 3 lobes (according to the number of seeds), waxy-red when ripe, attached to rachillae by pseudopedicels 

and subtended by the persistent perianth and staminal filaments; endocarp hard, brown, smooth; 

seeds nearly spherical, the raphe radiating on both sides in flexuous branches; endosperm hard, solid, 

white. Seedling leaf lanceolate, entire. A primarily West Indian genus of 4 species, one of wide 

distribution in the region, the other 3 confined to Hispaniola. 

TYPE: Pseudophoenix sargentii H. Wendl. ex Sarg. 

Reference: Read, R. 1968. Gentes Herb. 10: 169-213. 

1. Pseudophoenix sargentii H. Wendl. ex Sarg., 

Bot. Gaz. 11: 314. 1886. Type: United States; 

Florida. Sargent s. n. (holotype: A; isotype: 

GH). 

Small palm up to 8 m tall, the trunks mostly 

20-30 cm in diam. somewhat enlarged toward the 

base. Leaves ascending or arching, 1.5-3 m long; 

sheaths 15-30 cm long; petioles 20-50 cm long, 4- 

6 cm wide at base, narrowing to 1.5-2 cm at base of 

blade; blades asymmetric with an oblique outline 

at base, with 87-127 segments on each side of

146 


rachis, inserted singly or in groups of 3-5 at diverse 

angles; lowermost segments greatly reduced with 

7-17 occurring in the first 10 cm of rachis; bases of 

segments strongly complicate, 1-5 mm wide 

between the edges; mid rib at base of segments 

normally bearing tufts of brown scales. 

Inflorescences green, compact, divaricate, erect 

among the leaves and 75-120 cm long, or 

projecting immediately below the leaves and 90- 

170 cm long; 3 or 4 times branched; all branches at 

right angles to the main or secondary axes; 

ultimate branches 1-5.5 cm long; peduncle 35-75 

cm long; outer and inner primary bracts subequal, 

always shorter than the peduncle, splitting only 

near apex. Flower buds dark green, glaucous, 20- 

50 per branchlet; pseudo-pedicels mostly 3-6 mm 

long; up to 1.2 mm in diam.; calyx a triangular cup 

with apiculate angles; petals spreading or reflexed, 

6-6.6 mm long; staminal filaments cuspidate, 3.3- 

4 mm long, the dilated bases fused to form a very 

short cupule adnate to the petals. Ripe fruits 

globose if 1-seeded, or 2- or 3-lobed if 2- or 3seeded. 

Ripe 1-seeded fruits when fresh 1.2-1.7 

mm in diam.; pericarp wrinkled when dry; 

endocarp subspherical, 0.8-1.2 mm in diam. 

9. ROYSTONEA 

Roystonea O. F. Cook, Science, ser. 2, 12: 479. 1900. 

General distribution: Littoral habitats 

widespread through the northern Caribbean from 

the Yucatan Peninsula through the Florida Keys, 

northern Cuba Keys, some of the Bahamas, 

Hispaniola, Navassa, Saona, and Mona islands, 

and also an isolated station on Dominica in the 

Lesser Antilles. It is, however, absent from the 

Cayman Islands and Jamaica. 


Islands: Known only from Mona Island, where a 

small grove of 26 trees occurs. Seedlings 

originating from these trees have been distributed 

in Puerto Rico and St. Croix for horticultural use. 

The Mona Island trees grow near sea level in rather 

dry but dense coastal woodland over dolomitic 

limestone. These trees average ca. 6 m tall. 


Mona Island: 0.8 km WNW of Uvero, Proctor et 

al. 45905 (FTG). 


Pseudophoenix vinifera (Mart.) Becc. was 

cited by Martorell et al. (1981) as cultivated in 

Puerto Rico. 

Medium to very large, solitary, unarmed, monoecious palms; trunk columnar, or differentially 

swollen, mostly gray or light brownish to nearly white, often ringed by leaf-scars. Leaves pinnate, the 

segments not plane, extending at various angles. Sheaths tubular, forming a large, elongate, conspicuous, 

green crownshaft this eventually abscising cleanly. Petioles relatively short. Inflorescences intrafoliar, 

produced at the base of the crownshaft, much-branched; peduncle short, strongly fibrous, the first and 

second nodes bearing large, markedly unequal, coriaceous primary bracts, these soon shed as the 

inflorescence develops, the outer bract strongly bicarinate, winged, truncate at apex, the inner bract 1-2 m 

long, fusiform, smooth, splitting longitudinally, inflorescence branches white, prior to exposure 

embedded in a copious mass of free farinaceous trichomes. Flowers sessile, unisexual, but with prominent 

pistillode or staminodes, typically in triads of one pistillate flower accompanied by two staminate ones. 

But often just two or single staminate flowers occur. Staminate flowers with 3 imbricate sepals and 3 

petals, the latter joined at base, valvate above, and acute at apex; stamens usually 6, the pistillode 

prominent. Pistillate flowers with 3 broadly imbricate sepals and 3 valvate petals; staminodes joined in a 

crenulate or stellate ring; pistil conic, 3-pistillate and with 3 stigmas, the stigmatic remains persisting 

conspicuously on one side near base of the fruit. Fruits oblong, globose, or curved-pyriform, less than 2 

cm long, purplish to nearly black; exocarp very thin; mesocarp soft and juicy; endocarp a firm, thin shell 

within which the seed is usually free at maturity; usually 1-seeded; endosperm white, homogeneous. 

Seedling leaf lanceolate. A circum-Caribbean genus of 10 species (Zona, 1996), widely planted in tropical 

regions. A single species occurs naturally in Puerto Rico and the Virgin Islands. At least two other species 

are occasionally planted in our area. 

TYPE: Roystonea regia (Kunth) O. F. Cook (≡ Oreodoxa regia Kunth) 

Reference: Zona, S. 1996. Fl. Neotrop. Monogr. 71: 1-35. 1996.


1. Roystonea borinquena O. F. Cook, Bull. 

Torrey Bot. Club 28: 552. 1901. Type: Puerto 

Rico; Luquillo Mts. Sintenis 1605 (holotype: 

US!; isotypes: GH, MO, NY, P). 

Roystonea hispaniolana L. H. Bailey, Gentes 

Herb. 4: 268. 1939. Type: Dominican 

Republic. Bailey 242 (holotype: BH). 

Roystonea hispaniolana f. altissima Moscoso, 

Cat. Fl. Domingensis 63. 1943. Type: not 

seen, perhaps not extant. 

Roystonea peregrina L. H. Bailey, Gentes Herb. 8: 

127. 1943. Type: Guadeloupe? A cultivated 

plant of uncertain origin. Questel 51 

(holotype: BH). 

Fig. 32. E-G 

Trunk columnar 18 m tall or more, the grayish 

to grayish brown trunk mostly 40-50 cm in diam. 

in the lower part but often expanding to as much as 

60 cm just below the crownshaft, nearly smooth 

throughout. Leaves usually 12-15, ascending and 

arching, several hanging well below the horizontal, 

mostly 3-4 m long; crownshaft 1.5-1.7 m long; 

petioles 30-55 cm long; leaf-segments numerous, 

mostly 60-100 cm long (shorter toward the ends, 

longest near the middle), 2.5-5 cm broad, linear, 

folded toward base, with a prominent mid rib. 

Inflorescences 1-1.4 m long, twice branched, 

much shorter than the leaves. Staminate flowers 

whitish or creamy yellow with 6-9 stamens and 

bright purple anthers, the petals 6.3-6.4 mm long, 

the pistillode minute. Pistillate flowers also 

creamy yellow to whitish, but with shorter petals 

2.4-4.4 mm long; staminodial cup shallowly 6lobed. 

Fruits spheroid to ellipsoid, 11.6-15 mm 

long, light brown to nearly black when ripe. Seeds 

Sabal Adans., Fam. Pl. 2: 495. 1763. 

10. SABAL 

oblong-ellipsoid to globose, ca. 9 mm long. 

General distribution: Puerto Rico, Virgin 

Islands, and Hispaniola. 


Islands: This species grows naturally in moist, 

deep soils at low to middle elevations (near sea 

level to ca. 300 m), additionally it is widely 

planted. The statement by Zona (1996) that this 

species in Puerto Rico grows only on soils derived 

from limestone cannot be supported. Also, the 

synonymization of Roystonea hispaniolana under 

R. borinquena deserves further evaluation. This 

species is more common than its few records seem 

to indicate. Recorded from Arecibo, Coamo, 

Culebra, Dorado, Juana Díaz (planted), Naguabo 

(planted), Río Grande, San Germán, San Juan, San 

Sebastián, Utuado, Vieques, Villalba; St. Croix, 

St. John, and Tortola. 

Common names: Puerto Rico: Palma real, 

Palma real puertorriqueña, Palma de yaguas. 


Coamo: Coamo Springs, Goll 736 (US). Naguabo: 

N.L. Britton & E.G. Britton 7813 (US). Río 

Grande: Sierra de Luquillo, Sintenis 1605 (US). 

San Juan: Río Piedras, Little 16431 (US). Vieques: 

Playa Grande to La Mina, Shafer 3006 (US). ST. 

CROIX: Fountain Valley Golf Course, Fosberg & 

Hayes 58958 (US). 


Roystonea regia (Kunth) O. F. Cook., the 

Cuban Royal Palm, is occasionally cultivated on 

Puerto Rico and the Virgin Islands (St. Thomas). 

Roystonea oleracea (Jacq.) O.F. Cook and R. 

venezuelana L. H. Bailey were cited by Martorell 

et al. (1981) as cultivated in Puerto Rico. 

Small to large, solitary, unarmed palms; trunk massive or rarely slender or (in one species) decumbent 

and subterranean to ascending, often shaggy with persistent split petiole bases, or in some species or 

individuals these bases soon deciduous, leaving a clean, naked trunk ringed with leaf-scars. Leaves 

costapalmate, flabellate with a strong central costa that is an extension of the petiole. Petioles elongate, 

rounded beneath (abaxially) channeled on upper (adaxial) side, with sharp margins, split at base of sheath 

portion; leaf-costa usually elongate and down-curved in most species; segments plicate and connate 

through most of their length, the apices free and bifid, the margins often bearing numerous free threadlike 

fibers; hastula prominent at base of blade on upper (adaxial) side. Inflorescences interfoliar, compoundpaniculate 

the axes mostly covered by obliquely pointed, tubular, sheathing bracts, except the penultimate

148 


Fig. 32. A-D. Coccothrinax alta. A. Habit. B. Leaf. C. Infructescence. D. Fruit and embryo. E-G. Roystonea borinquena. E. 

Habit. F. Portion of leaf. G. Portion of infructescence. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. 

Gard. 78.


and ultimate branches. Flowers bisexual, sessile, solitary; calyx cupular, tridentate; petals concave, 

valvate toward apex; filaments subulate, connate at base and adnate to base of the corolla; ovary 3pistillate; 

style as long as or longer than the ovary, grooved on 3-sides; stigma undivided, slightly capitate, 

minutely papillose. Fruits globose to sub-pyriform, brown or black when ripe, 1-seeded; mesocarp fleshy, 

endocarp membranous. Seeds depressed-globose; endosperm homogeneous. Seedling leaf narrowly 

lanceolate. A broadly circum-Caribbean genus of 26 species (Read, 1979) or 15 species (Zona, 1990). One 

species occurs naturally in Puerto Rico and the Virgin Islands; another has been widely planted, especially 

in St. Croix, where it is somewhat naturalized. 

LECTOTYPE: Sabal adansonii Guers. (≡ Corypha minor Jacq.; Sabal minor (Jacq.) Pers.), designated 

by O.F. Cook, Bull. Torrey Bot. Club 28: 529-530. 1901. 

Reference: Zona, S. 1990. A monograph of Sabal (Arecaceae: Coryphoideae). Aliso 12 (4): 583-666. 

Key to the species of Sabal 

1. Trunk 20-35 cm in diam.; leaves evenly green, not glaucous; fruits black, 13-18 mm in diam 

............................................................................................................................……. 1. S. bermudiana 

1. Trunk 40-75 cm in diam.; leaves green or somewhat glaucous; ripe fruits dark brown, 5.9-8(10) mm in 

diam. ………...........................................................................................................….. 2. S. causiarum 

1. Sabal bermudiana L. H. Bailey, Gentes Herb. 

3: 326. 1934. Lectotype: Bermuda. Bailey et. 

al. 73 (BH), designated by Zona, Aliso 12: 

624. 1990. 

Sabal princeps Becc., Webbia 2: 59. 1907; Sabal 

beccariana L. H. Bailey, Gentes Herb. 4: 387. 

1940, nom. illeg. Lectotype: A plant 

cultivated in Palermo, Italy. Mattei s. n. (FI), 

designated by L.H. Bailey, Gentes Herb. 4: 

387. 1940. 

Sabal blackburniana sensu Britton & P. Wilson, 

Bot. Porto Rico 5: 116. 1923, non Glazebrook 

1829. 

A slow-growing palm with a stout trunk up to 

7 m tall, 20-35 cm in diam., gray and obscurely to 

prominently ringed with leaf-scars. Petioles 1-2 m 

× 2.7-4 cm, longer than the blades; hastula acute to 

acuminate, 8.5-18.5 cm long, glabrous or nearly 

so, the margins flat, revolute or involute; blades up 

to 1.5 m long, with 85-95 segments per leaf, these 

3.2-4.4 cm wide, joined at base for ca. 50 % of their 

length, the free portion long-bifurcate, the tissue 

firm and light green on both sides; margins bearing 

few to many hair-like filaments. Inflorescences 

arcuate, not exceeding the petioles in length; 

sheathing bracts usually lepidote; penultimate 

branches short and often not emergent beyond the 

subtending bracts of the main axis; ultimate 

branchlets 4-14 cm long. Flowers 4-6 mm long at 

anthesis; filaments 3. 1-4.5 mm long; pistil short, 

stout. Fruits pyriform, black when ripe, with thick 

pericarp; seeds oblate-concave, 7.5-12.5 mm in 

diam., often with a sharp funicular beak. 

General distribution: endemic to Bermuda, 

but widely cultivated elsewhere. 

Distribution in the Virgin Islands: Reported 

by Britton & P. Wilson (1923) as commonly 

planted in St. Croix and St. Thomas, where its 

leaves were “used for making hats and in 

basketry”. Subsequently it became somewhat 

widely naturalized on St. Croix, especially in the 

southern part, but more recently has been much 

reduced in numbers by housing development. No 

up-to-date information is available on the status of 

this species on St. Thomas. It is planted on St. 

John. 

Selected specimens examined: ST. THOMAS: 

Charlotte Amalie, N.L. Britton & Rose 1397 (US). 

2. Sabal causiarum (O. F. Cook) Becc., Webbia 

2: 71. 1907; Inodes causiarum O. F. Cook, 

Bull. Torrey Bot. Club 28: 531. 1901. Type: 

Puerto Rico. Underwood & Griggs 154 

(holotype: US!, isotype: NY). 

Inodes glauca Dammer in Urban, Symb. Antill. 4: 

127. 1903. Type: Puerto Rico; near Peñuelas. 

Sintenis 4844 (holotype: B, destroyed; 

isotypes: GH, MO, P, US!). 

Sabal haitensis Becc., Ann. Roy. Bot. Gard. 

(Calcutta) 13: 293. 1931(1933). Type: Haiti. 

Buch s. n. (holotype: B, destroyed, fragment

150 


FI). 

Sabal questeliana L. H. Bailey, Gentes Herb. 6: 

422. 1944. Type: St. Barthelemy. Questel 468 

(holotype: BH). 

Sabal umbraculifera sensu Bello, Anales Soc. 

Esp. Hist. Nat. 12: 114. 1883, non Martius, 

1824. 

Fig. 62. G 

Large, heavy-trunked palm up to 16 m tall, the 

trunk gray, slightly rough in texture, and faintly 

ringed with leaf-scars. Petioles 1-2 m long, up to 

nearly 5 cm wide, equaling or exceeding the blades 

in length; hastula acute, up to 21 cm long or more, 

glabrous or nearly so, with margins often 

prominently upturned; blades usually 1.5-2 m 

long, with 60-120 segments per leaf, these 2.5-5.8 

cm wide joined basally for ca. 49 % of their length, 

the free portion long-bifurcate; tissue firm, light 

green on upper (adaxial) side, usually somewhat 

glaucous beneath; numerous hair-like filaments 

borne from the sinuses between the segments. 

Inflorescences arcuate, conspicuously equaling or 

extending beyond the leaves; primary bracts 

strongly pointed; ultimate branchlets slender, 

glabrous, 3-8 cm long. Flowers white, slightly 

fragrant, 2.7-5.2 mm long at anthesis; filaments 

2.8-4.5 mm long; pistil short, stout. Fruits globose 

to globose-pyriform, often asymmetric; seeds 

oblate-concave, 5.9-8 (-10) mm in diam., dark 

glossy brown. 

General distribution: Hispaniola (southern 

coasts of Haiti and the Dominican Republic), 

Puerto Rico and the Virgin Islands. 


Thrinax Sw., Prodr. 4, 57. 1788. 

11. THRINAX 

Islands: This species, which grows naturally 

chiefly near sea level (up to 100 m), seems to be 

indifferent as to substrate, occurring on sandy soils 

in some areas but elsewhere over limestone and 

different types of igneous rock (e. g., andesite on 

Guana Island). It is frequently planted and 

cultivated for its massive, stately appearance. A 

famous avenue of these trees can be seen along the 

“Malecon”, the seafront boulevard of the city of 

Santo Domingo. Recorded from Aguadilla, Cabo 

Rojo, Camuy, Guánica, Isabela, Peñuelas, Ponce, 

Quebradillas, San Juan, San Sebastián, Yabucoa, 

and Yauco (although cultivated in many other 

localities); Anegada, Guana Island, St. Croix 

(sight record), and Tortola (cultivated). 

Common names: Puerto Rico: Palma de 

sombrero, Palma de escoba. 


Cabo Rojo: Cabo Rojo to Joyuda, N.L. Britton et 

al. 2403 (US). Camuy: Stevenson 5152 (US). 

Isabela: Bo. Jobos, Proctor et al. 41885 (US). 

Peñuelas: Sintenis 4844 (US). Ponce: Ponce to 

Peñuelas, N.L. Britton et al. 1775 (US). 

Quebradillas: Goll 955 (US-2). San Juan: Río 

Piedras, Little 16307 (US). Yabucoa: Sintenis 

5286 (US). Yauco: N.L. Britton & Shafer 1892 

(US). ANEGADA: N.L. Britton & Fishlock 1061 

(US); Acevedo-Rdgz. & Smith 11044 (US). 


Sabal palmetto (Walter) Lodd. ex Schult. f. is 

sometimes cultivated in Puerto Rico and St. Croix, 

its natural range includes southeastern United 

States, the Bahamas, and Cuba. This has thus far 

shown no tendency to become naturalized. 

Small to medium, solitary, unarmed palms; trunk rather slender, columnar, at base growing from a 

dense mass of roots forming a small mound above the rocky substratum, above smooth or fibrous, 

obscurely ringed with leaf-scars. Leaves palmate; sheaths fibrous, tubular at first, later becoming netlike; 

petioles flat on upper (adaxial) side, convex beneath, terminating at base of blade in the hastula, splitting 

at base within area of sheath; hastula conspicuous, with an adaxial ridge, flap or small projection. Leaf 

blade flabellate, lacking a central costa; upper (adaxial) surface glabrous; lower (abaxial) surface 

variously lepidote; segments joined basally to form a plicate continuous surface, free outwardly, tapering 

to a bifid apex. Inflorescences interfoliar, elongate, erect to arching, with several or numerous pendulous 

branches, these in turn usually simple-branched; lowest primary bract bicarinate, the others tubular with 

an oblique aperture; all primary bracts densely appressed-lepidote, each enclosing the base of a primary


branch; primary branches each with a bicarinate and bifid bract inserted midway along the flattened 

peduncle and partially enclosing the flowers before anthesis; each ultimate branch subtended by a narrow 

triangular bract. Flowers bisexual, mostly protandrous, solitary on prominent or very short bracteolate 

pedicels. Perianth a 6-lobed or-dentate cupule not enclosing the reproductive parts; stamens usually 6-12 

(sometimes fewer or more), the filaments straight, slender, and free, the anther locules opening by lateral 

slits. Pistil 1-locular, 1-ovulate, the style flaring upwardly to a laterally-compressed, funnel-shape stigma. 

Fruits small, depressed-globose, white when ripe; mesocarp thin, mealy; seeds depressed-globose, 

smooth, tan when fresh; endosperm homogeneous. Seedling leaf lanceolate, entire. A genus of four 

species, two of them endemic to Jamaica, the other two widely distributed from Honduras to Mexico and 

Florida, also in the Bahamas, and in the Antilles eastward to Barbuda. The two widespread species are not, 

however entirely sympatric, as only T. morrisii occurs in Puerto Rico, the Virgin Islands, and the western 


TYPE: Thrinax parviflora Sw. 

1. Thrinax morrisii H. Wendl., Gard. Chron. ser. 

3, 11: 104, f. 20, 21. 1892. Lectotype: 

Anguilla, Lesser Antilles. Nicholls, s. n. (K), 

designated by Read, Smithsonian Contr. Bot. 

19: 88. 1975. 

Thrinax microcarpa Sarg., Gard. & Forest 9: 162. 

1896. Type: United States; Florida. Curtiss 

2679 (holotype: NY). 

Thrinax praeceps O. F. Cook, Bull. Torrey Bot. 

Club 28: 536. 1901. Type: Puerto Rico. Cook 

850 (holotype: US!). 

Thrinax ponceana O. F. Cook, Bull. Torrey Bot. 


Ponce. Cook 1005 (holotype: US!). 

Fig. 62. H, I 

Small to medium palm 1-10.5 m tall, often 

producing flowers and fruits when very low in 

stature and remaining permanently low under 

harsh growth conditions; trunk (5-) 18-35 cm in 

diam. near base, slightly tapering upward, ashy 

gray, smooth to fibrous, with leaf and inflorescence 

scars sometimes evident, the whole trunk 

sometimes forming a network of intersecting 

cracks with age. Leaf sheath 28-60 cm long, soon 

breaking apart and separating from the petiole 

apically, becoming somewhat netlike, densely 

velvety at first, soon becoming glabrescent; 

petioles 27-54 cm long, at first densely whitelepidote 

abaxially, soon glabrescent; hastula 

ovate, obtuse or rarely pointed, erect, thin, 0.2-0.8 

cm long, up to 3 cm wide, densely velutinous, 

ciliate, at first, also with conspicuous tufts of white 

or silvery indumentum, becoming glabrescent. 

Developing leaf blades densely appressedlepidote 

on all surfaces; expanded leaf blades 

usually glossy green on upper surface, often 

becoming somewhat glaucous or blue-green with 

age, variously lepidote beneath, the minute scales 

hyaline, fimbriate, and somewhat interlocking, 

becoming glabrescent with age. Mature leaf blades 

nearly circular in outline, mostly 75-150 cm in 

diam., the 33-58 segments (rarely all of these lying 

in the same plane), rigid or flexuous, the longest 

middle ones 55-75 cm long and mostly 3.5-4.8 cm 

wide, widest at point of fusion, bifid at apex. 

Inflorescences 55-100 cm long, erect or arching 

equaling or exceeding the leaves; primary bracts 

silvery-white to tan-lepidote, often apically tufted 

with long white scales; primary branches 9-21, 

glabrous, pendent or at first tufted-arcuate, mostly 

9-30 cm long at anthesis; ultimate branches 

numerous, mostly 5.5-14.5 cm long, subtended by 

narrow triangular bracts. Flowers white, becoming 

pale yellow or pale orange with age, attached to 

inconspicuous disc-like pedicels with triangular 

bracteoles; perianth-lobes triangular-apiculate; 

stamens mostly 6, the filaments broadly connate at 

base, forming a cup; anthers exceeding the pistil in 

length; stigma minutely funnel-shaped, ciliate. 

Mature fruits globose, variable in size (often on the 

same inflorescence), (3.5-) 4-4.5 (-8 ) mm in 

diam., on pedicels 0.1-0.8 mm long. Seeds 

mahogany brown, usually with a small cavity at 

base. Flowering usually in May and June. 

General distribution: Florida, Bahamas, Turk 

& Caicos Islands, Greater Antilles and some 

associated small islands (but not Jamaica), 

eastward to Barbuda in the Leeward Islands. 


Islands: Occurs mainly on calcareous or neutral 

soils at low to lower middle elevations (sea level to 

450 m), often common in dry barren situations. 

Recorded from Arecibo, Bayamón, Cabo Rojo,

152 


Corozal, Fajardo, Guánica, Lajas, Lares, Mona 

Island, Peñuelas, Ponce, Quebradillas, Vega Baja, 

Vieques, and Yauco; Anegada. 

Common names: Puerto Rico: Palma de 

escoba, Pandereta. 



294 (SJ). Bayamón: on mogotes near Bayamón, 

Read 2058 (US-2). Cabo Rojo: Salinas de Cabo 

Rojo, Sintenis 600 (US). Camuy to Quebradillas, 

N.L. Britton & Cowell 1953 (US-2). Corozal: N.L. 

Britton & E.G. Britton 7829 (US). Fajardo: 

Sintenis 1324 (US). Guánica: N.L. Britton & 

Shafer 1922 (US). Lares: N.L. Britton et al. 2750 


(US). Mona Island: Otero & Chardon 911 (US). 

Ponce: Bo. Cañas, off Rd. 2 behind the Holiday Inn 

Hotel, Acevedo-Rdgz. & Cedeño 7761 (NY, 

UPRRP, US-2). Vieques: Lighthouse Peninsula, 

Shafer 2796 (US-2). Yauco: Susúa Forest 

Reserve, Acevedo-Rdgz. 11413 (NY, UPRRP, 

US). Vega Alta: S of communication tower, 

Acevedo-Rdgz. 12289 (US). Vega Baja: West of 

Vega Baja, N.L. Britton et al. 6811 (US-2). 

ANEGADA: West End, Fishlock 1062 (US-2). Along 

road from Pomato Point to West End, Acevedo- 

Rdgz. 11048 (US). West side of Island, directly S 

of Cow Wreck Bay, Acevedo-Rdgz. et al. 11574 

(UPRRP, US). 

The following taxa were listed by Britton & P. Wilson (1923) as being cultivated in Puerto Rico and/ 

or the Virgin Islands. 

Actinophloeus macarthurii (H. Wendl.) Becc., native to Australia, luxuriant in Puerto Rico gardens; 

Areca catechu L., Asiatic, planted in Puerto Rico and St. Thomas; Areca madagascariensis Mart., native 

of Madagascar, cultivated at the Mayagüez Agricultural Experiment Station; Arenga saccharifera Labill., 

East Indian, occasionally planted for ornament in Puerto Rico; Borassus flabellifer L., East Indian, 

recorded by West as found on St. Croix prior to 1793 and by Krebs on St. Thomas prior to 1851; Butia 

capitata (Mart.) Becc., from Brazil, grown at the Mayagüez Agricultural Experiment Station; Caryota 

mitis Lour., of Malaysian origin, cultivated at the St. Croix Agricultural Experiment Station in 1923; 

Caryota urens L., Asiatic, occasionally planted for ornament in Puerto Rico; Corypha elata Roxb., native 

of India, cultivated at the Mayagüez Agricultural Experiment Station; Dictyosperma rubrum H. Wendl. 

& Drude, East Indian, cultivated at the Mayagüez Agricultural Experiment Station; Dypsis lutescens (H. 

Wendl.) Beentje & J. Dransf. (Chrysalidocarpus lutescens H. Wendl.), from Madagascar, planted for 

ornament in Puerto Rico (San Juan); Elaeis guineensis Jacq., of African origin, cultivated at the Mayagüez 

Agricultural Experiment Station; Howea belmoreana (C. Moore & F. Muell.) Becc., native of Lord 

Howe’s Island in the Pacific Ocean, occasionally planted in Puerto Rico; Hyophorbe verschaffeltii H. 

Wendl., of the Mascarene Islands, occasionally planted for ornament in the Virgin Islands; Licuala 

grandis H. Wendl., native of New Britain Island, occasionally grown for ornament in Puerto Rican 

gardens; Livistona chinensis R. Br., from China, planted for ornament in Puerto Rico and the Virgin 

Islands; Martinezia caryotifolia Kunth, from South America, cultivated at the Mayagüez Agricultural 

Experiment Station; Phoenix dactylifera L., from Africa, planted in Puerto Rico and the Virgin Islands; 

Phoenix canariensis Chabaud, of the Canary Islands, cultivated at the St. Croix Agricultural Experiment 

Station; Phoenix reclinata Jacq., native of Africa, occasionally planted in Puerto Rico and Virgin Islands 

gardens; Phoenix rupicola T. Anderson, East Indian, occasionally planted for ornament in Puerto Rico and 

St. Thomas; Pritchardia pacifica Seem. & H. Wendl., native of the Fiji Islands, planted for ornament in 

Puerto Rico gardens; Rhapis excelsa (Thunb.) A. Henry ex Rehder (Rhapis flabelliformis L’Hér.), native 

of western Asia, frequently grown for ornament in Puerto Rico gardens; Rhopadostylis baueri (Hook.) H. 

Wendl. & Drude, native of the Norfolk and Chatham Islands, cultivated at the St. Croix Agricultural 

Experiment Station; Syagrus romanzoffiana (Cham.) Glassman (Cocos plumosa Lodd. ex Hook.), native 

of Brazil, occasionally planted for ornament in Puerto Rico; Washingtonia filifera (Linden ex André) H. 

Wendl. (Neowashingtonia filifera (Linden ex André) Sud.), native of southern California, U.S.A., 

cultivated at St. Croix Agricultural Experiment Station; Washingtonia robusta H. Wendl. 

(Neowashingtonia robusta (H. Wendl.) Britton), native of southern California, U.S.A., occasionally


planted for ornament in Puerto Rico. 

The following taxa were cited by Martorell et al. (1981) as being cultivated in Puerto Rico: 

Archontophoenix cunninghamiana (H. Wendl.) H. Wendl. & Drude, Arecastrum romanzoffianum 

(Cham.) Becc., Arenga pinnata (Wurmb) Merr., Arikuryroba schizophylla (Mart.) L. H. Bailey, 

Astrocaryum standleyanum L. H. Bailey, Bactris gasipaes Kunth, Chamaedorea elegans Mart., 

Chrysalidocarpus madagascariensis Becc., Corypha umbraculifera L., Cryosophila warscewiczii (H. 

Wendl.) Bartlett, Dictyosperma album (Bory) H. Wendl. & Drude ex Scheff., Elaeis oleifera (Kunth) 

Cortés ex Prain, Latania loddigesii Mart., Phoenix roebelinii O’Brien, Phoenix sylvestris (L.) Roxb., 

Pinanga kuhlii Blume, Ptychosperma elegans (R. Br.) Blume, Ptychosperma macarthurii (H. Wendl. ex 

H.J. Veitch) H. Wendl. ex Hook. f., Raphia farinifera (Gaertn.) Hyl., Syagrus coronata (Mart.) Becc., and 

Veitchia merrillii (Becc.) H. E. Moore. 

Family 20. COMMELINACEAE Spiderwort Family 

Commelinaceae R. Br., Prodr. 268. 1810, nom. conserv. 


Annual or perennial herbs with erect or trailing sub-succulent stems; leaves alternate (rarely in a 

rosette), simple, parallel veined and usually entire, mostly with sheathing or clasping bases. Flowers 

usually bisexual, actinomorphic or zygomorphic, 3-merous, in terminal, axillary, or leaf-opposed cymes 

or umbels, these subtended by spathe-like or leafy bracts. Sepals usually free and distinct, green and 

herbaceous. Corolla delicate and soon-withering, blue, pink or white (in our species), of free, equal or 

unequal clawed petals, or else united in a short tube. Stamens most often 6, in two whorls of 3, or in some 

genera reduced in number; anthers 2-locular. Ovary superior, 2- or 3-locular; ovules 1 to several in each 

locule; sometimes only 1 or 2 locules fertile; style simple, terminal, with a capitate or 3-lobed stigma. Fruit 

a capsule or berry. A tropical to warm-temperate family of 30 genera and ca. 650 species. There are 7 

genera represented by 14 species in Puerto Rico. 

TYPE: Commelina L. 

References: Faden, R. B. & Hunt, D. R., 1991. The classification of the Commelinaceae. Taxon 40: 

19-31. Howard, R. A., 1979. Commelinaceae in Fl. Lesser Antill. 3: 427-448. Hunt, D. R., 1983. 

Commelinaceae in Fl. of Trinidad & Tobago 3 (3): 255-275. Tucker, G. C., 1989. The genera of 

Commelinaceae in the southeastern United States. J. Arnold Arbor. 70: 97-130. 


1. Fertile stamens usually 6; filaments mostly hairy. …………...........................................................…. 2 

2. Inflorescences subtended by conspicuous, paired, foliaceous bracts …....……….. 6. Tradescantia 

2. Inflorescences subtended by solitary, inconspicuous, green bracts. …….............……………… 3 

3. Leaves entire: flowers white, actinomorphic; stamens all equal ………....……… 4. Gibasis 

3. Leaves finely serrulate; flowers pink (rarely white), zygomorphic; stamens dimorphic (3 

long, 3 short ) …....................................................................................…….. 7. Tripograndra 

1. Fertile stamens 1-3 (6 in Callisia fragrans), filaments glabrous. ……................................…………. 4 

4. Flowers in dense sessile clusters ………..............................................................……. 2. Callisia 

4. Flowers obviously pedicellate and sometimes also with long peduncles. …………..........……. 5 

5. Flowers strongly zygomorphic, the larger petals 5-12 mm long, bright blue, pale blue, or 

white …..................................................................................................…….. 3. Commelina 

5. Flowers actinomorphic, very small (2 mm long or less) whitish or dull bluish. …………. 6

154 


6. Plants diffuse, weakly erect, the stems loosely pilose; leaves ovate to ovate-lanceolate, 

less than 4 cm long, abruptly narrowed at base ……..............................……. 1. Aploleia 

6. Plants prostrate or nearly so, glabrous, forming mats; leaves linear or linear-lanceolate, 

up to 8 cm long, clasping stem at base …………..........................…….. 5. Murdannia 

1. APLOLEIA 

Aploleia Raf., Fl. Tellur. 2: 17. 1837 [“1836”] 

Weakly straggling herbs, the stems decumbent or ascending, much branched, the vegetative branches 

penetrating the sheaths at the nodes. Inflorescence terminal, open or diffuse, consisting of several or many 

pedunculate cymes arising from between minute paired bracts in the axils of ordinary or reduced leaves; 

peduncles always longer than the subtending leaves, terminating in small paired umbelliform cymes 

(cincinni). Flowers unequally long-pedicellate; sepals 2-3, herbaceous; petals 2-3, soon withering and 

deliquescent; stamens 1-3, opposite the sepals, the filaments glabrous, distinct, erect; anthers ovoid in bud, 

linear after dehiscence; ovary 2-3 locular with 2 ovules in each locule; style very short, with pedicellate 

stigma. A circum-Caribbean genus of two species, sometimes included in an expanded genus Callisia, to 

which it is alleged to be closely related. 

TYPE: Aploleia monandra (Sw.) H. E. Moore (≡ Tradescantia monandra Sw.). 

1. Aploleia monandra (Sw.) H. E. Moore, 

Baileya 9: 17. 1961; Tradescantia monandra 

Sw., Prodr. 57. 1788; Callisia monandra 

(Sw.) Schult. & Schult. f., Syst. Veg. 7. 1179. 

1830. Type: Hispaniola. Swartz s. n. 

(holotype: S-Sw, #R-6170). 

Callisia umbellulata Lam., Tabl. Enclycl. 1: 130, t. 

35, f. 2. 1791. Type: South America. 

Lamarck, Illus. 1: 130, t. 35, fig. 2. 1791. 

Figs. 33. A-E; 63. A 

Stems weak, glandular-pilose chiefly on one 

side, rooting at nodes. Leaves subsessile, the 

blades ovate or ovate-lanceolate, 2-4 cm long, 1-2 

cm broad, acuminate at apex, the margins ciliate. 

Peduncles filiform, 1-3 cm long; sepals ellipticoblong, 

1.6-2 × 0.9-1 mm, glandular-pilose or 

glabrous dorsally; petals lanceolate, smaller than 

the sepals, hyaline; stamens 1 or 2; ovary oblongellipsoid, 

compressed or angled, glabrous or pilose 

at apex; stigma 3-lobed. Capsule 1-1.3 mm long, 

apiculate. 


America, northern South America, Greater 

Antilles (except Jamaica), Guadeloupe, and 

Martinique. 

Distribution in Puerto Rico: Widespread in 

moist to wet districts and appears to prefer noncalcareous 

habitats, at medium to rather high 

elevations (100-1050 m). Recorded from Adjuntas, 

Aibonito, Arecibo, Barranquitas, Bayamón, 

Caguas, Cayey, Ciales, Jayuya, Las Marias, 

Maricao, Mayagüez, Naguabo, Patillas, Río 

Grande, Salinas, San Juan, Utuado, Villalba, 

Yabucoa, and Yauco. 

Common name: Puerto Rico: Cohítre morado. 


Adjuntas: Alto de la Bandera, N.L. Britton & 

Shafer 2112 (US). Aibonito: Between Aibonito 

and Cayey, Heller & Heller 525 (US). Arecibo: 

Río Abajo Forest Reserve, Acevedo-Rdgz. & 

Angell 9379 (UPR, UPRRP, US). Barranquitas: 

Bo. Barrancas, Proctor & Thomas 44442 (US). 

Bayamón: Bo. Santa Olaya, Proctor 43190 (US). 

Cayey: along Rd 1, km 65, Acevedo-Rdgz. 7911 

(NY, UPRRP, US). Las Marias: La Juanita, near 

Las Marias, N.L. Britton et al. 3927 (US). 

Maricao: Indiera Fría, Acevedo-Rdgz. 4716 (F, K, 

MO, US). Mayagüez: 5 mi. NE of Mayagüez, 

Heller 4452 (US). Naguabo: Sierra de Naguabo, 

Bo. Maizales, N.L. Britton & Cowell 2176 (US). 

Patillas: Bo. Marín, Proctor & Pinto 40288 (US). 

Río Grande: Sierra de Luquillo, Caribbean Natl. 

Forest, Road 191, km 9.9, at Quebrada Juan Diego, 

Proctor 44486 (US). Utuado: N.L. Britton & 

Cowell 988 (US). Villalba: Vicinity of Ala de la 

Piedra above Villalba, N.L. Britton & Earle 6073 

(US).


Fig. 33. A-E. Aploleia monandra. A. Habit. B. Inflorescence unit. C. Flower in fruit. D. Capsule. E. Seeds. F-I. Gibasis geniculata. 

F. Habit. G. Flower. H. Fruit. I. Seeds. J-O. Tradescantia zanonia. J. Habit. K. Inflorescence unit. L. Petal and stamens. M. 

Pistil. N. Upper portion of stamen showing anther. O. Seeds. (A, from Liogier 30289 and Proctor 40288; B-E, from Liogier 

30289; F, from Proctor 41221; G-I, from Cedeño 583; J, from Proctor 49928; K-O, from Proctor 44154).

156 

Callisia Loefl., Iter Hispan. 305. 1758. 


2. CALLISIA 

Perennial creeping or ascending herbs, the stems slender to rather stout and succulent. Leaves pale to 

dark green, at least at the upper ones subsessile and vaginate, the blades ovate to elliptic-lanceolate or 

oblong-lanceolate. Flowers actinomorphic, bisexual, typically in sessile fascicles representing pairs of 

fused abbreviated cymes in the axils of leaves; the latter either normal in size and shape or else reduced 

to a vestige little more than a sheath. Sepals 2-3, subequal, usually hyaline, erect, persistent in fruit; petals 

2-3, subequal, white or pink; marcescent; stamens 1-6, inserted opposite the petals, the filaments glabrous, 

anthers 2-locular, the locules rounded and separated by a band of sterile tissue; ovary oblong, sessile, 2or 

3-locular, each locule with 2 ovules; style long, with pedicellate or lobed stigma. Fruit a small capsule; 

seeds with punctiform hilum. A tropical American genus of 10 (Howard, 1979) to 20 species (Acevedo- 

Rodríguez, 1996). 

NEOTYPE: Callisia repens (Jacq.) L. (≡ Hapalanthus repens Jacq.), designated by Britton & P. Wilson, 

Bot. Porto Rico 5: 147. 1923. 

Key to the species of Callisia 

1. Stems decumbent-ascending, succulent, 5-10 mm thick; leaves 8-28 cm long; flowers densely clustered 

at intervals on terminal racemiform or panicle-like inflorescence, ovary and capsule 3-locular 

…............................................................................................................................……. 1. C. fragrans 

1. Stems prostrate, long creeping, very slender, 1-2 mm thick, leaves 1-4 cm long; flowers sessile in minute 

axillary paired cymules; ovary and capsule 2-locular …….......................................…….. 2. C. repens 

1. Callisia fragrans (Lindl.) Woodson, Ann. 

Missouri Bot. Gard. 29: 154. 1942; Spironema 

fragrans Lindl., Edward’s Bot. Reg. 26: t. 47. 

1840; Rectanthera fragrans (Lindl.) Deg, Fl. 

Hawaii 1: 62. 1932. Lectotype: Lindley, 

Edward’s Bot. Reg., 26: t. 47. 1840, based on 

a cultivated plant originating from Mexico, 

designated by D.R. Hunt in Davidse et al., Fl. 

Mesoamer. 6: 168. 1994. 

Fig. 34. A-E 

Decumbent stoloniferous herb with elongate 

succulent stems rooting at the nodes, the flowering 

branches becoming ascending or sub-erect. 

Leaves spreading, oblong-lanceolate to narrowly 

elliptic, 8-25 (-28) cm long, 2.5-4.5 cm broad, subsucculent, 

concave, the apex acute, the margins 

entire, the base slightly auriculate at the apex of the 

1 cm tubular sheath. Flowers small, aggregated in 

dense sessile clusters at intervals along an 

apparently naked simple or branched inflorescence 

arising from a normal leafy stem. Bracts whitish, 

ovate or 3-lobed, to 1 cm long. Sepals oblong, 

boat-shaped, hyaline, 4-5.5 mm long; petals white 

to pinkish, oblong-lanceolate, 4.5-5.5 mm long; 

stamens 6, the filaments white, filiform, the 

anthers elliptic, basal on a butterfly shaped white 

connective 2 mm long; ovary 3-locular, sharply 

trigonous, the style to 4 mm long. Capsule 6seeded. 

General distribution: Endemic to Mexico, but 

widely cultivated and naturalized in tropical 

countries, e. g., Jamaica, Bonaire, Hawaii, and 

India. 


Islands: Recorded as naturalized in Aguadilla, 

Camuy, Cayey, Ciales, Humacao, Maricao, and 

Toa Baja; St. Croix, St. John, Tortola, and Guana 

Island 


Ciales: Bo. Frontón, Proctor et al. 51148 (US). 

Humacao: Liogier et al. 31317 (US). ST. CROIX: 

Cotton Valley, just back of Yellow cliff Bay, 

Fosberg 53937 (US). ST. JOHN: Reef Bay Quarter; 

Fish Bay, Acevedo-Rdgz. & Siaca 3870 (JBSD, 

MO, NY, UPR, US). TORTOLA: Belmont Estate, S 

of Belmont Pond, Proctor 44414 (US). 

2. Callisia repens (Jacq.) L., Sp. Pl. ed. 2, 62. 

1762; Hepalanthus repens Jacq., Enum. Syst. 

Pl. 12. 1760. Type: Martinique. Jacquin, 

Select, Stirp. Amer. Hist. t. 11. 1763.


Prostrate, slender, creeping herb, the often 

purplish stems rooting at the nodes and often 

forming mats. Leaves ovate, 1-4 cm long; 1-2 cm 

broad, acute at apex, rounded to subcordate at 

base, sessile, sub-succulent, glabrous except for 

ciliate margins; pale green; sheath tubular, 3-3.5 

mm long, with a few long hairs at apex. Flowering 

branches often somewhat ascending with leaves 

progressively smaller; flowers-clusters barely 

exserted beyond the sheaths in leaf-axils; bracts 

filiform, ciliate, 6-7 mm long. Sepals 3, linearlanceolate, 

greenish, 2-5 mm long, minutely 

pubescent; petals 4, oblong, whitish hyaline, 

slightly shorter than or equaling the sepals; 

stamens exserted, typically 3, but may vary in 

number from 0 to 6; filaments minutely ribbonlike, 

coiled at first, to 10 mm long; anthers 

rounded-elliptic or elliptic, basal on a reniform 

white connective ca. 0.5 mm long. Ovary 2locular, 

pilose at apex, style to 4.5 mm long, the 

stigma trifid (Howard, 1979) or penicillate (Hunt, 

1983). Capsule lenticular, ca. 1.7 mm long, 

splitting from apex to base; seeds no more than 4. 


(Texas, Louisiana, and Florida), Mexico, Central 

and South America, Cuba, Cayman Islands, 

Jamaica, Puerto Rico, Virgin Islands, and the 



Islands: Frequent in moist shady situations or 

sometimes on rocky bands, without any obvious 

soil preferences, at low to upper middle elevations 

(sea level to 830 m). It has been suggested that this 

Commelina L., Sp. Pl. 40. 1753. 

3. COMMELINA 

species owes part of its present range to human 

introduction. It is sometimes planted in rock 

gardens and in hanging baskets. Recorded from 

Cabo Rojo, Coamo, Comerío, Culebra, Manatí, 

Mona Island, Río Grande, Salinas, Vega Baja, and 

Vieques; Guana Island, St. Croix, St. John, and 

Tortola. 

Common names: Puerto Rico: Belleza, 

Cohítre enano, Lenga de gallina. 

Note: Hunt (1963) gives the sepal length in 

Trinidad as 2-3 mm; Howard (1979) gives the 

sepal length in the Lesser Antilles as 3-4 mm; 

while Acevedo-Rodríguez (1996) gives the sepal 

length in St. John, Virgin Islands as 4-5 mm. These 

figures, if accurate, suggest a clinal change 

through the islands. 


Cabo Rojo: Salinas de Cabo Rojo, Sintenis 911 

(US). Culebra: N.L. Britton & Wheeler 35 (US). 

Salinas: Bo. Lapa, upper S slopes of Las Piedras 

Chiquitas, Proctor & Liogier 42738 (US). 

Vieques: Sierra Encantada, Shafer 2543 (US). ST. 

CROIX: Christiansted, Cane Bay Estate, Rose et al. 

3601 (US). pond 1 km NNW of Cathrine’s Rest, on 

Granard Road, Fosberg 54118 (US). Recovery 

Hill, Ricksecker 412 (US). ST. JOHN: Bethany to 

Rosenberg, N.L. Britton & Shafer 222 (NY, US); 

Road to Ajax Peak, Acevedo-Rdgz. & Chinea 2651 

(MO, NY, UPRRP, US, VINPS). TORTOLA: 

Belmont Estate, S of Belmont Pond, Proctor 

44413 (US). Great Mountain, Fishlock 59 (US-2). 

Road Town to High Bush, N.L. Britton & Shafer 

746 (US). 

Perennial or annual herbs with fibrous or tuberous roots; stems decumbent to erect or sometimes 

rhizomatous and bearing cleistogamous flowers. Leaves sessile or short-petiolate. Inflorescence cymose, 

subtended by a spathe-like bract. Flowers zygomorphic, bisexual or polygamous (some flowers with 

stamens only); sepals 3, unequal, free or nearly so, the outermost one hood-shaped; petals 3, free, blue or 

white (in ours), very unequal, the lower one very small or minute (rarely lacking), the upper two clawed 

at base with conspicuous expanded apical portion; stamens 6, three of which are fertile, with ellipsoid or 

saddle-shaped anthers, the others (staminodes) with X-shaped anthers; all filaments glabrous. Ovary 

sessile, glabrous, 2- or 3-locular, with 1 or 2 ovules per locule. Fruit a dry dehiscent capsule; seeds smooth 

or variously marked, with a linear hilum. A cosmopolitan tropical and warm-temperate genus of 230 

(Howard, 1979), 250 (G. C. Tucker, 1989) or 170 (Acevedo-Rodríguez, 1996) species. Whatever the 

correct figure, it is by far the largest taxon in the Commelinaceae. 

LECTOTYPE: Commelina communis L., designated by Britton & A. Brown, Ill. Fl. N. U. S. ed. 2. 1: 457. 

1913.

158 


Key to the species of Commelina 

1. Leaves finely pubescent, ovate with blunt apex; subterranean stolons present, bearing cleistogamous 

flowers ……..........................................................................................................… 1. C. benghalensis 

1. Leaves usually glabrous, lanceolate, narrowly elliptic, or ovate-lanceolate, acute or acuminate at apex; 

stolons with cleistogamous flowers absent. .................................................................……………… 2 

2. Flowers white; fruits indehiscent; leaves up to 17 cm long ……………. 4. C. rufipes var. glabrata 

2. Flowers blue or pale blue (rarely white); fruits dehiscent; leaves 3-10 cm long. ……………… 3 

3. Spathes free with ciliolate margins and without included mucilage; staminodes 2 

…..........................................................................................................……….. 2. C. diffusa 

3. Spathes closed toward base with included mucilage, the margins membranous, not ciliolate; 

staminodes 3 …….......................................................................................…… 3. C. erecta 

1. Commelina benghalensis L., Sp. Pl. 41. 1753, 

nom. conserv. Type: India. (holotype: Linn 

65.16 (BM), typ. conserv. 

Rhizomatous decumbent to sub-erect herb 

with basal leafless stolon-like branches bearing 

cleistogamous flowers. Aerial stems pubescent, up 

to 1 m long, rooting at lower nodes; petioles and 

sheaths ciliate with long pluricellular red hairs. 

Leaf blades 4-8 cm long, 1.5-3 cm broad, narrowed 

and inequilateral at base, freely pubescent on both 

surfaces. Spathes slightly funnel-shaped, auriculate 

on one side., 1-1.5 cm long, minutely pubescent, 

the margins joined toward the base. Sepals small, 

pubescent; petals orbicular, blue. Capsules 3locular, 

5-seeded: seeds minutely roughened and 

pitted. 

General distribution: Tropical Asia, introduced 

in the United States (Florida, Georgia, Louisiana, 

and California), Cuba, Jamaica, Puerto Rico, 

Martinique, Montserrat, St. Vincent, and Barbados. 


single collection from Cayey. 


Cayey: Axelrod et. al. 730 (SJ, UPR). 

2. Commelina diffusa Burm. f., Fl. Indica 18, t. 7, 

f. 2. 1768. Lectotype: India. Burman s.n. (G), 

designated by Merrill, J. Arnold Arbor. 18: 65. 

1937. 

Commelina longicaulis Jacq., Collectanea 3: 234. 

1791 [“1789”]. Lectotype: Venezuela. 

Caracas, Jacquin, Icon. Pl. Rar. 2: t. 294. 

1797, designated by R.A. Howard, Fl. Lesser 

Antill. 3: 432. 1979. 

Commelina cayennensis Rich., Actes Soc. Hist. 

Nat. Paris 1: 106. 1792. Type: French Guiana. 

LeBlond, s. n. (holotype: probably at P). 

Plants annual in areas with seasonal rainfall, 

long-persistent in areas or sites with constant 

moisture. Prostrate or ascending branched herb 

rooting at nodes. Leaves with scarious, ciliate 

sheaths, the blades lanceolate or ovate-lanceolate, 

acute or acuminate at apex, rounded at base. 

Peduncles very short or up to 5 cm long; spathes 

ovate-lanceolate, 1-2 cm long, acute or acuminate; 

cymes 1- to 3-flowered, polygamous; sepals 3-4 

mm long, green, scarious on margins; upper two 

petals 4-5 mm long, bright blue; lower petal much 

smaller or absent. Capsules 3-locular, the upper 

locule with 1 seed, the other two with 2 seeds 

each, seeds black, reticulate. 



Islands: A common weed of roadsides, moist 

ditches, and open waste land at low to high 

elevations (sea level to 1050 m). Recorded from 

Arecibo, Bayamón, Caguas, Cataño, Corozal, 

Guana Island, Guayama, Guayanilla, Manatí, 

Maunabo, Mayagüez, Naguabo, Naranjito, Patillas, 

Ponce, Río Grande, San Juan, Vega Baja, and 

Vieques; St. Croix, St. Thomas, and Tortola. 

Common name: Puerto Rico: Cohítre. 


Arecibo; Río Abajo State Forest, Acevedo-Rdgz. 

10602 (UPRRP). Bayamón: Goll 211, 213 (US). 

Cataño: Heller & Heller 115 (US). Guayama: 

Road from Guayama to Cayey, Underwood & 

Griggs 447 (US). Guayanilla: Guayanilla to 

Tallaboa, Shafer 1979 (US). Manatí: Road from 

Manatí to Vega Baja, Underwood & Griggs 966 

(US). Mayagüez: Road from Mayagüez to 

Joyuda, Underwood & Griggs 139 (US). Ponce:


Juana Mata, Goll 807 (US). Río Grande: 

Caribbean National Forest, Pico del Este, Boom 

7980 (US). San Juan: Río Piedras, Hioram 912 

(US). Vieques: Monte Pirata, Axelrod et al. 4888 

(US). ST. CROIX: Bassin, Ricksecker 56b, 175, 403 

(US). Christiansted, Rose et al. 3576, 3591, 3592 

(US). pond 1 km NNW of Catherine’s Rest, on 

Granard Road, Fosberg 54114 (US). Anna’s 

Hope, Thompson 100 (US). Cotton Valley, just 

back of Yellow cliff Bay, Fosberg 53938, 53939 

(US). East end of Concordia, scenic road, Fosberg 

54024 (US). Eliza’s Retreat, Jakobsberg (Lang’s 

Peak), Fosberg & Hayes 54166 (US). North slope 

Altona-Langford Road, Fosberg 54045 (US). 

Teague Bay, West Indies Lab campus, Fosberg 

53988, 60726 (US). Windsor Hill, Christiansted, 

Seligson 278 (US). ST. THOMAS: Raunkiaer s.n. 

(US). Kuntze s.n. (US). Eggers s.n. (US). Sugar 

Estate, Eggers 178 (US). TORTOLA: Beef Island, 

N.L. Britton & Shafer 884 (US). Experiment 

Station, Fishlock 237 (US). 

3. Commelina erecta L., Sp. Pl. 41. 1753. 

Lectotype: Virginia. Dillenius, Hort. Eltham. 

94, p. 77, t. 88. 1732, designated by C. B. 

Clarke in Alph. de Candolle & C. de Candolle, 

Monogr. Phan. 3: 181. 1881. 

Commelina elegans Kunth in Humb., Bonpl., & 

Kunth, Nov. Gen. Sp. 1[quarto ed.]: 259. 

1816. Type: Colombia. Humboldt & Bonpland 

s.n. (holotype: probably at P). 

Commelina virginica sensu some authors, non L., 

1763. 

Fig. 34. F-I 

Perennial herb with decumbent to erect stems 

arising from clustered, slender, tuberous roots; 

leaves lanceolate, mostly 3-9 cm long (rarely 

longer or shorter), 1-3 cm broad, the apex acute or 

acuminate, usually glabrous but sometimes 

minutely puberulous above. Inflorescences solitary 

or 2-4 clustered at top of stem; peduncles 0.7-2.2 

cm long; spathes broadly ovate, sharply acuminate, 

mostly 2-3 cm long (sometimes slightly longer or 

shorter) glabrous or minutely puberulous; cymes 1 

or 2 (usually only one fertile); flowers 1-3 (-6) per 

cyme; barely exserted beyond spathe, the pedicels 

ca. 3.5 mm long; sepals 4-5 mm long, whitish 

hyaline, the lower 2 connate; upper petals pale to 

deep blue (rarely white), 1-1.5 cm long (rarely 

longer). Fertile filaments curved, white; ovary 

ovoid-globose, the style elongate with minutely 

capitate stigma. Capsules 4-6 mm long, 3-locular, 

1 locule warty, indehiscent, the other two smooth, 

dehiscent, with 1 seed in each locule; seeds smooth 

light brown, ca. 2.7 (-4) mm long. 


States and Mexico south to Argentina; also in 

tropical Africa. 


Islands: Common in disturbed habitats and in dry 

to moist woodlands from low to high elevations 

(sea level to 1300 m). Recorded from Bayamón, 

Cabo Rojo, Cayey, Ciales, Coamo, Desecheo 

Island; Fajardo, Guayama, Manatí, Maricao, 

Mayagüez, Mona Island, Ponce, Río Grande, 

Salinas, Toa Alta, Vega Alta, Vega Baja, Vieques, 

andYabucoa; Anegada, George Dog, Great 

Camanoe, Guana Island, St. Croix, St. John, St. 

Thomas, Tortola, and Water Island. 

Common name: Puerto Rico: Cohítre azul. 


Bayamón: Goll 214 (US). Cayey: 7 mi. S of 

Caguas, Heller & Heller 312 (US). Ciales: 

Johnston 951 (US-2). Coamo: Coamo Springs, 

Goll 643, 663, 671, 683 (US). Desecheo Island: 

N.L. Britton et al. 1585 (US). Guayama: Guayama 

Road, Goll 573 (US). Manatí: Stevenson & 

Johnston 1975 (US). Mayagüez: Sintenis 84 (US). 

Mona Island: vic. Cueva de Doña Gena, Acevedo- 

Rdgz. & Siaca 4395 (FTG, NY, SJ, US). Ponce: 

Juana Mata, Goll 803, 804, 810 (US). Salinas: 

Sabana Llana, Goll 163, 164 (US). Toa Alta Road, 

Puente Flaco, Goll 885 (US-2). Vega Baja: Goll 

1012 (US). Vieques: Isabel Segunda to Sierra 

Encantada, Shafer 2516 (US). Yabucoa: Punta de 

la vaca, Sintenis 5130 (US). ANEGADA: Area 1 km 

W of town, Acevedo-Rdgz. et al. 11507 (US). ST. 

CROIX: Teague Bay, Fosberg 60726 (US). ST. 

JOHN: East End Quarter, E of Southside Pond, 

Acevedo-Rdgz. et al. 1832 (VINPS, NY, US). ST. 

THOMAS: Raphune hill, Acevedo-Rdgz. 11224 (US, 

MAPR, UPRRP). TORTOLA: Beef Island, N.L. 


4. Commelina rufipes var. glabrata (D. R. Hunt) 

Faden & D. R. Hunt, Ann. Missouri Bot. 

Gard. 74: 122. 1987; Commelinopsis glabrata 

D. R. Hunt, Kew Bull. 36: 195. 1981. Type: 

Trinidad. Broadway 6716 (K). 

Tradescantia portoricensis Bello, Anales Soc. 

Esp. Hist. Nat. 12: 122. 1883. Type: Puerto 

Rico. Bello s.n. (destroyed).

160 


Fig. 34. A-E. Callisia fragrans. A. Habit. B. Inflorescence unit. C. Flower. D. Stamen. E. Pistil. F-I. Commelina erecta. F. 

Flowering branch. G. Flower. H. Inflorescence with subtending bract. I. L.s. flower. J-L. Tradescantia zebrina. J. Habit. K. 

Flower, top view. L. Stamen. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78.


Athyrocarpus persicariifolius sensu Britton and P. 


Commelinopsis persicariifolia Delile, 1816. 

Sprawling, decumbent to erect, sub succulent 

perennial herb, rooting freely at the lower nodes, 

the erect portions of stems up to 1 m tall. Leaves 

lanceolate or oblong-lanceolate, 8-19 cm long, 

1.5-4 cm broad, acuminate at base, unequally 

cuneate at base, glabrous or nearly so; sheaths 

closed. Peduncles 1-4 at apex of stem, less than 1 

cm long; spathes 2-4 cm long, broader than long, 

the margins free; cymes 2, the lower with (0)1-3 

flowers, these staminate; upper cyme with 2-10 or 

more bisexual flowers; sepals 5 mm long; 2-3 mm 

broad; upper petals white, long-clawed; fertile 

stamens long, staminodes short. Capsules ellipsoid, 

5.5-7 mm long, crustaceous, shining grayish 

white; anterior loculi 2-seeded; posterior loculus 

containing 1 large seed. 

General distribution: Greater Antilles (except 

Jamaica), Grenada, Trinidad, and northern South 

4. GIBASIS 

Gibasis Raf., Fl. Tellur. 2: 16. 1837 [“1836”]. 

America. 

Distribution in Puerto Rico: Frequent on 

moist to wet forest slopes (mostly non-calcareous) 

at middle to upper middle elevations (100-830 m). 

Recorded from Aguada, Dorado, Las Marias, 

Luquillo, Maricao, Naguabo, Quebradillas, Río 

Grande, Salinas, Utuado, and Yabucoa. 

Note: The closely related var. rufipes occurs 

only in South America. A large-leaved variant in 

Grenada and Trinidad has been suggested to be 

taxonomically distinct. 


Dorado: grounds of Dorado Beach Hotel, Maas 

6464 (US). Las Marias: La Juanita, N.L. Britton et 

al. 3911 (US). Maricao: coffee plantation, Liogier 

10773 (US). Río Grande: El Verde Research 

Station, Boom 10059 (US). Río Grande: Sierra de 

Luquillo, Caribbean Natl. Forest, along W side of 

Río Mameyes, Proctor 44484 (US). Utuado: Bo. 

Don Alonso, Acevedo-Rdgz. 10534 (JBSD, 

MAPR, NY, UPRRP, US). 

Annual or perennial herbs, the stems creeping, ascending, or tufted, the roots often with tubers. 

Leaves linear to ovate or oblanceolate, often pubescent or pilose with glandular hairs. Inflorescences 

terminal and in the axils of upper leaves, branched to form panicles or corymbs, these few- or manyflowered; 

bracts small but leaf-like. Flowers pedicellate, actinomorphic, bisexual, borne in abbreviated, 

ebracteate cincinni, aggregated in pairs or small umbels. Sepals and petals 3, free, the sepals persistent in 

fruit; stamens 6, all fertile or 3 fertile and 3 sterile; filaments usually pubescent and the anthers broadly 

triangular. Ovary 3-locular, with 2 ovules per locule; stigma minutely capitate. Capsules apically 

dehiscent with 3 valves; seeds rugose and with linear hilum. A genus of 6 species, widely distributed in 

tropical America except for one occurring in India. A single species is found in Puerto Rico. 

TYPE: Gibasis pulchella (Kunth) Raf. (≡ Tradescantia pulchella Kunth). 

1. Gibasis geniculata (Jacq.) Rohweder, Abh. 

Auslandsk., Reihe C. Naturwiss. 18: 143. 

1956; Tradescantia geniculata Jacq., Enum. 

Syst. Pl. 18. 1760; ; Aneilema geniculata 

(Jacq.) Woodson, Ann. Missouri Bot. Gard. 

29: 147. 1942. Lectotype: Martinique. 

Plumier (Burman ed.), Pl. Amer., t. 116, fig. 2. 

1757., designated by R.A. Howard, Fl. Lesser 

Antill. 3: 438. 1979. 

Figs. 33. F-I; 63. B 

Perennial, trailing-ascending, diffuselybranched 

herb, rooting at the decumbent lower 

nodes. Leaves ovate to ovate-lanceolate, mostly 3- 

7 × 1-3 cm, acute at apex, rounded to cordate at 

base, dark glossy green on upper side, often tinged 

purplish beneath, usually sparsely pubescent on 

both sides. Peduncles glabrous 2.5-5 cm long, the 

inflorescence loosely cymose or seemingly 

paniculate; pedicels filiform, 4-12 mm long; 

sepals lanceolate, green, 2-3 mm long, glabrous or 

nearly so; petals ovate, 3-5 mm long, white; 

filaments 2.5-3 mm long, villous toward base. 

Capsules 2-3 mm long; seeds 1 mm long, gray, 

obscurely rugose. 

General distribution: widespread in the

162 


Neotropics, but in the West Indies confined to 

Hispaniola, Puerto Rico, and the Lesser Antilles. 

Distribution in Puerto Rico: Frequent in 

moist wooded sites at low to rather high elevations 

(25-1000 m). This species grow on both 

calcareous and non-calcareous soils. Recorded 

from Adjuntas, Aibonito, Arecibo, Caguas, 

Cayey, Ciales, Dorado, Fajardo, Florida, Guayama, 

Isabela, Jayuya, Luquillo, Maricao, Mayagüez, 

Naguabo, Salinas, San Juan, Utuado, Vega Alta, 

Yabucoa, and Vieques. 

Common name: Puerto Rico: Cohítre azul. 


Arecibo: Río Abajo State Forest, along Juan Ruiz 

5. MURDANNIA 

Murdannia Royle, Ill. Bot. Himal. Mts. 403, t. 95. f. 3. 1840, nom. conserv. 

trail, Acevedo-Rdgz. 10686 (US). Ciales: along 

trail Camino de la Ceiba towards Quebrada del 

Pozo Azul, Acevedo-Rdgz. & Vicens 11881 

(MAPR, NY, UPRRP, US). Guayama: Bo. 

Carmen, Proctor & Rivera 46940 (US). Maricao: 

Bosque Estatal de Maricao, Acevedo-Rdgz. et al. 

7696 (F, MO, NY, US). Naguabo: Eggers 629 

(US). Salinas: Bo. Lapa, Proctor & Thomas 44153 

(US). San Juan: Río Piedras, Stevenson 208 (US). 

Utuado: Bo. Don Alonso, Acevedo-Rdgz. et al. 

13409 (US). Vega Alta: Bo. Sabana, Proctor 

42899 (US). Vieques: Sierra Encantada, Shafer 

2562 (US). 

Annual or perennial prostrate or ascending herbs; stems often elongate and rooting at nodes; leaves 

various. Cymes aggregated in panicles or verticils, or else solitary; flowers zygomorphic; sepals 3, equal, 

free; petals 3, subequal, free, stamens 3-6. the outer whorl usually fertile, or consisting of one staminode, 

or absent; staminodes hastate or 3-lobed at apex. Capsules subequally 3-valved; seeds with punctiform or 

linear hilum. A tropical Asiatic and African genus of 50 species, a single one of which has become 

naturalized in the Neotropics. 

TYPE: Murdannia scapiflora (Roxb.) Royle (≡ Commelina scapiflora Roxb.) 

1. Murdannia nudiflora (L.) Brenan, Kew Bull. 

7: 189. 1952; Commelina nudiflora L., Sp. Pl. 

41. 1753. Lectotype: China or Java. Osbeck 2 

(LINN-65.12), designated by Merrill, J. 

Arnold Arbor. 18: 64. 1937. 

Fig. 35. A-F 

Prostrate or decumbent sub-succulent annual 

herb, rooting at nodes; leaves linear to narrowly 

lanceolate, usually glabrous, 3-13 × 0.1-0.9 cm, 

acute to subacuminate at apex, slightly clasping at 

the narrowed base. Inflorescences short- to longpedunculate, 

the slender peduncles 2-15 cm long; 

cymes glabrous, usually solitary with ca. 5-8 

flowers; pedicels to 3 mm long at anthesis, 

increasing to 4-5 mm long in fruit. Sepals 2 mm 

long; petals ca. 2 mm long; usually bluish or 

purplish pink; fertile stamens 2, the sterile ones 2- 

4; filaments bearded. Capsules 3-4 mm long, the 

valves apiculate, persistent. 

General distribution: Tropical Asia and 

Africa, adventive as an aggressive weed at 

scattered localities in the Neotropics and in 

Florida. 

Distribution in Puerto Rico: Weedy herb of 

wet open or disturbed situations from low to upper 

middle elevations (5-800 m), locally frequent. 

Apparently occurs only on acidic or noncalcareous 

soils. Recorded from Loíza, Luquillo, 

Naguabo, Río Grande, and Vega Alta. 

Note: The flowers are said to open only 

between the hours of 11:00 a.m. to 1:00 p.m. 

(Hunt, Fl. Trin. & Tob., 1985). 


Luquillo: Bo. Sabana, Caribbean Natl. Forest, 

along Río del Cristal, Axelrod & Axelrod 7260 

(US). Vega Alta: Bo. Sabana, Estáncias del Mar, 

Proctor 46076 (US).


Fig. 35. A-F. Murdannia nudiflora. A. Habit. B. Flower. C. Capsule. D. Pistil. E. Stamen. F. Seed. G-N. Tripogandra serrulata. 

G. Habit. H. Inflorescence. I. Flower. J. Sterile stamen. K. Fertile stamen. L. Pistil. M. Capsule. N. Seed. (A, from Proctor 

46076; B-F, from Proctor 50169; G-N, from Proctor 49728).

164 

Tradescantia L., Sp. Pl. 288. 1753. 


6. TRADESCANTIA 

Perennial herbs of diverse habit, with fibrous or tuberous roots and simple or branched stems. Flowers 

bisexual, actinomorphic, in terminal or axillary umbel-like cymes subtended and often more or less 

enclosed by paired, sometimes boat-shaped bracts, these either similar to or strongly differentiated from 

ordinary leaves. Sepals 3, free, (rarely united), rarely accrescent and fleshy in fruit; petals 4, variously 

colored or white, usually free, sessile or sometimes clawed, rarely united into a slender tube; stamens 6, 

all similar, equal, and fertile or occasionally subequal; filaments glabrous or bearded, the antherconnectives 

broad, versatile, ovary 3-locular, with usually 2 ovules per locule; stigma minutely capitate. 

Capsules 3-valved; seeds rugose-reticulate, with usually a linear hilum. An American genus of more than 

60 species occurring in both tropical and temperate areas. All the species recorded from Puerto Rico and 

the Virgin Islands are introduced ornamentals except for T. zanonia, and each one was formerly placed in 

a different genus. The introduced species have escaped from cultivation and have become more or less 

naturalized. 

TYPE: Tradescantia virginica L. 

Key to the species of Tradescantia 

1. Stems elongate, creeping or decumbent-ascending; flowers pink to rose-purple. ……............……… 2 

2. Leaves violet-glaucous, not striped; petals free …………...................................…. 1. T. pallida 

2. Leaves bicolorous with broad longitudinal stripes of green and silver on upper side, usually plain 

purplish beneath; petals united at base into a tube ……............................……….. 4. T. zebrina 

1. Stems erect (occasionally or partly decumbent); flowers white. ……....................................……….. 3 

3. Stems concealed by a dense leaf-rosette; leaves usually purple beneath (rarely green); sepals pale 

green, remaining thin and membranous in fruit ………….............................…..2. T. spathacea 

3. Stems evident, not concealed by leaves, up to 1 m tall; leaves pale green beneath; sepals becoming 

purplish black and fleshy in fruit ……………................................................…… 3. T. zanonia 

1. Tradescantia pallida (Rose) D. R. Hunt, Kew 

Bull. 30: 452. 1975; Setcreasea pallida Rose, 

Contr. U. S. Natl. Herb. 13: 294. 1911. Type: 

Mexico; Tamaulipas, near Victoria. Palmer s. 

n. (holotype: US!; isotype US!). 

Setcreasea purpurea Boom, Acta Bot. Néerl. 4: 

167. 1955. Type: Mexico. Sent by C.A. 

Purpus to Leyden where it was cultivated and 

later vouchered by Boom 28046 (holotype: L). 

Perennial herb with elongate decumbentascending 

stems with sub-erect flowering branches. 

Leaf sheaths clasping, 1-2.5 cm long, ciliate; 

blades oblong, 10-18 cm long, 2-3.5 cm broad, 

acute at apex, the upper one somewhat smaller. 

Inflorescences terminal and in upper leaf-axils; 

peduncles mostly 3-10 cm long; flowers in small 

densely cymose clusters subtended by 2 or 3 

bracts, these similar to leaves but smaller; pedicels 

umbellate, to 7 mm long, pilose toward apex; 

sepals oblong, 8-10 mm long; petals pink to rose- 

purple, 15-20 mm long; stamens 6, with 3 

filaments epipetalous and 3 adherent to petal 

margins, glabrous or variably pubescent. Fruit 

unknown, the plants always propagated by 

cuttings. 


(Florida and Louisiana), Mexico and Central 

America; widely grown as an ornamental 

throughout the tropics. 


Islands: Often seen in Puerto Rico as a cultivated 

plant, but voucher specimens not collected. In the 

Virgin Islands it has been collected on Guana 

Island, St. Croix, and St. Thomas as an escaped 

plant becoming naturalized. 

Common name: Puerto Rico: Cohítre morado. 


West Indies Laboratory, Teague Bay, Fosberg 

55281 (US). ST. THOMAS: Hull Bay road, Acevedo- 

Rdgz. 11354 (MAPR, US).


2. Tradescantia spathacea Sw., Prodr. 57. [June- 

July] 1788; Rhoeo spathacea (Sw.) Stearn, 

Baileya 5: 195. 1957. Type: Jamaica. Swartz 

s.n. (holotype: S-Sw, R-6172). 

Tradescantia discolor L’Hér., Sert. Angl. 8, t. 12, 

[late 1788 or Jan] 1789; Rhoeo discolor 

(L’Hér.) Hance ex Walp. Ann. Bot. Syst. 3: 

660. 1853. Type: L’Hériter, Sert. Angl. t. 12, 

late 1788 or Jan 1789, based on cultivated 

material. 

Erect perennial sub-succulent herb with short 

stout erect stems to 20 cm tall, often clustered and 

forming large colonies. Leaves imbricate, crowded, 

sub-erect , linear-lanceolate to oblong-lanceolate, 

20-35 cm long, mostly 3-5 cm broad, acuminate at 

apex, scarcely narrowed at base above the sheath, 

usually dark green above, red-purple beneath. 

Inflorescences axillary; peduncles 2-4.5 cm long, 

simple or branched; bracts deeply boat-shaped, 

broadly ovate, 2-4.5 cm long, 2.5-5 cm broad. 

Flowers numerous, scarcely exserted; pedicels ca 

1.5 cm long, recurved in fruit; sepals ovatelanceolate, 

3-4 mm long; petals free, white, very 

broadly ovate, 5-8 × 5-7 mm broad. Stamens free; 

filaments 4-7 mm long, bearded toward base; 

ovary with 1 ovule in each locule. Seeds oblongellipsoid 

with linear hilum. 

General distribution: Mexico to Guatemala 

and Belize; widely cultivated in tropical countries, 

easily escaping and often becoming naturalized. 

Found almost throughout the Greater and Lesser 

Antilles. 


Islands: Recorded from Cataño, Manatí, 

Quebradillas, San Juan (cult.), and Yabucoa 

(cult.); St. John. This species occurs much more 

widely but voucher specimens are seldom 

collected. 

Common names: Puerto Rico: Sangría, 

Sanguinaria. 


Cataño: Goll 971 (US). Manatí: Stevenson 549 

(US). Quebradillas: Sargent B19 (US). San Juan: 

Río Piedras, Stevenson s.n. (US). Yabucoa: 

Sintenis 5054 (US). ST. JOHN: On trail to Margaret 

Hill, Acevedo-Rdgz. 2310 (US). 

3. Tradescantia zanonia (L.) Sw., Fl. Ind. Occid. 

1: 604. 1797; Commelina zanonia L., Sp. Pl. 

41. 1753; Campelia zanonia (L.) Kunth in 

Humb. Bonpl. & Kunth, Nov. Gen. Sp. 1 

[quarto ed.]: 264. 1816. Lectotype: French 

Guiana. Plumier, Nov. Pl. Amer. T. 38. 1703, 

designated by D.R. Hunt in Davidse et al., Fl. 

Mesoamer. 6: 161. 1994. 

Fig. 33. J-O 

Perennial herb with thick, succulent stems, 

these erect or partly decumbent, up to 1 m tall or 

more, often unbranched, exuding glutinous 

mucilage when broken. Leaves lanceolate to 

oblanceolate, 10-18 (-30) cm long, 3-5 (-8) cm 

broad, acuminate at apex, narrowed at base, dark 

green above, paler beneath, glabrous. 

Inflorescences axillary, the peduncles simple or 

branched, up to 20 cm long; bracts 2, often 

unequal, up to 3 (-6) cm long; pedicels up to 6 mm 

long; sepals 3-5 mm long, cucullate, accrescent 

and enclosing the mature fruit in a black, juicy 

envelope; petals free, broadly obovate-quadrate, 

6-10 mm long, 5-8 mm broad, usually white; 

stamens free, filaments 4-8 mm long, glabrous or 

bearded; anther-connectives sagittate. Seeds 

black, smooth, lustrous, flat, with linear hilum. 

General distribution: Continental Neotropics 

from Mexico to Brazil, including Trinidad and the 

Greater Antilles, but absent from the Lesser 

Antilles. 

Distribution in Puerto Rico: Occurs in moist 

sheltered woodlands and thickets at middle to 

upper elevations (200-890 m). Recorded from 

Adjuntas, Arecibo, Cayey, Ciales, Coamo, 

Orocovis, Ponce, Salinas, Utuado, and Yauco. 

Common name: Puerto Rico: Cohítre blanco. 


Adjuntas: Sintenis 4207 (US-2). Between Alto de 

la Bandera and Jayuya, Cowell 5491 (US). 

Arecibo: Bo. Río Arriba, just SE of Río Abajo 

Forest Reserve boundary, Axelrod & Ward 8819 

(US). Ciales: Bo. Toro Negro, Axelrod & Stenzel 

11134 (US). Coamo: Sintenis 3126 (US). Ponce: 

Bo. San Patricio, from S end of Rt. 515 along path 

of proposed Rt. 10, Axelrod et al. 8965 (US). 

Salinas: Bo. Lapa, SE base of Las Tetas de Cayey, 

Proctor 44154 (SJ). Yauco: Bo. Sierra Alta, 

Proctor & Díaz 44741 (US). 

4. Tradescantia zebrina Heynh., Alp. Aufz. Gew. 

735. 1847. Type: Not known, based on 

material cultivated in Europe. 

Zebrina pendula Schnizl., Bot. Zeit. 7: 870. Dec.

166 


1849; Tradescantia pendula (Schnizl.) D. R. 

Hunt, Kew Bull. 36: 197. 1981. Type: 

Unknown. 

Fig. 34. J-L 

Decumbent, branched, long trailing subsucculent 

herb, rooting at nodes, the stems 

glabrous or sometimes pilose. Leaves usually 

glabrous, broadly to narrowly or oblong-ovate, 3- 

8 cm long, 1.5-3.2 cm broad, acute to acuminate at 

apex, rounded-inequilateral at base, the sheaths 

villous, the upper broadly striped bluish green and 

silver, or the silvery portions replaced partly or 

wholly by purple, the underside uniformly reddish 

purple. Peduncles short to elongate, up to 12 cm 

long; flowers clustered between two unequal, 

imbricate, leaf-like bracts, the outer bract 2-5 cm 

long, the inner 1-2 cm long; pedicels to 3 mm; 

calyx white, 5-8 mm long, irregularly lobed, 

tubular toward base; corolla-tube white, 6-10 mm 

long, 1.3 mm in diam., the lobes ovate, 5-10 mm 

long, 3-7 mm broad, usually bright pink (rarely 

violet-blue); stamens epipetalous, subequal, the 

hairy filaments 3-5 mm long; ovary glabrous, 3locular, 

with 2 ovules in each locule. Capsules 

dehiscent, gray-brown; seeds with punctiform 

7. TRIPOGANDRA 

Tripogandra Raf., Fl. Tellur. 2: 16. 1837 [“1836”]. 

hilum. 

General distribution: United States (Florida, 

Louisiana, and Kentucky), Mexico and Central 

America; introduced and naturalized throughout 

the West Indies and South America. 


Islands: Naturalized on shady banks and in moist 

secondary woodlands from near sea level to almost 

900 meters. Recorded from Adjuntas, Aibonito, 

Arecibo, Arroyo, Bayamón, Carolina, Ciales, 

Lajas, Maricao, Morovis, Naguabo, Orocovis, San 

Germán, San Juan, Utuado, and Yauco; St. Croix, 

St. John, and Tortola. 

Common names: Puerto Rico: Cohítre 

morado, Judío errante. 


Adjuntas: Sintenis 4153 (US). Aibonito: Heller & 

Heller 892 (US). Arecibo: Río Abajo State Forest, 

Acevedo-Rdgz. 10589 (US). Ciales: Bo. Frontón, 

Proctor et al. 51147 (US). Maricao: Monte 

Montoso, N.L. Britton & Cowell 4150 (US). San 

Germán: Sargent 58 (US). San Juan: Río Piedras, 

Stevenson 1398 (US). ST. CROIX: Bassin, 

Ricksecker s.n. (US); Ricksecker & Ricksecker 

141 (US). ST. JOHN: Cruz Bay Quarter, Center Line 

Road, Acevedo-Rdgz. & Siaca 4250 (US). 

Annual or perennial sub-succulent herbs with fibrous roots; stems simple or branched; trailing to 

erect, rarely scandent, often bearing long adventitious roots. Leaves with sheating bases, the blades linear 

to ovate. Inflorescences axillary, long pedunculate; flowers bisexual, zygomorphic, borne in small paired 

and fused cincinni not subtended by evident bracts but borne on a common peduncle, these double cincinni 

variously clustered or aggregated. Sepals 3, free; petals 3, free; stamens 6, dimorphic, the outer 3 shorter, 

the inner 3 longer; filaments glabrous or pilose, the longer filaments curved in order to be erect in front of 

the upper petal. Ovary 3-locular with 2 ovules in each locule; stigma more or less capitate. Capsules 3locular, 

with 1 or 2 seeds in each locule; seeds original, reticulate, with punctiform or linear hilum. A 

Neotropical genus of about 20 species, one of which commonly occurs in Puerto Rico. 

TYPE: Tripogandra multiflora Raf. ( Tradescantia multiflora Jacq. 1790, non Sw. 1788). 

1. Tripogandra serrulata (Vahl) Handlos, 

Baileya 17: 33. 1970. Commelina serrulata 

Vahl, Eclog. Amer. 2: 4. 1798. Type: 

probably from Montserrat. Ryan s. n. 

(holotype: C-Vahl). 

Tradescantia elongata G. Mey., Prim. Fl. Esseq. 

146. 1818. Tripogandra elongata (G. Mey.) 

Woodson, Ann. Missouri Bot. Gard. 29: 152. 

1942. Type: Guyana. collector unknown 

(holotype: probably at GOET). 

Figs. 35. G-N; 63. C 

Perennial gregarious herb, the trailing lower 

stems purplish and rooting at nodes; flowering 

stems erect, up to 1 m tall or more, glabrous. 

Leaves lanceolate to oblong-lanceolate, or 

sometimes nearly ovate, 3-12 cm long, 1.5-3 cm 

broad, acute to acuminate at apex, narrowed and


slightly oblique at base, the margins minutely 

serrulate, the upper surface sometimes purplestreaked; 

sheaths ciliate. Inflorescences terminal 

and in the uppermost leaf-axils; peduncles up to 5 

cm long, glabrous; pedicels 1-5 mm long; sepals 

elliptic, 3-5 mm long, glabrous; petals pink, pale 

pink, or rarely white, ovate-elliptic, 3.5-6 mm 

long, 2.3-4.3 mm broad; stamens with villous 

filaments, the anthers dimorphic. Capsules 3 mm 

long; seeds 1-1.5 mm long, gray or grayish brown, 

reticulate. 


America, northern South America, Hispaniola, 

Puerto Rico, Guadeloupe, Dominica, Martinique, 

St. Vincent and Trinidad. 

Distribution in Puerto Rico: Frequent in 

moist to wet, sheltered, non-calcareous habitats at 

low to upper middle elevations (near sea level to 

700 m). Recorded from Adjuntas, Arecibo, 

Arroyo, Caguas, Ciales, Corozal, Maricao, 

Maunabo, Río Grande, San Juan, Trujillo Alto, 

and Utuado. 


Adjuntas: 7 mi. N from Adjuntas, Grant & Rundell 

93-02295 (US). Arecibo: Río Abajo State Forest, 

Acevedo-Rdgz. 10676 (US). Maricao: Bosque 

Estatal de Maricao, Acevedo-Rdgz. et al. 7697 

(NY, US). Río Grande: Acevedo-Rdgz. et al. 

10760 (MAPR, UPRRP, US); Sierra de Luquillo, 

Proctor & Thomas 44496 (US); Bo. Herreras, 

Proctor & Thomas 45326 (US). San Juan: Heller 

& Heller 662 (US). 

Family 21. HAEMODORACEAE Bloodroot Family 

Haemodoraceae R. Br., Prodr. 299. 1810, nom. conserv. 


Perennial herbs with rhizomes or tubers, sometimes stoloniferous, the rhizomes often containing a red 

latex (colored by haemocorin, an arylphenalenone). Leaves linear or lance-linear, parallel-veined, all 

basal or sometimes distichous along erect stems, equitant, and sheathing at base, glabrous or pubescent. 

Inflorescences terminal on erect stems, cymose, in racemes or panicles; flowers regular, bisexual; perianth 

either 6-parted with separate parts, or 6-lobed above and tubular below, persistent; stamens 6 or 3 with free 

filaments; ovary inferior or superior, 3-locular, with few or many ovules; styles slender with small 

stigmas. Fruit a loculicidal 3-valved capsule; seeds few to many, tuberculate, hairy or glabrous, with 

abundant starchy endosperm. A family of 14 genera and ca. 80 species (Maas & Maas-van de Kamer, 

1993), widely distributed in South Africa, Australia, and in temperate and tropical America. 

TYPE: Haemodorum Sm. 

References: Howard, R. A. 1979. Flora of the Lesser Antill. 3: 467-469. Maas, P. J. M. & H. Maasvan 

de Kamer. 1993. Fl. Neotrop. Monogr. 61: 1-44. Maas, P. J. M. & H. Maas-van de Kamer. 1993. Fl. 

Guianas, fasc. 15, 198. Haemodoraceae. Pp. 103-112. 

1. XIPHIDIUM 

Xiphidium Aubl., Hist. Pl. Guiane 33. 1775. 

Stoloniferous herbs with creeping rhizomes and erect, sub-succulent, leafy stems. Leaves equitant, 

distichous, sheathed at base. Inflorescence terminal, paniculate with the flowers secund in two rows on the 

branches. Perianth 6-parted with spreading lobes. Stamens 3, with short filaments; anthers oblong; ovary 

superior, 3-locular, with numerous ovules; styles filiform with apical stigmas. Capsules subglobose, with 

numerous seeds. A Neotropical genus of 2 species, one (the type) widespread, the other endemic to 

western Cuba. 

TYPE: Xiphidium caeruleum Aubl.

168 


1. Xiphidium caeruleum Aubl., Hist. Pl. Guiane 

33. 1775. Lectotype: French Guiana. Aublet, 

Hist. Pl. Guiane t. 11, 1775, designated by 

Maas & Maas-van de Kamer, Fl. Neotrop. 

Monogr. 61: 27. 1993. 

Ixia xiphidium Loefl., Iter Hispan. 179. 1758; 

Xiphidium floribundum Sw., Prodr. 17. 1788; 

Fl. Ind. Occid. 1: 80. 1797, nom. Illeg.; 

Xiphidium albidum Lam., Tabl. Encycl. 1: 

131. 1791, nom. Illeg. Type: Venezuela; 

Macarapa. Loefling s. n. (holotype: probably 

at LINN). 

Figs. 36. A-H; 63. D 

Rhizomes horizontally creeping, up to 20 cm 

long and 1 cm thick, bearing slender stolons up to 

55 cm long, rooting at nodes; roots fibrous. Erect 

stems mostly 20-60 cm tall, sometimes taller. 

Leaves linear-lanceolate, glabrous, 45-60 cm 

long, mostly 1.5-5 cm wide (sometimes wider), 

acuminate at apex, clasping at base, the margins 

finely serrulate. Inflorescence a thyrse up to 30 cm 

long or more, the branches (cincinni) 5- to 25flowered, 

simple or sometimes once-branched; 

main axis and branches minutely pubescent; 

pedicels 2-3 mm long; perianth-segments oblonglanceolate, 

to 6.5 mm long, white to yellowish 

white, sometimes pale bluish within. Small 

deciduous plantlets sometimes produced in axils 

of inflorescence-bracts. Capsules 5-6 mm in 

diam., maturing shiny orange, then turning black; 

seeds many, black. 


America, Greater and Lesser Antilles, Trinidad, 

and tropical South America. 


Islands: Commonly cultivated as a garden plant, 

scaped and apparently naturalized in moist shaded 

banks and forest slopes at low to middle elevations 

(90-500 m). It is more widespread than the few 

collections indicate. Recorded from Caguas, 

Mayagüez, Río Grande, and San Germán. The 

only Virgin Islands record is from a cultivated 

plant collected in St. Thomas in 1923. 

Common names: Puerto Rico: Cola de 

paloma, Mano poderosa. 


Río Grande: Sierra de Luquillo, Caribbean Natl. 

Forest, along W side of Río Mameyes 0.2-0.5 km 

above road 988 bridge, Proctor 44483 (US). 

Family 22. PONTEDERIACEAE Water-hyacinth Family 

Pontederiaceae Kunth in Humb., Bonpl., & Kunth, Nov. Gen. Sp. 1 [quarto ed.]: 265. 1816, nom. conserv. 


Annual or perennial aquatic herbs of fresh water habitats, floating or rooted in mud, rhizomatous or 

stoloniferous, with sympodial branching. Stems usually short, often somewhat spongy or succulent. 

Leaves distichous, petiolate, the petioles vaginate, sheathing at base; stipules often present. Inflorescences 

terminal or pseudolateral, spicate, racemose, or paniculate, each subtended by a spathe-like leaf sheath. 

Flowers bisexual, actinomorphic or zygomorphic with a perianth of 6 imbricate segments, these free or 

basally connate. Stamens 3 or 6, rarely 1, inserted on the perianth, often unequal, with free filaments. 

Ovary superior, 3-locular with axile placentation or 1-locular with parietal placentation; ovules numerous 

or solitary; styles slender, the stigmas 3- to 5-lobed. Fruit a 3-valved capsule or indehiscent nutlet; seeds 

numerous, usually ribbed. A small pantropical family of 7 genera and ca. 30 species. Two genera are found 

in Puerto Rico. 

TYPE: Pontederia L. 

References: Barrett, S.C.H. 2004. Pontederiaceae. Pp. 474-476. In: Smith, N. et al. (eds.) Flowering 

Plants of the Neotropics, Princeton University Press, Princeton. Horn, C. 1994. Fam.197. Pontederiaceae. 

Pp. 91-102. In: Fl. Guianas, fasc. 15. Horn, C. 2002. Fam. 220. Pontederiaceae. Pp. 37-46. In: Flora of 

North America Editorial Committee (eds.), Fl. North Amer., vol. 26.


Fig. 36. A-H. Xiphidium caeruleum. A. Plant with inflorescence and detail of leaf margin. B. Part of inflorescence. C. Top view 

of flower. D. Lateral view of flower with three tepals removed. E. Dorsal and frontal views of stamens. F. Infructescence. G. Part 

of infructescence and transverse section of fruit. H. Seed. From Mori, S. et al. 1997. Vascular plants of central French Guiana. 

Mem. NYBG Vol. 76(1).

170 



1. Flowers blue or violet, showy in one species; stamens 6; petioles sometimes swollen …… 1. Eichhornia 

1. Flowers white or blue, small; stamens 3; petioles not swollen ……...................…….. 2. Heteranthera 

1. EICHHORNIA 

Eichhornia Kunth, Eichhornia. 1842, nom. conserv. 

Floating or rooted herbs. Leaves often with swollen petioles and usually with broadly expanded 

blades (some forms outside our area may have linear submerged leaves). Inflorescences apparently 

terminal, the flowers in spikes, racemes or panicles. Flowers with perianth segments in two series, united 

below, often large and showy; stamens 6, irregularly adnate to the perianth, 3 of them exserted and 3 

included; filaments flattened at the base; ovary 3-locular with numerous ovules. Fruit a capsule included 

in the withered perianth. Seven species distributed in the Neotropics, one in Africa. 

TYPE: Eichhornia azurea (Sw.) Kunth (≡ Pontederia azurea Sw.), typ. conserv. 

Key to the species of Eichhornia 

1. Plant floating; leaves in a basal pseudo-rosette, petioles usually swollen; inflorescence 4- to 15flowered, 

to 15 cm long; flowers with central distal lobe with a dark blotch in center and yellow spot 

within blotch ……...........................................................................................….. 1. E. crassipes 

1. Plant rooted; leaves produced at intervals along stem, not in a basal pseudo-rosette, petioles not swollen; 

inflorescence 2- to 4-flowered, 2-5 cm long; flowers with lobes lacking a blotch with yellow spot 

………….........................................................................................….. 2. E. diversifolia 

1. Eichhornia crassipes (Mart.) Solms in Alph. de 

Candolle & C. de Candolle, Monogr. Phan. 4: 

527. 1883; Pontederia crassipes Mart., Nov. 

Gen. Sp. Pl. 1: 9, t. 4. 1823; Piaropus 

mesomelas Raf., Fl. Tellur. 2: 81. 1837, nom. 

illeg. Lectotype: Brazil; Bahia. Martius 60 

(M), designated by C.N. Horn in Görts-van 

Rijn, Fl. Guianas fasc. 15, Fam. 197. 

Pontederiaceae: 94. 1994. 

Fig. 63. E 

Aquatic perennial, typically free-floating or 

sometimes rooting in mud. Flowering stems to 25 

cm long, erect, often bending over after flowering. 

Leaves in a basal pseudo-rosette, the petiolate ones 

floating or emersed; leaf petioles inflated below 

the middle, or at least slightly swollen, 3.5-35 cm 

long; blades of petiolate leaves leathery, ovate, 

orbicular or reniform, 2.5-16 × 3.5-14 cm, flat or 

concave-convex, the apex short-cuspidate, 

acuminate or rounded at base. Inflorescence 

spicate, densely or sub loosely 4- to 15-flowered, 

to 15 cm long; spathes obovate, 4-11 cm long; 

peduncle 5-12.5 cm long, glabrous. Flowers 

opening individually within 2 hours after sunrise, 

wilting by night; perianth blue or mauve-blue, 

limb lobes obovate, 1.6-4 × 1.5-2.5 cm, margins 

entire, central distal lobe with dark blotch in center 

and yellow spot within blotch; perianth tube 1.5-2 

cm long, pubescent with glandular-tipped hairs; 

anthers 1.7-2.1 mm long; ovary 3-6 mm long, 

glabrous; style ca. 2 cm long, 3-lobed, pilose. 

Capsule 1.5 cm long; seeds columnar, 1.1-2.1 × 

0.6-0.9 mm, horizontally striate, 11- to 14-winged. 

General distribution: Introduced in the 

tropics and subtropics worldwide, native to 

Northern South America. 


Islands: Floating in water of lakes, ponds, and 

streams. Recorded from Coamo, Lajas, Loíza, 

Mayagüez, Rincón, San Juan, Utuado, and Yauco; 

St. Croix. 

Common names: Puerto Rico: Flor de agua, 

Jacinto de agua. 


Coamo: Coamo Dam, Liogier et al. 29454 (US). 

Lajas: Laguna Cartagena, Caudales s.n. (UPR). 

Loíza: Vacia Talega area, Woodbury et al. s.n. 

(UPR). Mayagüez: Sintenis 489 (US). Rincón: 

Cayures, Caudales s.n. (UPR). San Juan: Río 

Piedras, Stevenson 1104 (US). Utuado: Dos 

Bocas, Caudales s.n. (UPR). Yauco: Río Yauco,


Barinas, Caudales s.n. (UPR). ST. CROIX: 

Katherine’s Rest, Ricksecker 19, 99 (US). 

2. Eichhornia diversifolia (Vahl) Urb., Symb. 

Antill. 4: 147. 1903; Heteranthera diversifolia 

Vahl, Enum. Pl. 2: 44. 1805; Piaropus 

diversifolius (Vahl) P. Wilson in Britton & P. 

Wilson, Bot. Porto Rico 5: 149. 1923. Type: 

French. Guiana. L.C. Richard s.n. (holotype: 

C). 

Aquatic annual, rooted in mud. Stems 

elongate, slender, floating. Flowering stems 1.5-3 

cm long, erect. Leaves produced at intervals along 

stem, not in a basal pseudo-rosette, dimorphic, the 

sessile ones with linear blades, 3-7 cm × 2-4 mm, 

acuminate at apex, the petiolate blades rounded to 

cordate, 1.3-3.2 × 0.6-4 cm, with obtuse to acute 

apex and a cordate base; leaf petioles not inflated, 

2-7 cm long. Inflorescence a spike, 2- to 4flowered, 

2-5 cm long (from base of spathe); 

spathes linear, 1.3-1.9 cm long; peduncles 0.9-1.9 

cm long, glabrous. Flowers all opening the same 

day; perianth blue to purplish blue or lavender, the 

tube 8-15 mm long, yellowish within (fide Stahl 

2. HETERANTHERA 

Heteranthera Ruiz & Pav., Fl. Peruv. Prodr. 9. 1794, nom. conserv. 

watercolor), the lobes 4-10 (-12) × 3-4.5 mm, the 

upper one often with a pale yellow spot; stamens of 

two different lengths, the upper 2.5-6 mm long, the 

lower 5.1-7.6 mm long; anthers 0.7-1.3 mm long; 

ovary ellipsoid, 1-4 × 3 mm, glabrous; style 1.5 cm 

long, the stigma capitate, homostylous, pilose. 

Capsule 6-10 mm long; seeds cylindric, 0.4.1 × 

0.2-0.5 mm, longitudinally 9- to 12-winged, 

brown. 

General distribution: Central America, Greater 

Antilles, and South America. 

Distribution in Puerto Rico: In lakes and 

streams. Known from a single collection of 

Woodbury s.n. and a botanical illustration 

prepared by Agustin Stahl in the 1880’s. 

Common name: Puerto Rico: Flor de agua. 


No locality or date given, Woodbury s.n. (UPR). 


Liogier & Martorell (1982) cite Eichhornia 

azurea (Sw.) Kunth as occurring in a pond near 

Cayey, Puerto Rico. However, no specimens have 

been seen to confirm this record nor has it been 

recently collected there. 

Annual or perennial herbs, rooting in mud. Leaves sessile or petiolate, those at base sessile, forming 

a rosette, the blades linear or sometimes oblanceolate, acuminate to obtuse at apex; petiolate leaves 

emersed or floating, the blade reniform, cordate, or oblong, membranous, acute to obtuse at apex. 

Inflorescences spicate or solitary, 1- to 30-flowered; spathes folded or clasping, with acute to caudate 

apex. Flowers with perianth segments connate ½ or more of their length, yellow, blue-mauve, mauve, or 

white, tubular or salverform, limb actinomorphic or zygomorphic, the lobes linear to oblanceolate or 

narrowly elliptic, obtuse to acuminate at apex; stamens 3, unequal, 2 lateral ones usually shorter; filaments 

yellow or purple, inflated in some species, glabrous, glandular-pubescent, or pilose; anthers yellow or 

purple, rounded, oblong, or sagittate; ovary incompletely 3-locular; ovules 10-many; style 3-lobed. Fruit 

an elongate capsule; seeds 10-200, ovoid, testa with longitudinal wings. Twelve species distributed in the 

Americas and Africa. 

TYPE: Heteranthera reniformis Ruiz & Pav. 

Reference: Horn, C. N. 1985. A systematic revision of the genus Heteranthera (sensu lato; 

Pontederiaceae). Ph.D. Dissertation, University of Alabama. 260 pp. 

Key to the species of Heteranthera 

1. Petiolate leaves emersed, erect, the blades ovate to elliptic or elliptic-lanceolate, cuneate to truncate at 

base; inflorescence 1-flowered; perianth blue or white, actinomorphic …..... 1. Heteranthera limosa 

2. Petiolate leaves floating, or if emersed weakly ascending, the blades round-reniform; inflorescence 2to 

8-flowered; perianth white, zygomorphic ……..................................….. 2. Heteranthera reniformis

172 


1. Heteranthera limosa (Sw.) Willd., Ges. 

Naturf. Freunde Berlin, Neue Schriften 3: 

439. 1801; Pontederia limosa Sw., Prodr. 57. 

1788. Type: Jamaica. Swartz s.n. (lectotype: 

S-Sw, R-5197; isolectotype: C). here 

designated*. 

Plant annual, emersed and rooting in shallow 

water. Flowering stems erect, 2–24 cm long. 

Leaves strongly ascending to erect, forming a 

basal rosette, the petiolate ones emersed; leaf 

petioles stout at base, gradually narrowed distally, 

2–14 cm long; blades of petiolate leaves ovate to 

elliptic or elliptic-lanceolate, 1–5.5 cm × 4–33 

mm, acute to broadly so at apex, truncate to 

cuneate at base. Inflorescences 1-flowered, 4-6 cm 

long; spathes 0.9–4.5 cm long, glabrous. Flower 

opening within 1 hour after dawn, wilting by 

midday; perianth blue or white, salverform, limbs 

essentially actinomorphic, lobes equal, narrowly 

elliptic, 5.2–16 × 3–3.5 mm, the distal lobes 

yellow toward base; perianth tube 15–44 mm long, 

essentially glabrous; stamens unequal, lateral 

stamens 2.3–7.8 mm long, central stamen 3.3–7.2 

mm long, the filaments linear, glandularpubescent; 

style glabrous. Capsule linear-oblong 

or narrowly ovoid-ellipsoid, 12-23 × 2.5-5.5 mm; 

seeds 0.5–0.8 × 0.2–0.6 mm, longitudinally 9– to 

14–winged, grayish brown. 

General distribution: United States, Mexico, 

Central America, West Indies, and South America. 


from a single collection made in a roadside ditch; 

Mayagüez. 

Note: *The lectotype specimens designated 

here follow those chosen by Horn (1985) but were 

not validly published (see reference above for 

Heteranthera). 


Mayagüez: About 5 km N of Mayagüez, Caudales 

33256 (UPR). 

2. Heteranthera reniformis Ruiz & Pav., Fl. 

Peruv. 1: 43. 1798. Type: Peru. Ruiz & Pavón 

s.n. (lectotype: MA; isolectotypes: B, BM, 

K), here designated*. 

Plant annual or facultatively perennial, 

trailing and rooting at the nodes in mud or floating 

in shallow water. Flowering stems 1–9 cm long, 

often deflexed. Leaves strongly ascending to erect, 

forming a basal rosette, the petiolate ones floating 

or emersed; leaf petioles stout, 1–15 cm long; 

blades of petiolate leaves round-reniform, 1–4 × 

1–7.5 cm, the apex obtuse, cordate at base. 

Inflorescences spicate, sub loosely 2– to 8flowered, 

to 5 cm long; spathes folded, 0.8–5.5 cm 

long, glabrous; peduncle 0.5–4.2 cm long, 

glabrous. Flowers opening ca. 3 hours after 

sunrise, wilting by early afternoon; perianth white, 

salverform, limbs zygomorphic, lobes narrowly 

elliptic, 3–6.5 mm long, distal central lobe with 

yellow or green region at base, sometimes with 

distal brown spot; perianth tube 6–10 mm long, 

densely glandular-pubescent; stamens unequal, 

lateral stamens 0.9–2.2 mm long, the filaments 

linear, pubescent with white multicellular hairs 

toward apex; central stamen 2.2–4.7 mm long, the 

filament sparsely pubescent; style pubescent with 

multicellular hairs. Capsule narrowly ellipsoidoblong, 

8-12 mm long, 2-3 mm in diam., acute and 

short-beaked; seeds oblong-ellipsoid, 0.5–0.9 × 

0.3–0.5 mm, 8– to 14-winged, light brown. 

General distribution: Eastern United States, 

Mexico, Central America; scattered in South 

America (Argentina, Brazil, Paraguay); naturalized 

in Italy. 


Islands: Wet mud of streams and roadside ditches. 

Widely distributed, although seldom collected. 

Note: *The lectotype specimens designated 

here follow those chosen by Horn (1985) but were 

not validly published (see reference above for 

Heteranthera). 


Carolina: Bo. Cangrejo Arriba, in small park 

opposite jct. Of Road 187 extension w/Highway 

26 (Ave. Baldorioty de Castro), Proctor 45228 

(US). Humacao: Bo. Punta Santiago, beside Hwy 

3 at Río Antón Ruiz bridge, Proctor 44204 (US). 


Pontederia cordata L (Fig. 63. F) is 

sometimes cultivated in ponds and aquatic gardens 

in Puerto Rico.


Family 23. MUSACEAE Banana Family 

Musaceae Juss, Gen. Pl. 61. 1789, nom. conserv. 


Large perennial herbs with a massive sympodial corm and adventitious root systems, monocarpic (in 

Ensete) or suckering (in Musa). Leaves large, spirally arranged, with large, broad, entire blades and stout, 

long petioles with closely-overlapping basal sheath, forming a pseudostem. Inflorescence terminal, and 

usually pendulous, with spirally arranged, keeled, leathery, colorful bracts enclosing a few-flowered, 

axillary monochasial cyme. Flowers zygomorphic, functionally unisexual or less often bisexual (in 

Ensete), pistillate flowers subtended by the lower bracts, staminate ones by the upper bracts; sepals 3, 

petaloid, adnate to 2 petals to form a tubular structure that soon splits along one side, the remaining petal 

free. Stamens 5 or 6, the 6th one usually staminodial, anthers linear. Ovary inferior, 3-locular, with 

numerous axile ovules per locule, style terminal, the stigma 3-lobed, papillate. Fruit a fleshy capsule with 

separable exocarp. Seeds stony, few to numerous, operculate, without an aril. A palaeotropical family of 

two genera and ca. 42 species. The common cultivated banana or plantain are sterile triploid that produce 

no seeds. 

TYPE: Musa L. 

References: Purseglove, J.W. 1972. Tropical Crops: Monocotyledons. Longman. 607 pp. Ríos, P.G. 

1930. Cultivo del banano en Puerto Rico. Ins. Agric. Exp. Sta. Río Piedras Bull. 36: 1-58. Simmonds, 

N.W. 1982. Bananas, 2nd edition. Longman, London. 

Note: The Musaceae is partially treated in this work because of the ubiquitous presence of Musa 

cultivars on the islands. They are propagated by asexual means and have only been recorded as persistent 

(not naturalized) in secondary forests and abandoned farms. 

Musa L., Sp. Pl. 1043. 1753. 

1. MUSA 

Characters as given for the family. Musa are predominantly clumped perennial herbs with unisexual 

flowers. About 35 species are naturally distributed in the Old World tropics. Musa is cultivated as a source 

of fibers, ornamental plants, and most importantly as a food crop. Cultivation of Musa sterile diploids or 

triploids (bananas and plantains) constitutes a major crop throughout tropical and subtropical regions of 

the world. Traditionally, the cultivated bananas are referred to Musa sapientum L. while the plantains to 

M. paradisiaca L. as if they were real species. The use of these binomials is confusing since they have been 

applied indiscriminately to wild species and numerous cultivars alike. Both names are typified by triploid 

hybrids (AAB) and may be applied only to clones that resemble them. The usage of these binomials are 

of little value and therefore should be abandoned in favor of a system that describes the cultivar’s genome. 

Banana and plantain cultivars have been derived from two main species, Musa acuminata Colla (AA) and 

Musa balbisiana Colla (BB), through parthenocarpy, interspecies crossing, and polyploidy. A more 

precise way to refer to these cultivars would be by groups of letters describing their genome (Simmonds, 

1982). The following cultivars are known to occur in Puerto Rico. 

1. AA group. A sterile diploid commonly known as Guineito, Datil, or Niño. 

2. AAA group. Includes the Guineo gigante or Guarán, the Monte cristo, the Monte cristo 

enano, the Guineo enano, the Morado or Colorado, and the Morado verde or Colorado blanco. 

3. AAB group (Fig. 63. H). Includes the Manzano (the type of Musa sapientum), the Plátano 

común, Cuarenteno, Jartón, or Maricongo, the Plátano enano, and the Plátano congo (the type 

of Musa paradisiaca). 

4. ABB group (Fig. 63. G). Includes the Chamaluco, Piche, Mafafo, Malango, Cuatrofilos, or 

Forrongo, the Chamaluco enano, and the Cenizo.

174 


The fertile species Musa velutina H. 

Wendl. & Drude is sometimes grown as an 

ornamental plant. This species usually reaches 1- 

1.5 m tall and has an erect inflorescence with rosy 

fruits that produce seeds. 

Family 24. HELICONIACEAE Heliconia Family 

Heliconiaceae Nakai, J. Jap. Bot. 17. 201. 1941. 


Large perennial rhizomatous herbs with erect pseudostems and distichous, petiolate leaves with 

broad simple blades. Inflorescence terminal, erect or recurved and pendulous with distichous, colored 

bracts enclosing axillary glomerate or cincinnate flowers. Flowers bisexual; sepals 3, two of them adnate 

at base of the corolla, the third sepal free and larger; petals united into a tube, 3-toothed at the apex; fertile 

stamens 5, with linear anthers, the single staminode short and petaloid; ovary inferior, 3-locular, with one 

basal ovule in each locule, style filiform, the stigma clavate or capitate and 3-lobed. Fruit a capsule or 

drupe-like, separating into 3 cocci or dehiscent, usually blue or black. Seeds stony, grayish, without an aril. 

A primarily Neotropical family of one genus and more than 100 species, all of them occurring in Central 

and South America and the West Indies except 6 species and several varieties indigenous to islands of the 

southwest Pacific area from Samoa to Indonesia. The Neotropical species are pollinated by 

hummingbirds, while those of the Pacific islands are pollinated by nectar-feeding bats. 

TYPE: Heliconia L. 

Reference: Berry, F. & W. J. Kress. 1992. Heliconia: an identification guide. Smithsonian Institution 

Press. 334 pp. 

1. HELICONIA 

Heliconia L., Mant. Pl. 2: 147, 211. 1771, nom. conserv. 

Characters as given for the family. Distribution and notes as given above for the family. 

TYPE: Heliconia bihai (L.) L. (≡ Musa bihai L.) 

Key to the species of Heliconia 

1. Inflorescence pendulous …..................................................................................……… 4. H. rostrata 

1. Inflorescence erect. ………………..................................................................................................… 2 

2. Plants mostly ca. 1 m tall, rarely to 1.5 m. ……...............................................................…….. 3 

3. Bracts ascending at an angle of 10 o -45 o to the rachis; flowers orange with black tips 

……….................................................................................................…. 3. H. psittacorum 

3. Bracts nearly horizontal; flowers yellow to green, not black-tipped (sometimes white-tipped) 

………...................................................................................…….. [ H. acuminata] 

2. Plants usually 2 m tall or more. ……..............................................................................……… 4 

4. Bracts yellow, or yellow variously flushed with red, deeply boat-shaped with bases 

overlapping ………..................................................................................…. 2. H. caribaea 

4. Bracts crimson or deep red, sometimes green-margined towards the apex, narrowly boatshaped, 

not overlapping at bases. …...........................................................……………… 5 

5. Bracts more or less spirally arranged; bracteoles soon decaying into a fibrous mass; floral 

tube more than 10 mm long ……......................……………………. 1. H. bihai 

5. Bracts distichous; bracteoles not decaying into a fibrous mass; floral tube less than 5 mm 

long ……………...........................................................................….. 5. H. stricta


1. Heliconia bihai (L.) L., Mant. Pl. 2: 211. 1771; 

Musa bihai L., Sp. Pl. 1043. 1753; Bihai bihai 

(L.) Griggs, Bull. Torrey Bot. Club 31: 445. 

1904, nom. inadmiss. Neotype: Surinam 

(LINN-286.1), designated by L. Andersson, 

Nordic J. Bot. 1: 765, 769. 1981. 

Plants 1.5 m tall. Petioles variable, up to 2 m 

long; leaf blades narrowly oblong, up to 2 m long, 

and 20 cm or more wide, glaucous when young, 

but often becoming green on both surfaces; apex 

cuspidate-acuminate, the base rounded to acute. 

Inflorescence erect with a short peduncle; bracts 

shallowly boat-shaped, 2.5-4 cm wide, narrowly 

triangular, succulent, the middle ones ca. 13 cm 

long or more, widely separated and never 

overlapping at the base, usually scarlet, crimson, 

or dusky red, often with the upper margins yellow; 

flowers with perianth 3 cm long, the sepals and 

petals white-tipped. Fruits bright blue. 


Rico, Lesser Antilles, and northern South 

America. 


Luquillo, Río Grande, Santa Isabel, Trujillo Alto, 

and Utuado; this species is very rare in Puerto 

Rico. 


Luquillo: Bo. Sabana, Rt. 988, Axelrod et al. 

12548 (UPRRP-2). Santa Isabel: Santa Isabel, Bo. 

Felicia 2, Paso Seco, Pomales 12 (UPRRP). 

Utuado: Bo. Caniaco, Acevedo-Rdgz. & Chinea 

11808 (US). 

2. Heliconia caribaea Lam., Encycl. 1: 426. 1785. 

Type: Martinique. Plumier (Burman ed.), Pl. 

Amer., t. 59. 1756. 

Heliconia borinquena Griggs, Bull. Torrey Bot. 

Club 30: 658. 1903; Bihai borinquena 

(Griggs) Griggs, Bull. Torrey Bot. Club 31: 

445. 1904. Type: Puerto Rico. Underwood & 

Griggs 363 (holotype: US!; isotype: NY!). 

Bihai bihai of Britton & P. Wilson, 1924, non 

Musa bihai L., 1753. 

Figs. 37. A-E; 63. I 

Plants 3-6 m tall. Petioles up to 2 m long; leaf 

blades oblong or elliptic-oblong, up to 1 m long or 

more, 20-35 cm wide, green on upper side, usually 

more or less glaucous beneath, abruptly shortacuminate 

at the apex, the base acute or rounded. 

Inflorescence erect with short peduncle, 30-50 cm 

long, of heavy texture; bracts deeply boat-shaped, 

acute to long-acuminate, 6-15 cm long, closely 

overlapping at base, yellow, often variously 

flushed with red with a yellow edge; flowers ca. 3 

cm long, green or greenish yellow with sepals and 

petals green-tipped. Ripe fruits deep blue. 


Antilles. 


Islands: Frequent in humid ravines and on moist or 

wet hillsides at low to middle elevations (50-700 

m). Recorded from Barranquitas, Bayamón, 

Caguas, Cayey, Fajardo, Guayama, Las Piedras, 

Luquillo, Naguabo, Patillas, Río Grande, and 

Yauco; Tortola. 

Common names: Puerto Rico: Botecitos, 

Guineo cimarrón, Guineo silvestre, Pámpano, 

Plátano de indio. 


Barranquitas: Sargent 682 (US). Bayamón: 

Sintenis 1131 (US). Fajardo: Río Arriba, N.L. 

Britton & Shafer 1684 (US). Naguabo: Sierra de 

Naguabo, Shafer 3352 (US-2). Río Grande: Sierra 

de Luquillo, Eggers 1278 (US). TORTOLA: Sage 

Mountain, Fishlock 467 (US). 

3. Heliconia psittacorum L. f., Suppl. Pl. 158. 

1782. Lectotype: Surinam. Dahlberg 13 

(LINN-286.4), designated by L. Andersson, 

Opera Bot. 82: 50, 51. 1985. 

Stoloniferous herb usually ca. 1 m tall, 

growing in dense clumps. Leaves several, 

petiolate, the blades narrowly oblong, divaricate, 

up to 40 cm long, 5-10 cm wide, green, the apex 

acuminate, the base rounded to subcordate, often 

oblique. Inflorescence erect, long-pedunculate, 

the peduncle 15-55 cm long, slender, 2-3 mm 

thick, glabrous; bracts 3-6, usually orange-yellow, 

distichous, moderately spaced, 3-6 cm long, 1 cm 

wide. Flowers ca. 4.5 cm long, yellow-orange, the 

segments tipped with black. Ripe fruits subglobose, 

to 9 mm in diam., variously colored yellow, 

orange, bluish, or black. 

General distribution: Native to the Lesser 

Antilles and tropical South America; commonly 

cultivated as an ornamental in gardens and beside 

houses in most tropical countries, where it tends to 

persist or become naturalized. 

Distribution in Puerto Rico & Virgin Islands:

176 


Fig. 37. A-E. Heliconia caribaea. A. Inflorescence and leaf. B. L.s. bract showing flowers. C. Flower. D. Sepal. E. Capsule. (A, 

from Proctor 44647; B-E, from Proctor 44728).


Recorded as a naturalized plant from Arecibo, 

Cayey, Río Grande, and San Juan. In the Virgin 

Islands recorded under cultivation on Guana 

Island 

Note: This species also has a number of 

horticultural variants, and it hybridizes with other 

species. The hybrid H. psittacorum × 

spathocircinata cv. “Golden Torch” has been 

recorded in Puerto Rico under cultivation in Ciales 

and Río Grande. 


Arecibo: Bosque de Río Abajo, Camp Radley, 

Acevedo-Rdgz. 274 (SJ); Río Abajo State Forest, 

Proctor 43014 (US). San Juan: Río Piedras, Jardín 

Botánico University of Puerto Rico, Acevedo- 

Rdgz. & Serrano 7907; 7909; 7910 (US). ST. 

THOMAS: Crown Mountain, Acevedo-Rdgz. 11215 

(US). 

4. Heliconia rostrata Ruiz & Pav., Fl. Peruv. 3: 

71. 1802; Bihai rostrata (Ruiz & Pav.) 

Griggs, Bull. Torrey Bot. Club 31: 445. 1904. 

Lectotype: Peru; Huánuco, Pavón s. n. (MA), 

designated by L. Andersson in G. Harling and 

L. Andersson, Fl. Ecuador, 221. Musaceae, 

57. 1985. 

Erect herb usually 3-5 m tall; leafy shoots in 

groups of 8 to 29; pseudostem green, glabrous, 

1.25-2 m tall, 3.4-4.5 cm thick; leaves 6 to 19 per 

shoot; petioles 20-30 × 1.1-1.4 cm; blades oblong, 

the longest 100-125 × 15-28 cm, the apex acute to 

acuminate, the base usually truncate, the surfaces 

green and glabrous. Inflorescence pendent, to 70 

cm long; peduncle green to deep red, scurfy to 

puberulous, 6-20 cm long, 1-1.1 cm thick; rachis 

flexuous, red, scurfy to minutely puberulous, 0.7- 

1 cm thick, the bracts spaced 1.2-1.5 cm apart. 

Fertile bracts distichous or spirally arranged, 14- 

33 per inflorescence, 7-8.2 cm long, 3-4 cm wide at 

base, narrowed apically from the distal side to 

form a blunt, beak-like point, the whole bract red 

with a broad yellow margin distally and apically. 

Flowers ca. 19 per fertile bract; perianth 4.5-5.4 

cm long, white to pale yellow at base, yellow 

distally. Stamens with anthers usually connivent 

within apex of corolla-tubes; ovary white to pale 

yellow, glabrous, 6-8 mm long. Ripe fruits blue, 9- 

12 mm long, 5-8 mm in diam. 

General distribution: Venezuela, Colombia, 

Ecuador, Peru and Bolivia; widely cultivated 

elsewhere because of its handsome inflorescences. 


Río Grande, in the Caribbean National Forest, 

Sierra de Luquillo, from a roadside collection, 

perhaps planted or persistent after cultivation. 


Río Grande: Caribbean National Forest, Rt. 191, 

Rodríguez & Vélez 46 (UPRRP). 

5. Heliconia stricta Huber, Bol. Mus. Paraense 

Hist. Nat. 4: 543. 1906. Lectotype: Peru; 

Loreto. Huber 2038 (MG), designated by L. 

Andersson, Nordic J. Bot. 1: 778. 1981. 

Heliconia humilis of authors, non Musa humilis 

Aublet, 1775. 

Plants slender to rather stout, (0.75-) 1-3 (-4) 

m tall. Petioles slender, 25-90 cm long; leaf blades 

ovate to obovate, or oblong, 28-75 × 7-20 cm, 

sharply acuminate at the apex, obtuse to acute or 

shortly attenuate at the base, green on both sides. 

Inflorescence erect, sessile or with a short 

peduncle up to 8.5 cm long, glabrous; bracts rather 

few (3-9), stout, distichous, rather deeply boatshaped, 

dark red with greenish or yellowish green 

margins, overlapping at base and strongly 

ascending, the lowermost much the longest, 10-26 

cm long, long-acuminate at the apex. Flowers 

white at the base and apex, the middle part dark 

green, 5-5.5 cm long, glabrous. Fruits not 

described. 

General distribution: Tropical western South 

America, extending eastward to Manaus in the 

Amazon basin and in the north to Surinam. 

Distribution in Puerto Rico: In swamps and 

wet forests. Recorded as a cultivated plant in the 

Jardín Botánico, Río Piedras and as naturalized in 

Río Grande, Bo. Jiménez in the Sierra de Luquillo, 

in disturbed secondary rain-forest at 210-240 m 

elevation. 


Río Piedras: Jardín Botánico University of Puerto 

Rico, Acevedo-Rdgz. & Serrano 7908 (US). 


The following additional species of Heliconia 

have been noted under cultivation in Puerto Rico: 

Heliconia acuminata Rich. (Bo. Tejas, Las 

Piedras, no voucher) H. irrasa R. R. Sm. (San 

Juan: Río Piedras, Jardín Botánico, Acevedo-Rdgz. 

& Serrano 7906, US-2; 7903, US), Heliconia

178 


latispatha Benth. (Río Grande: Caribbean National 

Forest, in front of forest headquarters, Acevedo- 

Rdgz. 11197, US-2. San Juan: Río Piedras, Jardín 

Botánico, Acevedo-Rdgz. & Serrano 7905, US-2), 

Heliconia marginata (Griggs) Pittier (for 

descriptive information about this species see 

Howard, F1. Lesser Antill. 3: 522. 1979), 

Heliconia platystachys Baker (Río Piedras: Jardín 

Botánico University of Puerto Rico, Acevedo- 

Rdgz. & Serrano 7902, NY, UPRRP, US), and 

Heliconia subulata Ruiz & Pav. 

Family 25. ZINGIBERACEAE Zingiber Family 

Zingiberaceae Adans., Fam. Pl. 2: 61. 1763, nom. conserv. 


Rhizomatous perennials, usually aromatic herbs; rhizomes well-developed, often horizontal and 

creeping, generally rich in starch. Stems usually short, the leaf sheaths forming elongate pseudostems 

above. Leaves distichous, primarily cauline, the basal ones often reduced to bladeless sheaths; sheaths 

elongate, usually open or sometimes closed; ligule present, membranous; petioles present or absent; 

blades lanceolate, oblanceolate, oblong, or linear, acuminate or obtuse at apex, the venation pinnate with 

a prominent midvein and secondary lateral veins which are parallel or arching. Inflorescence a terminal 

spike or thyrse; involucral bracts often scale-like, spirally-arranged. Flowers bisexual, zygomorphic, 

epigynous, the perianth in 2 whorls of 3; calyx tubular or spathe-like, sometimes split on one side, 3-lobed 

or -toothed, green or colorless; corolla fused basally into a tube, 3-lobed, generally longer than the calyx, 

white, yellow, or red; androecium composed of an outer whorl of 2 lateral tooth- to petal-like staminodes 

and a reduced median staminode and an inner whorl composed of a single fertile stamen subtended by 

several lateral, connate, often petaloid staminodes which form a petaloid labellum (lip). Nectar glands 

occur at the base of the floral tube surrounding the style base, or else are sunken in the apex of the ovary. 

Ovary inferior, 3-carpellate, (1-) 3-locular, the locules all fertile; placentation parietal, axile, or ± basal; 

ovules several to many per locule, anatropous; placentation, axile, parietal, or basal; style entire, straight, 

linear-filiform, clasped by the thecae of the anther; stigma funnelform, sometimes slightly 2-lobed, 

papillate, the margin often ciliate. Fruit a dry loculicidally dehiscent capsule or fleshy and berry-like and 

indehiscent, the sepals often persistent at base. Seeds hard, operculate, with a thin lobate or laciniate aril; 

embryo linear. A pantropical family of approximately 50 genera and 1400 species with its center of 

distribution in southeast Asia (with 80-90% of the taxa). The only indigenous genus in Puerto Rico is 

Renealmia, but several other taxa have been introduced, with some of the species more or less naturalized. 

TYPE: Zingiber Mill. 

Reference: Wu Delin & Kai Larsen. 2000. Zingiberaceae. Pp. 322-377. In: Wu Zhengy & P. H. 

Raven, eds., Flora of China, Vol. 24, Flagellariaceae through Marantaceae, Science Press, Beijing, China 

and Missouri Botanical Garden Press, St. Louis, U.S.A. 


1. Inflorescence cone-like, the bracts closely imbricate. …….........................................................…… 2 

2. Inflorescence terminating a long, leafy stem, the leaves with well-developed blades 

…….............................................................................................................................. 3. Hedychium 

2. Inflorescence terminating a long or short stem with sheaths or scale-like cataphylls only, lacking 

well-developed leaf blades …….......................................................................................…….. 3 

3. Inflorescence with a showy involucre of pink to red sterile bracts; corolla pink to red, 

sometimes white; labellum deep crimson with white or yellow margin ………. 2. Etlingera 

3. Inflorescence without a showy involucre of sterile bracts; corolla and labellum pale yellow 

to white …………...........................................................................................… 5. Zingiber


1. Inflorescence an elongated thyrse or raceme, the bracts not closely imbricate. ……….......……….. 4 

4. Ligule tongue-like, 5-12 mm long; labellum (lip) sessile or short-clawed ……….……. 1. Alpinia 

4. Ligule a narrow band of tissue, 1-2 mm long; labellum (lip) long-clawed …………. 4. Renealmia 

1. ALPINIA 

Alpinia Roxb., Asiat. Res. 11: 350. 1810, nom. conserv. 

Coarse aromatic herbs,1-3 m tall, with stout creeping rhizome. Pseudostems well-developed, many, 

leafy. Leaves numerous; blades oblong or lanceolate; ligule generally elongated and tongue-like. 

Inflorescence a terminal panicle, thyrse, or raceme, dense or open, the flowers often disposed in cincinni; 

bracts of main axis remote, minute and scale-like, ovate to lance-oblong or lanceolate; bracteoles tubular 

or open to base, rarely hooded, large or small, sometimes absent; cincinni stalked, 1- to 3-flowered. 

Flowers showy; calyx tubular to turbinate, shallowly 3-lobed, sometimes split on one side; corolla tube 

cylindric, about as long as the calyx, the lobes oblanceolate to elliptical, the central lobe +/- hooded; anther 

with or without a filament, the apex often lacking a terminal appendage; lateral staminodes very small or 

absent, subulate or tooth-like, or sometimes connate with base of lip; lip ovate, tubular-incurved, variously 

lobed or entire; ovary 3-locular with axile placentation, the ovules few to many in each cell; stigma 

subglobose or sometimes clavate. Fruit a subglobose, many-seeded capsule, dry or sometimes fleshy, 

indehiscent or irregularly dehiscent, often red; seeds numerous, globose or angled, with a fleshy or spongy 

aril. Approximately 230 species native to tropical and subtropical Asia, Australia, and the Pacific region, 

introduced in North America, Mexico, West Indies, Central America, and South America. In Puerto Rico 

and the Virgin Islands, only A. zerumbet is known to spread outside cultivation. 

TYPE: Alpinia galanga (L.) Willd. (≡ Maranta galanga L.), typ. conserv. 

1. Alpinia zerumbet (Pers.) B. L. Burtt & R. M. 

Sm., Notes Roy. Bot. Gard. Edinburgh 31: 

204. 1972; Costus zerumbet Pers., Syn. Pl. 1: 

3. 1805, nom. nov. for Zerumbet speciosum J. 

C. Wendl. in Schrad., Sert. Hannov. 4: 3, t 19. 

1798; Languas speciosa (J.C. Wendl.) Small, 

Fl. s.e. U.S., ed. 2. 307, 1375. 1913; Alpinia 

speciosa (J.C. Wendl.) K. Schum. in K. 

Schum. & Hollrung, Fl. Kais. Wilh. Land 29. 

1889, non D. Dietr., 1839. Type: China. “In 

Caldario adferuari debet” J.C. Wendland s.n. 


Fig. 64. A 

Robust herb. Pseudostems 2-3 m tall. Ligule 

tongue-like, coriaceous, 5-10 mm long, obtuse, 

pubescent abaxially; petioles 1-1.5 cm long; 

blades oblong to oblong-lanceolate, 30-70 × 5-14 

cm, glabrous, margins brownish strigose, often 

densely so distally, cuneate at base, acuminate or 

sub-abruptly narrowed at apex to a spirallytwisted, 

caudate tip. Inflorescence a raceme-like 

panicle, drooping, 10-30 × 7-12 cm (in flower); 

rachis purple-red, velvety; panicle branches short 

and stout, often densely pubescent, bearing 1- to 2 

(-3)-flowers; bracteoles elliptic, 1.8-3.5 cm long, 

enveloping the buds, glabrous, white with a pink 

apex, deciduous. Flower pedicels 1-2 cm long; 

calyx subcampanulate, toothed at apex, 1.8-2.5 cm 

long, white; corolla tube shorter than calyx, the 

lobes oblong, 2.5-3 cm long, milky white tipped 

with pink, the central lobe larger than lateral ones; 

lateral staminodes subulate, ca. 2 mm long; 

labellum (lip) ovate or broadly ovate-spatulate, 

3.5-6 cm long, the margins crisped and incurved, 

yellow with purple-red stripes; stamen 2.5-3 cm 

long; ovary hirsute, golden yellow. Capsule 

subglobose, ca. 2 cm in diam., ribbed, vermillion; 

seeds angled. 

General distribution: Native of Asia, cultivated 

for ornament in the tropics and subtropics 

worldwide. 


Islands: Moist roadsides and river banks. 

Commonly planted for ornament and persisting in 

some areas. Recorded from Arecibo, Bayamón, 

Caguas, Canóvanas, Cayey, Mayagüez, and 

Naguabo; St. Thomas. 

Common names: Puerto Rico: Boca de 

dragón, Pimienta Angola. 


180 



10854 (US). Bayamón: Sintenis 1295 (US). 

Caguas: Las Cruces de Caguas, Goll 444 (US). 

Canóvanas: Caribbean National Forest, La 

Condesa Section, Axelrod & Schulz 5619 (US). 

Cayey: S of Cayey, Liogier et al. 28483 (UPR). ST. 

THOMAS: Eggers s.n. (US). 

2. ETLINGERA 

Etlingera Giseke, Prael. Ord. Nat. ad 202, 209. 1792. 


Alpinia purpurata (Vieill.) K. Schum. (syn: 

Guillainia purpurata Vieill.) (Fig. 64. B), a native 

of Polynesia, is grown as an ornamental in Puerto 

Rico and throughout the tropics. Alpinia sanderae 

Sander, a native of New Guinea, was cited by 

Martorell et al. (1981) as cultivated in Puerto Rico. 

Robust, erect herbs; rhizomes creeping, nodose, branched, aromatic. Pseudostems (shoots) welldeveloped, 

leafy, terete. Leaves short-petiolate, lanceolate, large; ligule well-developed. Inflorescence a 

ovoid or subglobose head, or spike, with flowers arranged in 3 or 4 concentric circles on a flat receptacle, 

surrounded at base by numerous, imbricate, often conspicuous, sterile involucral bracts; peduncles arising 

laterally from rhizomes near bases of pseudostems, growing well above ground and elongate or 

subterranean and short; fertile bracts subtending a single bracteole and flower; bracteoles long-tubular, 2to 

3-toothed, persistent; calyx tubular, membranous, unilaterally split, the apex 3-toothed; corolla tube 

equaling or longer than the calyx, 3-lobed at apex, much shorter than the tube. Lateral staminodes absent or 

represented as rudimentary often hairy teeth or bumps. Labellum tongue-shaped, more or less 3-lobed, 

adnate at base to filament and forming a distinct tube, the central lobe with entire or 2-lobed apex, the lateral 

lobes basally folded over stamen; stamen shorter than the labellum, the free part of filament very short and 

rather broad, the anthers bent forward, truncate and emarginate at apex, connective appendage absent; style 

slender, the stigma broad with a dorsal callus; ovary 3-locular, the ovules numerous in each locule. Capsule 

indehiscent, fleshy, obovoid or subglobose, subterete or angular, smooth, longitudinally ridged or with 

obtuse warts in rows. Approximately 70 species distributed in tropical or subtropical regions of China, 

India, Indonesia, Malesia, Philippines, Thailand and northern Australia. 

TYPE: E. littoralis (J. König) Raeusch. (≡ Amomum littorale J. König). 

Reference: Burtt, B. L. & R. M. Smith. 1986. Etlingera: the inclusive name for Achasma, Geanthus and 

Nicolaia (Zingiberaceae). Notes Roy. Bot. Gard. Edinburgh 43: 235-241. 

1. Etlingera elatior (Jack) R. M. Sm., Notes Roy. 

Bot. Gard. Edinburgh. 43: 244. 1986; Alpinia 

elatior Jack, Malayan Misc. 2 (7): 2. 1822; 

Nicolaia elatior (Jack) Horan., Prodr. 

Monogr. Scitam. 32. 1862. Type: Sumatra. 

Jack s.n. (presumably lost). 

Coarse herb often growing in large colonies. 

Pseudostems (shoots) 3-6 m tall. Leaves 

numerous; ligule to 1.5 cm long, shortly 2-lobed, 

glabrous; petiole 1.5-4 cm long; leaf blades 

lanceolate, 38-85 × 8-18 cm, glabrous. 

Inflorescence a terminal, ovoid head of spirally 

imbricated flowers, surrounded at base by showy 

involucral bracts; peduncle 60-150 × 0.8-1.5 cm, 

clothed with green, glabrous sheaths; involucral 

bracts 3-6 × 2-3 cm, spreading, the upper obtuse to 

emarginate, the lower abruptly narrowed to a 

caudate tip, crimson-pink, glabrous; floral bracts 

similar to involucral bracts but pinkish, smaller; 

bracteoles tubular, ca. 2 cm long, unilaterally split; 

calyx 3-4 cm long, unilaterally split, the apex 3toothed; 

corolla pink to red, sometimes white; 

labellum deep crimson with white or yellow 

margin; filament short, flat, whitish pubescent; 

anther red, longer than filament. Fruiting head 

greenish or reddish, globose, 2-2.5 cm in diam., 

short-pubescent; seeds many, black. 

General distribution: Native of Indonesia, 

Malesia, and southern Thailand; widely cultivated 

as an ornamental or for food and often becoming 

naturalized in tropical and subtropical regions 

worldwide. 


Islands: Naturalized populations of this species 

occur in the Luquillo Mountains; common in one


area along a roadside (Acevedo-Rdgz., pers. 

comm., 2005). 

3. HEDYCHIUM 

Hedychium J. König in Retzius, Observ. Bot. 3: 73. 1783. 

Common names: Puerto Rico: Antorcha, 

Torch Ginger. 

Epiphytic or terrestrial, aromatic herbs with tuberous rhizomes. Pseudostems erect, leafy. Leaf blades 

typically oblong or lanceolate; sheaths open; ligule large, membranous. Inflorescence a terminal, spicate 

thyrse, densely-flowered; bracts densely imbricate, 1- to several-flowered; bracteoles tubular. Flowers 

showy; calyx tubular, often split on one side, the apex truncate or shortly 3-lobed; corolla with a long and 

slender tube, the lobes linear, shorter than the tube, reflexed at anthesis; labellum large, suborbicular, 

deeply 2-lobed or –cleft at apex, with a long or sometimes short claw; lateral staminodes 2, petaloid, larger 

than the corolla lobes; stamen usually with an elongate filament, the anther dorsifixed, divaricate at base, 

unappendaged at apex; ovary 3-locular with axile placentation. Fruit a globose, 3-valved, loculicidal 

capsule; seeds numerous; aril lacerate, red. 

TYPE: H. coronarium J. König 

1. Hedychium coronarium J. König in Retzius, 

Observ. Bot. 3: 73. 1783. Type: Asia. J. G. 

König s.n. (holotype: probably at BM; 

isotype: C). 

Fig. 64. D 

Herb, the pseudostems 1-3 m tall. Leaves 

sessile; sheaths whitish or tawny floccose distally; 

ligule 2-3 cm long, bi-lobed at apex, membranous, 

glabrous or with appressed-pubescence; blades 

oblong-lanceolate or lanceolate, 20-40 × 4.5-8 cm, 

long-acuminate to apex, acute at base, adaxially 

glabrous to sparsely puberulous, abaxially 

sparsely to densely whitish floccose. Inflorescence 

10-20 × 4-8 cm, ellipsoid; bracts imbricate, 2- to 3flowered, 

ovate, 4.5-5 × 2.5-4 cm, chartaceous 

with membranous margins, green; bracteoles 30- 

35 mm long. Flowers white, fragrant; calyx split 

on one side, 3-4.5 cm long; corolla tube 7-8 cm 

long, 3 mm in diam., lobes reflexed, the central 

lobe spatulate, the lateral ones linear-lanceolate, 3- 

5 cm × 3 mm; lateral staminodes oblonglanceolate 

or elliptic, 3.5-5.5 × 1.5-3 cm, white 

with yellow at base; labellum obcordate, 2-lobed, 

short-clawed at base, 4-6 cm long, 5-6 cm wide 

when spread out, white, pale yellow at base; 

filament 2.5-3.5 cm long, white, the anther 12-15 

mm long; ovary sericeous. Capsule oblong, 

glabrous, the valves orange-yellow within; seeds 

with a crimson aril. 

General distribution: Native to the Himalayan 

region; cultivated in tropical regions worldwide. 


Islands: Moist roadsides, ditches, and river banks. 

Commonly planted for ornament, naturalized and 

spreading. Recorded from Adjuntas, Bayamón, 

Cayey, Ciales, Maricao, Mayagüez, Naguabo, 

Ponce, Río Grande, and Toa Alta. Not collected in 

the Virgin Islands but very likely cultivated there. 

Common names: Puerto Rico: Jazmín 

cimarrón, Jazmín del río, Mariposa blanca, 

Narciso, Nardo, Dulce nieve. 


Bayamón: Heller & Heller 419 (US). Cayey: 

Sintenis 1881 (US). Ciales: Bo. Fronton, Otero 

354 (UPR). Maricao: Sintenis 494 (US). 

Mayagüez: Sargent 437 (US). Naguabo: Sierra de 

Naguabo, Shafer 3417 (US). Ponce: 20 mi. N of 

Ponce, along Hwy 139, Hansen et al. 9209 (UPR). 

Río Grande: El Yunque, Luquillo Mts., Liogier et 

al. 29299, 29474, 3322 (UPR). Toa Alta: 

Stevenson 3047 (US). 


Hedychium flavum Roxb., Fl. Ind. 1: 81. 1820. 

Several specimens of a Hedychium collected in the 

Luquillo Mountains of Puerto Rico and recorded 

as having yellow or green flowers were reported 

by Liogier & Martorell (2000) as H. flavum Roxb. 

However, the whitish floccose to sparsely whitish 

floccose abaxial leaf surface and ellipsoid 

inflorescence spike are characteristic of H. 

coronarium while H. flavum Roxb. has a glabrous

182 


abaxial leaf surface and an oblong inflorescence 

spike. The specimens in question also have the 

bilobed ligule characteristic of H. coronarium, 

wich separates them from the closely related 

yellow-flowered H. flavescens Carey ex Roscoe. 

Heavy insect damage of the flowering material of 

4. RENEALMIA 

Renealmia L. f., Suppl. 7, 79. 1782, nom. conserv. 

the specimens precluded comparison of any 

further diagnostic characteristics between the 

yellow-flowered species and H. coronarium. The 

specimens probably represent only a color 

variation of H. coronarium. 

Aromatic rhizomatous herbs; rhizomes sympodial, subterranean. Pseudostems well-developed or 

short. Leaves cauline, the lowermost reduced to sheaths; sheaths open, reticulate, striate, or sometimes 

smooth, often with short prickle-like hairs; petioles well-developed or wanting; ligule a narrow band of 

tissue 1-2 mm long; blades narrowly elliptic to narrowly oblong-elliptic or elliptic-oblanceolate, cuneate 

at base, acute to acuminate at apex, essentially glabrous. Inflorescence a terminal thyrse or raceme, rarely 

a spike, borne on a separate shoot, the leaves of which are reduced to bladeless sheaths (cataphylls) or 

sometimes they are blade-bearing; axis and branches of inflorescence often pubescent; bracts herbaceous 

to membranous, deciduous, subtending single flowers or cincinni of 2-13 (-17) flowers; bracteoles 

herbaceous, often tubular and enveloping flowers at anthesis or sometimes open and cup-shaped before 

anthesis; calyx coriaceous, tubular, turbinate, or less often urceolate, 3-lobed, the lobes deltate to 

triangular; corolla tubular proximally, the tube slightly longer than the calyx, 3-lobed, the lobes narrowly 

elliptic, the 2 lateral lobes often smaller than the dorsal one; labellum (lip) clawed at base, often with 2 

lateral, small, tooth-like staminodes. Stamen filament short, ca. 1 mm long or wanting; stigma cup-shaped. 

Capsule 3-locular, globose to ellipsoid, rather fleshy, loculicidally dehiscent, often black or red; seeds 

irregularly ellipsoid, glossy, brown, with a large, yellow, orange, or white aril. 

TYPE: Renealmia exaltata L. f. 

Reference: Maas, P. J. M. 1977. Renealmia (Zingiberaceae-Zingiberoideae), Costoideae 

(Zingiberaceae) Additions. Fl. Neotrop. Monogr. 18: 1-161. 

Key to the species of Renealmia 

1. Leaves of inflorescence shoots blade-bearing; corolla white or light pink; labellum burgundy with white 

center ………......................................................................….. 3. R. jamaicensis var. puberula 

1. Leaves of inflorescence shoots with sheaths only (blades wanting); corolla and labellum yellow, orange, 

or red .……......................................................................................................................................… 2 

2. Inflorescence a raceme, rarely a thyrse, the cincinni 2- to 6-flowered; calyx (12-) 15-20 (-25) mm 

long; corolla 18-32 mm long; capsule (10-) 15-35 (-40) × (6-) 8-20 (-25) mm; seeds more than 

25 …………............................................................................................…… 1. R. alpinia 

2. Inflorescence a thyrse, the cincinni 2- to 17-flowered; calyx 3-10 mm long; corolla 13-16 mm 

long; capsule 4-12 (-14) × 4-9 mm; seeds 10-25.......…….................……………. 2 R. aromatica 

1. Renealmia alpinia (Rottb.) Maas, Acta Bot. 

Néerl. 24: 474. 1976; Amomum alpinia 

Rottb., Soc. Med. Havn. Collect. 2: 245, 248, 

t. 1. 1775. Type: Surinam. Rolander s.n. 

(holotype: probably at C). 

Renealmia exaltata L. f., Suppl. Pl. 79. 1782 

[“1781”]; Alpinia exaltata (L. f.) Roem. & 

Schult., Syst. Veg. 1: 21. 563. 1817. 

Holotype: Surinam. Dahlberg 64 (LINN). 

Renealmia bracteosa Griseb., Fl. Brit. W. I. 601. 

1864. Type: Dominica. Imray s.n., 1859 

(holotype: GOET; isotype: K). 

Renealmia exaltata var. [β ] gracilis K. Schum. in 

Urban, Symb. Antill. 4: 158. 1903. Syntypes:


Puerto Rico; Luquillo Mountains. Eggers 888 

(BR, GH, L, M, P, PR, W) and Eggers 1161 (C 

in part, US!). 

Robust herb, 1-6 m tall; rhizomes 1-3 cm 

thick. Leaves sessile or short-petioled, the petioles 

often pubescent; sheaths smooth and glabrous or 

sparsely setose hairs; ligule short, 1-2 mm long; 

blades narrowly elliptic, 30-110 (-150) × 5-20 cm, 

acuminate at apex, cuneate at base, glabrous or 

sometimes sparsely pubescent abaxially; scape of 

inflorescence erect, 15-50 (-70) × 0.3-1 cm, 

reddish, clothed with sheaths 6-21 × 1-2.5 (-3.5) 

cm. Inflorescence a basal raceme or sometimes 

thyrse, 12-55 × 4-8 cm, with 2- to 6-flowered 

cincinni, more or less hirsute, the rachis, bracts, 

bracteoles, peduncles, and calyx pink to red; bracts 

ovate to narrowly ovate, 1.5-10 (-17) × 0.3-1 (-2) 

cm, membranous; peduncles 5-25 (-35) mm long; 

pedicels 2-15 (-20) mm long; bracteoles 15-30 (- 

35) mm long; calyx tubular, (12-) 15-20 (-25) × 4- 

8 (-10) mm, membranous, appressed-pubescent, 

pink to red; corolla glabrous,18-32 mm long, 

yellow, orange, or red, the tube 12-17 mm long, the 

lobes oblong, 13-15 mm long; labellum 10-12 × 6- 

10 mm when spread out, yellow, limb erect, 5-6 × 

6-10 mm, more or less lobed, the middle lobe 

entire and rounded, lateral lobes involute, 

rounded, 3-5 × 2-3 mm; lateral staminodes ca. 2 

mm long. Anther 8-12 mm long, dark yellow; style 

18-30 mm long; nectarial glands surrounding the 

style base, multilobular; ovary ellipsoid, red. 

Capsule ellipsoid, crowned by the circumcissile 

calyx, (10-) 15-35 (-40) × (6-) 8-20 (-25) mm, red, 

maturing black-purple; seeds numerous, angled, 

2-4 mm in diam.; aril orange. 


America, Puerto Rico, the Lesser Antilles, and 

tropical South America. 

Distribution in Puerto Rico: In wet forests 

and river banks. Recorded from Bayamón, Juncos, 

Manatí, Naguabo, and Río Grande. 

Common names: Puerto Rico: Bihao, Bijao, 

Jengibre de jardín. 


Bayamón: Stahl 685 (US). Fajardo: Río Arriba, 

N.L. Britton & Shafer 1682 (US). Juncos: Sintenis 

2551b (US). Manatí: near Manatí, Sintenis 6626 

(US). Naguabo: Vicinity of La Florida, Shafer 

3132 (UPR, US). Río Grande: El Verde, Luquillo 

Mts., Liogier et al. 28619 (UPR); Sierra de 

Luquillo, Sintenis 1591 (US). 

2. Renealmia aromatica (Aubl.) Griseb., Abh. 

Königl. Ges. Wiss. Göttingen 7: 275. 1857; 

Alpinia aromatica Aubl., Hist. Pl. Guiane 3. 

1775. Lectotype: Plumier, manuscript no. 5, t. 

27 & 28 (P), designated by P. Maas & H. 

Maas, Moscosoa 6: 149. 1990. 

Renealmia occidentalis (Sw.) Sweet, Hort. Brit. 

ed. 2. 493. 1830; Alpinia occidentalis Sw., 

Prodr. 11. 1788. Lectotype: Jamaica. Swartz 

s.n. (M), designated by Maas, Fl. Neotrop. 

Monogr. 18: 105. 1977. 

Robust herb, (0.8-) 1-3 (-4) m tall; rhizomes 

1-1.5 cm thick. Leaves sessile or sometimes shortpetioled, 

the petioles glabrous; sheaths finely 

veined, glabrous; ligule short, 1-2 mm long, 

glabrous; blades narrowly elliptic, 13-55 × (2-) 4- 

12 cm, acute or acuminate at apex, cuneate at base, 

glabrous; scape of inflorescence erect, (15-) 40-65 

(-100) cm long, up to 5 mm thick, reddish, clothed 

with sheaths 4-15 × 0.5-1.5 cm. Inflorescence a 

basal, rarely a terminal, thyrse, 10-35 × 2-7 cm, 

with 2- to 17-flowered cincinni, more or less 

hirsute, the rachis, bracteoles, peduncles, and 

calyx red; bracts narrowly triangular-ovate to 

triangular-ovate, acute or obtuse, 0.6-7.5 × 0.3-1 

cm, membranous, pale green; peduncles 4-10 (- 

12) mm long; pedicels 3-10 (-17) mm long, 

bracteoles 7-17 mm long; calyx turbinate, 3-10 × 

4-7 mm, membranous, appressed-pubescent 

proximally, pink to red, the lobes depressed-ovate, 

obtuse, 1-3 × 3-5 mm; corolla glabrous,13-16 mm 

long, yellow, the tube 12-17 mm long, the lobes 

oblong, 13-15 mm long; labellum 7-9 × 5-9 mm 

when spread out, yellow, limb erect, 4-7 × 5-9 mm, 

more or less lobed, the middle lobe reflexed, 

rounded and entire, lateral lobes rounded, 3-5 × 1- 

2 mm; lateral staminodes ca. 1-1.5 mm long. 

Anther 5-7 mm long, dark yellow; style 14-17 mm 

long; nectarial glands partly surrounding the 

excentric style base; ovary ellipsoid, red. Capsule 

globose to ellipsoid, 4-12 (-14) × 4-9 mm, red, 

rarely purple-black; seeds 10-25, obtusely angled, 

2-3 × 2-4 mm; aril orange. 


America, West Indies, and northwestern South 

America. 

Distribution in Puerto Rico: In forested areas. 

Recorded from Arecibo, Cayey, and Río Grande.

184 


Common names: Puerto Rico: Bihao, Bijao, 

Narciso. 


Arecibo: Bosque de Río Abajo, Camp Radley, 

Acevedo-Rdgz. 536 (SJ). Cayey: Guavate, Liogier 

et al. 29833 (UPR). Río Grande: Coco Beach, 

Liogier et al.29264 (UPR). 

3. Renealmia jamaicensis var. puberula 

(Gagnep.) Maas, Acta Bot. Néerl. 24: 477. 

1976; Renealmia antillarum var. puberula 

Gagnep., Bull. Soc. Bot. France 50: 203. 

1903; Alpinia antillarum var. puberula 

(Gagnep.) Moscoso, Cat. Fl. Doming. 88. 

1943. Lectotype: Dominican Republic. 

Richard s.n. (P), designated by Maas, Fl. 

Neotrop. Monogr. 18: 88. 1977. 

Alpinia antillarum sensu Urban., 1920 and Britton 

& P. Wilson, 1924, non Roem. & Schult., 

1817. 

Fig. 64. E 

Robust herb, 1-3 m tall; rhizomes ca. 1 cm 

thick. Leaves sessile or sometimes short-petioled; 

sheaths finely veined, glabrous or sometimes with 

furcate or stellate prickles near margins and ligule; 

ligule short, 0.5-1 mm long, essentially; blades 

narrowly elliptic, 10-31 (-40) × 2.7-8 cm, the 

uppermost leaves often strongly reduced, acute or 

shortly acuminate at apex, cuneate at base, 

glabrous. Inflorescence a dense, thyrse, 6-17 × 

(1.5-) 3-8 cm, with 2- to 5-flowered cincinni, the 

peduncles and pedicels densely to sparsely 

covered with sessile or short-stalked furcate to 

plurifurcate hairs, often mixed with simple hairs; 

rachis pink to red, pubescent or sometimes 

glabrous; bracts ovate to triangular, acute or 

obtuse, 0.8-3 (-4) × 0.3-1 (-1.5) cm, membranous, 

red, pinkish red, or scarlet; peduncles 2-10 (-12) 

mm long; pedicels 2-15 (-20) mm long; bracteoles 

(7-) 8-11 (-13) mm long, red; calyx tubular to 

5. ZINGIBER 

Zingiber Mill., Gard. Dict. Abr. ed. 4. [as “Zinziber”], nom. et orth. conserv. 

slightly turbinate, 5-11 (-13) × 3-6 mm, 

membranous, essentially glabrous, dark pink to 

magneta, the lobes obtuse, 1-2 × 2-4 mm; corolla 

glabrous, (15-) 20-25 mm long, white or light pink, 

the tube 6-12 mm long, the lobes oblong, 6-13 × 3- 

5 mm; labellum 5-8 × 5-8 mm when spread out, 

burgundy with white center, limb erect, 3.5-6 × 5- 

8 mm, more or less lobed, the middle lobe 

obscurely divided into two broadly ovate-deltate 

to ovate-triangular lobules, 0.5-1 × 1.5-4 mm; 

lateral staminodes 0.8-1.3 × 0.30-0.5 mm. Anther 

6 mm long, red; style 20-23 mm long; nectarial 

glands multilobulate, completely surrounding the 

style base; ovary ellipsoid, sparsely pubescent, 

red. Capsule globose to ellipsoid, 5-10 × 5-8 (-10) 

mm, red, maturing black or puple; seeds 5-25, 

obtusely angled, 1.5-3 × 2-3.5 mm; aril orange. 

General distribution: Hispaniola and Puerto 

Rico. 

Distribution in Puerto Rico: Wet or damp 

montane forest and shaded places. Recorded from: 

Adjuntas, Barranquitas, Cayey, Ciales, Guayama. 

Humacao, Luquillo, Maricao, Naguabo, Ponce, 

Río Grande, and Utuado. 

Common name: Puerto Rico: Narciso 

colorado. 


Adjuntas: Alto de la Bandera, E.G. Britton & 

Marble 2034 (US). Barranquitas: Monte Torrecilla, 

N.L. Britton et al. 5606 (US). Guayama: Mountain 

between Guayama & Cayey, N.L. Britton et al. 

6572 (US). Humacao: Mt. Britton, Luquillo 

National Park, Jones 11023 (US). Maricao: 

Sintenis 216 (US). Naguabo: Sierra de Naguabo, 

Shafer 3225 (US). Ponce: Toro Negro Forest 

Reserve, Acevedo-Rdgz. & Breckon 7832 (US). 

Río Grande: Sierra de Luquillo, Sintenis 1578 

(US); Luquillo Mountains, Caribbean National 

Forest, El Yunque area, Acevedo-Rdgz. 6224 (NY, 

US). Utuado: Bo. Gaonico, Proctor et al. 47107 

(US). 

Rhizomatous aromatic herbs; rhizomes tuberous, branched. Pseudostems erect, elongate, leafy. 

Leaves distichous, with short petioles or sessile; petioles swollen, cushion-like; ligule small, deeply 2lobed 

or entire; blades oblong, lanceolate, or linear. Inflorescence spicate, conical, borne on a separate


shoot from the rhizome, the leaves of which are reduced to bladeless sheaths (cataphylls); bracts closely 

imbricate, 1-flowered, persistent; bracteoles split to the base, not tubular. Flowers solitary in the axils of 

the bracts; calyx tubular, split on one side, 3-toothed at apex; corolla slender, 3-lobed, the central lobe 

often wider than the lateral ones, white or cream; lateral staminodes adnate to the labellum forming a 3lobed 

organ, the central lobe retuse or cleft at apex; stamen with a short filament, the connective elongating 

and enfolding the style; style slender, the stigma not expanded. Capsule loculicidally or irregularly 

dehiscent; seeds black, shiny, covered by an aril; aril white, the margin irregularly lacerate. 

TYPE: Zingiber officinale Roscoe (≡ Amomum zingiber L.). 

Reference: Theilade, I. 1996. Revision of the genus Zingiber in Peninsular Malaysia. Gard. Bull. 

Singapore 48: 207-236. Theilade, I. 1999. A synopsis of the genus Zingiber (Zingiberaceae) in Thailand. 

Nord. J. Bot. 19: 389-410. 

Key to the species of Zingiber 

1. Leaf blades linear to linear-lanceolate or broadly lanceolate, 2-4 (-5) cm wide; ligule ca. 2-10 mm long, 

sparsely pubescent, bi-lobed; bracts red or purplish brown …...............…….. 1. Zingiber montanum 

2. Leaf blades broadly lanceolate or oblong-lanceolate, 3-8.5 cm wide; ligule 15-35 mm long, scarious, 

entire; bracts initially green, maturing red……….......................................…… 2. Zingiber zerumbet 

1. Zingiber montanum (J. König) Link ex Dietr., 

Sp. Pl. 1. 52. 1831; Amomum montanum J. 

König in Retz., Observ. 3: 51. 1783. Type: 

Thailand; Phuket. Koenig s.n. (holotype: C). 

Zingiber purpurem Roscoe, Trans. Linn. Soc. 8: 

348. 1807. Type: Cultivated in the Liverpool 

Botanic Garden, no specimen known. 

Zingiber cassumunar Roxb., Asiat. Res. 347, t. 5. 

1810. Type: Burtt and Smith in Dassanayake 

(Flora of Ceylon 4: 494), indicate the 

lectotype is probably Roxburgh’s drawing 

501 which is at K. 

Rhizomes internally pale carrot color. 

Pseudostems 1.2-1.8 m tall. Leaves sessile or 

short-petiolate; sheaths glabrous or pubescent 

along edges, green; ligule 2-10 mm long, sparsely 

pubescent, bi-lobed; blades linear to linearlanceolate 

or broadly lanceolate, 13-35 (-60) × 2-4 

(-8) cm, hirsute adaxially, acuminate at apex, 

narrowly cuneate at base. Inflorescence fusiform 

or cylindric-ovate, 8-15 × 3-4 cm, acute at apex, 

the shoot (scape) erect, 8-60 cm tall with 5-7 

cataphylls; mature bracts ovate, obtuse to broadly 

so at apex, 3-3.5 × 1-1.7 cm, pubescent, with a 

subscarious, greenish, black-lineolate margin, red 

or purplish brown; bracteoles ovate, 1-1.5 cm 

long, 3-dentate. Calyx 1.2-1.5 cm long, 

membranous, truncate, unilaterally split, white, 

glabrous; corolla 4-6 cm long, the lobes linearlanceolate, 

pale yellow to white, reddish lineolate 

on margins; labellum 6 cm long and 2.5 cm wide, 

white or pale yellow, the central lobe broadly 

rounded, the lateral lobes oblong, free nearly to 

base. Stamen ca. 1 cm long; ovary 3-4 mm long, 

pubescent. Capsule ovoid, ca. 1.5 cm diam. 

General distribution: Probably native to 

India; widely cultivated in southeast Asia for 

medicinal uses. Often escaping cultivation in the 

Greater Antilles. 

Distribution in Puerto Rico: Naturalized in 

moist second-growth forest in limestone hills, 

200-250 m; Isabela. 

Common name: Puerto Rico: Jengibre 

colorado. 


Isabela: Bo. Arenales, trail on W side of Rt. 112, 

Axelrod & Thomas 11949 (UPRRP). 

2. Zingiber zerumbet (L.) Roscoe ex Sm., Exot. 

Bot. 2: 105. 1805; Amomum zerumbet L., Sp. 

Pl. 1. 1753. Type: To be sought among several 

syntypes.* 

Costus glabratus sensu Bello, 1883, non Swartz, 

1788. 

Fig. 64. C 

Rhizomes internally light yellow to yellow. 

Pseudostems 0.6-2 m tall. Leaves sessile or shortpetiolate; 

sheaths glabrescent, green; ligule 1.5- 

3.5 cm long, scarious, entire; blades broadly 

lanceolate or oblong-lanceolate, 15-40 × 3-8.5 cm, 

glabrescent, acuminate at apex, cuneate at base. 

Inflorescence ovoid to ovoid-oblong or ellipsoid,

186 


6-15 × 2.5-5 cm, obtuse at apex, the shoot (scape) 

erect, 10-45 cm tall with 6-8 cataphylls; mature 

bracts broadly obovate or spatulate, broadest 

above middle, broadly rounded or obtuse at apex, 

3-3.5 × 2-3 cm, appressed-pubescent proximally, 

with a subscarious, red-lineolate margin, initially 

green, maturing red; bracteoles linear to lanceolate, 

1.5-3.5 cm long. Calyx 1.2-2.5 cm long, 

membranous, white; corolla 3.5-5.5 cm long, the 

lobes lanceolate, pale yellow to white; labellum 

1.5-2.5 cm long, pale yellow to white, the central 

lobe emarginate, the lateral lobes free nearly to 

base. Stamen ca. 1 cm long, the connective ca. 8 

mm long; ovary ca. 4 mm long, glabrous. Capsule 

ellipsoid or obovoid, 0.8-1.5 cm long, red. 

General distribution: Native of southeast 

Asia. Cultivated in tropical regions worldwide for 

culinary or medicinal uses. 

Distribution in Puerto Rico: Wet roadsides, 

stream and river banks, and shaded forested areas. 

Recorded from: Arecibo, Bayamón, Cataño, 

Cidra, Fajardo, Maricao, Mayagüez, Naguabo, 

Utuado, Vega Alta, Vega Baja, and Yabucoa. 

Note.* A lectotypification made by Theilade 

(Gard. Bull. Singapore 48: 228. 1996) is erroneous 


because the proposed lectotype is not part of the 

original material. 

Common name: Puerto Rico: Jengibre 

amargo. 



11382 (US). Bayamón: Hioram 326 (US). Cataño: 

Goll 1058 (US). Cidra: Pueblo Viejo, Stevenson 

6620 (US). Fajardo: Río Arriba, Liogier et al. 

34479 (UPR). Maricao: Maricao State Forest, 

Liogier 35674 (UPR). Mayagüez: Sintenis 124 

(US). Naguabo: Sierra de Naguabo, Shafer 3143 

(US). Utuado: Bo. Don Alonso, Acevedo-Rdgz. 

13419 (FTG, MAPR, NY US). Vega Alta: 

Between Vega Alta & Corozal, Liogier & Liogier 

37356 (UPR). Vega Baja: Rd. 155, km 5, 

Woodbury s.n. (UPR). Yabucoa: vic. Sta. Helena, 

Sintenis 5117 (US). 


Zingiber officinale Roscoe, native to Asia, 

was cited by Martorell et al. (1981) as cultivated in 

Puerto Rico. 

Species from several genera of Zingiberaceae are cultivated in Puerto Rico and are sometimes 

persistent after cultivation but have not become naturalized. Curcuma longa L., native to Asia, is recorded 

by specimen records from Aguada (Sargent 585, US), Maricao (Near Rosario, Liogier et al. 30943, UPR), 

Mayagüez (Sintenis 491, US), and Moca. Kaempferia rotunda L., native to the East Indies, was cited by 

Liogier & Martorell (1982, 2000) and is based on a specimen record from the Río Abajo Forest Reserve 

(Arecibo: Río Abajo, Liogier et al. 30879, UPR). However, the identity of the specimen is uncertain 

because it is sterile and has uncharacteristic leaves for that species. 

Family 26. COSTACEAE Costus Family 

Costaceae Nakai, J. Jap. Bot. 17: 203. 1941. 


Mostly tall, non-aromatic, perennial, glabrous or pubescent herbs with creeping tuberous rhizomes; 

aerial stems unbranched (except occasionally leafy branch shoots arising from the base of an old 

inflorescence or from leaf-axils), terete, straight or spirally contorted, with closed leaf sheaths and a 

truncate or 2-lobed ligule. Leaves short-petioled, spirally attached to the stem, mostly elliptic (rarely 

lanceolate or obovate), glabrous or variously pubescent. Inflorescence a terminal strobiloid or capitate 

spike, or else flowers solitary in leaf-axils, or rarely terminating a short leafless stem unlike the taller 

sterile leafy ones. Bracts closely imbricate, each bract subtending one flower (or 2 flowers in some African 

species), and each with a linear nectarial callus just below the apex. Bracteole folded or tubular. Calyx


tubular, shortly 3-lobed, the lobes subequal. Corolla tubular at base, the tube equaling the calyx; outer 

portion 3-lobed, the lobes narrowly elliptic, the dorsal one larger than the two lateral ones. Labellum 

equaling or much exceeding the corolla, oblong-obovate to broadly obovate when spread out, often more 

or less 3-lobed, the middle lobe small, the lateral lobes small to very large, their margins crisped, fimbriate, 

or entire. The single stamen petaloid, the anther usually attached at the middle. Style filiform; stigma 

bilamellate, provided with a 2-lobed dorsal appendage, or else cup-shaped and without an appendage. Two 

nectar glands present toward apex of the ovary. Ovary 3-locular or 2-locular; ovules numerous. Fruit a 3or 

2-locular capsule crowned by the persistent calyx, longitudinally dehiscent, or indehiscent and 

rupturing irregularly when ripe. Seeds angular-ovoid to four-sided prismatic, arillate, glossy black or 

brown; endosperm copious. A pantropical family of 4 genera and ca. 100 species, most of them in the 

genus Costus. Only Costus occurs in Puerto Rico. 

TYPE: Costus L. 

Reference: P. J. M. Maas. 1972. Costoideae (Zingiberaceae). F1. Neotrop. Monogr. 8: 1-140. 

Costus L., Sp. Pl. 2. 1753. 

1. COSTUS 

Characters chiefly as given for the family. Large, tall, low, or acaulescent herbs, with flowers in 

strobilaceous spikes and ovary 3-locular. 

TYPE: Costus arabicus L. 

Key to the species of Costus 

1. All bracts provided with foliaceous appendages ………...…... 1. C. guanaiensis var. macrostrobilus 

1. Bracts without appendages (or appendages only on lowest bracts). ………...........................……… 2 

2. Bracts sharp-pointed (pungent) at the apex, red; underside of leaves densely sericeous; calyx 22- 

25 mm long. ……………............................................................................... 4. C. speciosus 

2. Bracts never pungent; calyx 3-16 mm long. …….........................................................……….. 3 

3. Plants less than l m tall; ligule hardly 1 mm long; leaves broadly obovate or elliptic, marked 

with faint bands of light and dark green, densely villous on upper side, glaucous and densely 

puberulous beneath ………....................................……… 2. C. malortieanus 

3. Plants up to 2.5 m tall or more; ligule 2-15 mm long; leaves narrowly elliptic, not marked, 

variously pubescent but never villous on upper side. ………….....................…………… 4 

4. Exposed part of bracts orange-red; calyx 3-7 mm long …………..………. 3. C. scaber 

4. Exposed part of bracts greenish; calyx 9-16 mm long ………….…………. 5. C. spicatus 

1. Costus guanaiensis var. macrostrobilus (K. 

Schum.) Maas, F1. Neotrop. Monogr. 8: 52. 

1972; Costus macrostrobilus K. Schum. in 

Urban, Symb. Antill. 4: 159. 1903. Type: 

Puerto Rico; Sierra de Luquillo. Eggers 1289 

(lectotype: US!; isolectotype: C), designated 

by Maas, Fl. Neotrop. Monogr. 8: 54. 1972. 

Plants stout, mostly 2-6 m tall; sheaths 10-40 

mm in diam., glabrous to puberulous or villose. 

Ligule truncate, 5-10 (-15) mm long; petioles 5-20 

mm long; leaf blades narrowly ovate to narrowly 

obovate, shortly acuminate at the apex, cuneate, 

rounded, or rarely cordate at the base, mostly 20- 

65 × 5-15 cm, the upper surface strigose and 

somewhat scabrid to the touch. Inflorescence 

ovoid, obtuse, 5-15 (-30) cm: long, 4-6 (-10) cm 

thick, terminating a leafy stem; bracts green on the 

exposed part, usually red on the covered part, 

coriaceous, broadly ovate, 2-4 (-6) cm long, 1.5-5 

cm wide; appendages foliaceous, rarely absent, 

rigid, green, ascending or slightly reflexed, 

narrowly to broadly deltate, 2-4 cm long or more, 

1.5-2 cm wide or more; bracteoles red, often 

green-tipped, 2.5-4 cm long, glabrous to densely 

puberulous. Calyx red, 14-18 (-22) mm long. 

Corolla usually white, 7-10 cm long, glabrous to 

puberulous, the tube ca. 2 cm long, the lobes

188 


narrowly obovate; labellum white or yellowish 

white, broadly obovate when spread out, 7.5-11 × 

6-7 cm, the lateral lobes usually striped with red, 

the middle lobe reflexed, blotched with yellow to 

orange in the center. Stamen narrowly elliptic, 4-6 

cm long, up to 2 cm wide; anther 1-1.4 cm long. 

Ovary 0.5-1 cm long, densely puberulous to 

glabrous. Capsule 1.2-2.2 cm long, puberulous to 

glabrous; seeds black. 

General distribution: This variety occurs in 

Central America, Puerto Rico, Trinidad, and 

northern South America (to Peru). Three other 

varieties exist. 

Distribution in Puerto Rico: Grows in moist 

or wet thickets at low to upper middle elevations 

(90-860 m), locally frequent. Recorded from 

Arecibo, Ceiba, Florida, Río Grande, and Utuado. 

Common name: Puerto Rico: Caña india. 


Arecibo: Río Abajo State Forest, along Igartua 

trail, Acevedo-Rdgz. 10597 (US). Río Grande: 

Sierra de Luquillo, Sintenis 1740 (US); Proctor 

44752 (US). 

2. Costus malortieanus H. Wendl., Hamburger. 

Garten-Blumenzeitung 19: 30. 1863. Type: 

Costa Rica. Wendland s. n. (probably at 

GOET). 

Plants less than 1 m tall; sheaths reddish near 

insertion of the petiole, to 2 cm in diam., glabrous 

to variously hairy; ligule truncate, hardly 1 mm 

long; petiole to 2 mm long, densely villose; leaf 

blades obovate to elliptic, mostly 15-26 × 9-15 cm, 

blunt and very shortly apiculate at the apex, 

cuneate at base, the upper side lightly to densely 

villose, light green marked with faint darker green 

bands, the underside paler green or glaucous and 

puberulous. Inflorescence globose to shortcylindrical, 

4-9 cm long, ca. 4 cm in diam.; 

exposed part of bracts green, the covered part 

reddish; bracts coriaceous, broadly ovate, obtuse, 

3-4 cm long and wide, glabrous; bracteoles 

reddish, 1.4-2 cm long, glabrous. Calyx reddish to 

greenish, 5-9 mm long. Corolla yellow to cream, 5- 

7 cm long, glabrous, the tube 1-2 cm long, the 

lobes narrowly obovate; labellum yellow, broadly 

obovate when spread out, 4-6 cm long, 3.5-5 cm 

wide with slightly crenulate margins, the lateral 

lobes usually striped with dark red, the middle lobe 

reflexed with 3 spatulate lobules. Stamens white, 

slightly tinged with purple, narrowly elliptic, 4.2- 

4.5 cm long, with recurved, slightly 3-lobulate 

apex; anthers 6-7 mm long. Ovary 5-8 mm long, 

glabrous. Capsule ellipsoid, trisulcate, 12 mm 

long, glabrous; seeds black. 

General distribution: Native to Nicaragua 

and Costa Rica in moist to wet lowland forests; 

cultivated worldwide for its attractive foliage. 

Distribution in Puerto Rico: Found naturalized 

at one site in Río Grande. 


Río Grande: El Verde area, disturbed secondary 

rain-forest, ca 240 m elevation, Proctor 50425 

(SJ). 

3. Costus scaber Ruiz & Pav., Fl. Peruv. 1: 2, t. 3. 

1798. Type: Peru. Ruiz & Pavón s. n. (BM, 

fragment). 

Fig. 38. A-H 

Slender plants up to 3 m tall; sheaths glabrous 

to puberulous, mostly 0.5-1.5 cm in diam.; ligules 

obliquely truncate, 2-12 mm long; petioles 2-10 

mm long, puberulous to glabrous; leaf blades 

narrowly elliptic, 10-32 × 3-11 cm, acuminate at 

the apex, cuneate to rounded (rarely cordate) at 

base, the upper surface glabrous to sparsely 

puberulous but with the costa densely strigulose, 

the underside glabrous to minutely puberulous 

with the costa often densely sericeous. 

Inflorescence ovoid to narrowly cylindrical, 4-10 

cm long elongating to 22 cm in fruit, 1.5-3.5 cm 

thick and to 4.5 cm in fruit; bracts orange-red to 

red, broadly ovate, 2-3.5 cm long, obtuse, 

coriaceous, glabrous to puberulous or rarely 

strigose, the margins of the covered part usually 

lacerating into fibers; bracteoles reddish, 0.9-1.2 

cm long or more, glabrous to puberulous. Calyx 

reddish, 3-7 mm long. Corolla orange to yellow, 

3.5-4 cm long, glabrous, the tube ca. 1 cm long, the 

lobes narrowly obovate, 3 cm long. Labellum 

yellow, oblong-obovate when spread out, 2-3 × 

1.5-2 cm, the lateral lobes rolled inward and 

forming a tube. Stamen narrowly elliptic, equaling 

or slightly exceeding the labellum, red to orangered, 

yellowish at the apex, 2-2.5 cm long; anther 5- 

7 mm long. Ovary 3-8 mm long, glabrous to 

densely puberulous. Capsule ellipsoid to 

subglobose, 7-12 mm long, glabrous to densely 

puberulous at the apex; seeds black. 

General distribution: Mexico, Central


Fig. 38. A-H. Costus scaber. A. Habit. B. Upper part of stem with inflorescence. C. Flower and bracteole. D. L. s. of flower. E. 

Stigma. F. Young fruit with bract and bracteole. G. Seeds. H. Seed with aril. From Mori, S. et al. 1997. Vascular plants of central 


190 


America, Hispaniola, Puerto Rico, the southern 

Lesser Antilles, and tropical South America to 

Peru, Bolivia, and Brazil. 


Islands: Recorded as a wild plant only from 

Arecibo and Corozal; St. Thomas. 


Arecibo: 0.5 km S of Biáfara, Acevedo-Rdgz. 

11703 (US). Corozal (in a “forested gully”), 

Britton et al. 8357 (NY). Mayagüez: at the 

Agricultural Experiment Station, Britton & Hess 

2825 (NY, US). ST. THOMAS: Road 33, by entrance 

to Mountain Top, Acevedo-Rdgz. 11367 (US). 

4. Costus speciosus (J. König) Sm., Trans. Linn. 

Soc. London 1: 249. 1791; Banksia speciosa 

J. König in Retz., Observ. Bot. 3: 75. 1783. 

Type: Asia. Koenig s. n. (C-Schum.). 

Costus spicatus var. [β ] pubescens Griseb., F1. 

Brit. W. I. 602. 1864. Type: St. Vincent. 

Guilding s. n. (K). 

Fig. 64. F. 

Plants up to 3 m tall; sheaths green, tinged 

reddish brown, 0.7-1.2 cm in diam., minutely 

puberulous to glabrous; ligules truncate, 1-2 mm 

long; petioles 2-5 mm long, densely puberulosericeous; 

leaf blades narrowly elliptic, 12-25 cm 

long, acuminate at the apex, tipped by a subulate 

point, cuneate to rounded at the base, the upper 

surface glabrous, the underside densely sericeous 

or puberulous and very soft to the touch. 

Inflorescence ovoid to broadly ovoid, 4-7 × 3-5 

cm; bracts red to brownish red, narrowly ovatetriangular, 

acute, 2.5-3.5 × 1-2.5 cm. Calyx 

reddish brown to green, 2.2-2.5 cm long. Corolla 

yellow to pinkish white, ca. 6 cm long, minutely 

sericeous; tube ca. 1.5 cm long, the lobes elliptic to 

obovate, 4-5 cm long. Labellum white to pinkish 

white, 6-7 cm long, broadly obovate when spread 

out and up to 10 cm wide, the margins crenate; 

middle lobe blotched with yellow in the center. 

Stamen white to yellowish white, narrowly elliptic 

to narrowly ovate, 4.5-5 cm long with obtuse apex; 

anthers 1. 1 cm long. Ovary 5-10 mm long, 

sericeous; capsule not described. 

General distribution: Native to the Indo- 

Malayan region, occurring from India to New 

Guinea; widely cultivated in the Neotropics and 

naturalized in the Greater and Lesser Antilles. 

Distribution in Puerto Rico: Grows in 

disturbed moist to wet thickets and roadside banks 

at low to lower middle elevations (50-200 m). 

Recorded from Caguas, Ceiba, Naguabo, and Río 

Grande. 

Note: The following collection was cited by 

Maas (1972) as Costus speciosus for Puerto Rico: 

Wagner 415 (U). 

5. Costus spicatus (Jacq.) Sw., Prodr. 11. 1788; 

Alpinia spicata Jacq., Enum. Syst. Pl. 11. 

1760. Lectotype: Martinique. Jacquin, Select. 

Stirp. Amer. Hist. t. 1. 1763, designated by 

Maas, Fl. Neotrop. Monogr. 8: 104. 1972. 

Costus cylindricus Jacq., Fragm. Bot. 54, t. 77. 

1809. Type: Martinique. Jacquin, Fragm. Bot. 

54, t. 77. 1809 ¹. 

Plants up to 2.5 m tall; sheaths 1-2 cm in 

diam., glabrescent; ligules truncate, 2-10 mm 

long; petioles 2-10 mm long, puberulous to 

glabrous; leaf blades narrowly elliptic, 7-33 × 3.5- 

8.5 cm or more, shortly acuminate at the apex, 

rounded to cordate at base, glabrescent on both 

surfaces. Inflorescence ovoid to cylindrical, 5-27 

× 3-4.5 cm; bracts greenish or reddish on the 

exposed part, reddish on the covered part, broadly 

ovate, 2-4 cm long and broad, obtuse at the apex, 

glabrous and coriaceous, the margin of the covered 

parts lacerating into fibers; bracteoles 1.7-3 cm 

long. Calyx 9-16 mm long. Corolla yellow to pink, 

4-5 cm long, glabrous, the tube 1 cm long, the 

lobes narrowly obovate, 3-5 cm long. Labellum 

yellow, broadly oblong-obovate when spread out, 

2.5-5 cm long and wide, the lateral lobes rolled 

inward and forming a slender tube, the margins 

crenulate. Stamen narrowly elliptic, 3-4 cm long; 

anther 7-8 mm long. Ovary 4-9 mm long, sericeous 

or rarely glabrous. Capsule ellipsoid, 10-15 mm 

long; seeds black. 


Rico, and the Lesser Antilles. 

Distribution in Puerto Rico: Recorded as a 

wild plant from Humacao; in addition, Maas 

(1972) cites a specimen (without locality) 

collected by Boye (S). This species is cultivated at 

the Botanical Garden, Río Piedras. Its occurrence 

as a wild plant in Puerto Rico needs further 

confirmation. 

Common name: Puerto Rico: Caña amarga. 

Note: ¹ Howard (Fl. Lesser Antill. 3: 535. 

1979) regarded Jacquin’s fig. 77 as the holotype of


Costus cylindricus. However, Jacquin’s original 

publication of this name contained two elements 

[Plumier’s manuscript no. 5 t. 30 (cited twice 

through a reference to Aublet) and Jacquin’s own 

illustration plate 77] and therefore Jacquin’s fig. 

77 should be considered a syntype instead. Further 

study of the original material is needed in order to 

make a sound decision of a type for this name. 


Humacao, Pterocarpus forest, Woodbury s.n. 

(UPR). 

Family 27. CANNACEAE Indian-shot Family 

Cannaceae Juss., Gen. Pl. 62. 1789, nom. conserv. 


Erect glabrous perennial herbs; usually with tuberous, starchy rhizomes. Leaves spirally arranged, 

with large simple blades having a prominent mid rib and numerous fine parallel vertical veins, the petioles 

forming open sheaths around the stems; ligules lacking. Inflorescence a terminal racemiform or paniculate 

thyrse often with several to many short, 2-flowered cymules axillary to the bracts; branches each with one 

large primary bract (10-30 cm long), and two branch bracts (5-20 cm long). Flowers bright-colored, 

bisexual, zygomorphic; individual flowers sessile or short-pedicellate; sepals 3, imbricate, equal, free, 

usually green or purplish; petals 3, unequal, connate at base into a short tube. Stamen one, petaloid, with 

a single marginal pollen-sac (theca), this more or less adnate at base to the fleshy, petaloid style and the 

petaloid inner staminode. Inner staminode (labellum) one, often recurved, smaller than the outer ones; 

outer staminodes 0-3, longer than the petals, connate at base with the labellum, style, and stamen into a 

tube. Septate nectar glands present. Ovary inferior, 3-locular, verrucose or tuberculate; ovules numerous. 

Fruit a verrucose to tuberculate, loculicidally dehiscent capsule crowned by the persistent sepals. Seeds 

numerous, globose to ellipsoid, very hard, black to dark brown; without an aril. A Neotropical family of 

a single genus and ca. 9 species, some of which have become naturalized in Africa and tropical Asia. 

Hybridization has produced many variants often planted for their ornamental flowers. 

TYPE: Canna L. 

References: Maas, P. J. M. & H. Maas. 1988. Cannaceae in Flora of Ecuador no. 32, 9 pp. Tanaka, N. 

2001. Taxonomic revision of the family Cannaceae in the New World and Asia. Makino New Ser. 1: 1- 

74. 

Canna L., Sp. Pl. 1. 1753. 

1. CANNA 

With characters as given above for the family. 

LECTOTYPE: Canna indica L., designated by Britton, Fl. Bermuda 86. 1918. 

Key to the species of Canna 

1. Plant conspicuously glaucous; leaf blades less than 15 cm wide; flowers pure yellow; rhizomes slender, 

long-creeping …………..................................................................................… 1. C. glauca 

1. Plant not glaucous; leaf blades usually wider than 15 cm; flowers red to orange, but never pure yellow; 

rhizomes short, tuberous. …….............................................................................................……….. 2 

2. Bracts persistent; flowers usually crimson (rarely yellowish orange); seeds globose, 4-5 mm in 

diam. ………...................................................................................................…….. 2. C. indica 

2. Bracts caducous; flowers pure orange; seeds narrowly ellipsoid, 4-7 × 2.5-4 

mm...................................................................................................................... 3. C. jaegeriana

192 


1. Canna glauca L., Sp. Pl. 1. 1753. Type: Type: 

To be sought among several syntypes.* 

Plants gregarious in wet, marshy sites; 

rhizomes long-creeping, fleshy, to 1.5 cm thick, 

beset with numerous roots; stems up to 1.5 m tall or 

more. Sheath and leaf blades glaucous; blades 

narrowly ovate to narrowly elliptic, 30-50 × 3-15 

cm, very gradually narrowed toward the apex, the 

base narrowly cuneate. Inflorescences usually 

simple (rarely branched), bearing 2-flowered 

cincinni of short-pedicellate light yellow flowers, 

the pedicels up to 1 cm long (in fruit). Floral bracts 

ovate to broadly ovate, 0.7-2.5 cm long; bracteoles 

broadly ovate to ovate-triangular, 0.3-2 cm long. 

Sepals narrowly elliptic-triangular, 1-2.5 cm long. 

Corolla 7-9 cm long, the tube ca. 2 cm long, the 

lobes narrowly ovate, 5-7 cm long. Outer 

staminodes 3, pale yellow, narrowly obovate to 

narrowly elliptic, 8-10 cm long (including basal 

tubular part); inner staminode (labellum) pale 

yellow, strongly recurved, narrowly elliptic to 

narrowly obovate, 7.5-8 cm long. Stamen pale 

yellow, the free part 3.5-4.5 cm long. Style yellow, 

the free part 4-5 cm long. Capsule ellipsoid to 

globose, 2-6 cm long. 

General distribution: Widespread throughout 

the Neotropics. 

Distribution in Puerto Rico: Widespread but 

not common, occurring in wet fresh-water habitats 

near sea level. Recorded from Cabo Rojo, 

Carolina, Guánica, Mayagüez, San Juan, Vega 

Alta and Vega Baja. 

Common names: Puerto Rico: Maraca 

amarilla, Maraca boba, Maraca de pantano. 

Note. * A lectotypification made by N. 

Tanaka (Makinoa New Ser. 1: 53. 2001) is 

erroneous because the proposed lectotype is not 

part of the original material. 


Cabo Rojo: Sintenis 665 (US). Guánica: Guánica 

Lake, Sargent 264 (US). Mayagüez: Road from 

Mayagüez to Joyuda, Underwood & Griggs 130 


Vega Alta: Bo. Sabana, Proctor 44331 (US). Vega 

Baja: Bo. Cibuco, Proctor et al. 45562 (US). 

2. Canna indica L., Sp. Pl. 1. 1753. Lectotype: 

Herb. van Royen 912. 356-390 (L), designated 

by Maas in C.E. Jarvis et al., Regnum Veg. 

127: 29. 1993. 

Canna lambertii Lindl., Bot. Reg. 6: t. 470. 1820. 

Type: Trinidad. Lindley, Bot. Reg. 6: t. 470. 

1820. Based on a plant grown by Lambert 

from seed collected in Trinidad. 

Canna sylvestris Roscoe, Monandr. Pl. Scitam. t. 

10. 1828. Lectotype: Tropical America. 

Herbarium Hookerianum, M1818/72 42 (K), 

designated by N. Tanaka, Makinoa New Ser. 

1: 35. 2001. 

Canna portoricensis Bouché, Linnaea 12: 147. 

1838. Type: Puerto Rico; Moritz s.n. (B, 

destroyed; L?). 

Canna coccinea sensu Britton & P. Wilson, 

1923, non Miller, 1768. 

Figs. 39. A-F; 64. G 

Rhizomes stout, more or less tuberous; stems 

1-2 m tall, with glabrous leaf sheaths; leaf blades 

mostly narrowly ovate to narrowly elliptic, 20-60 

× 10-30 cm, the apex acute to shortly acuminate, 

the base rounded to narrowly cuneate. Inflorescence 

simple or branched, bearing several solitary 

flowers or 2-flowered cincinni. Floral bracts ovate 

to obovate, 1-3 cm long; bracteoles (narrowly) 

ovate-triangular, 0.5-1.5 cm long; sepals ovate to 

narrowly ovate, 0.7-2 cm long. Corolla red (rarely 

yellow), 4-6.5 cm long, the tube 1-2 cm long, the 

lobes linear, 3-5 cm long. Outer staminodes 2 or 3, 

red or yellow, narrowly elliptic to narrowly 

obovate, 5-7.5 cm long, with obtuse or emarginate 

apex. Inner staminode (labellum) mostly red and 

dotted yellowish or pinkish at base (rarely 

completely yellow), recurved, narrowly oblongovate, 

4.5-6.5 cm long, up to 1 cm wide, the apex 

usually emarginate. Stamen 4-6 cm long. Style 

4.5-6.5 cm long. Capsules ellipsoid to subglobose, 

1.5-4 cm long; seeds black, 4-5 mm in diam. 

General distribution: Very common 

throughout the more moist parts of tropical and 

subtropical America; widely naturalized in 

tropical Asia and Africa. Often cultivated and 

escaping from cultivation. Readily hybridizes 

with other species to form a wide spectrum of 

horticultural variants with showy flowers. 


Islands: Common in moist shaded secondary 

vegetation from near sea level to over 700 m 

elevation. Recorded from Adjuntas, Aibonito, 

Añasco, Arecibo, Barranquitas, Barceloneta, 

Carolina, Cataño, Cayey, Ciales, Coamo, Florida, 

Guaynabo, Humacao, Jayuya, Loíza, Lares,


Fig. 39. A-F. Canna indica. A. Part of stem and inflorescence. B. Flower. C. L. s. of flower (left) and transverse section of ovary 

(right). D. Stamen. E. Fruit and c.s. of fruit. F. Seed. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. 

Gard. 78.

194 


Maricao, Mayagüez, Naguabo, Patillas, Ponce, 

Rincón, San Germán, San Juan, San Lorenzo, San 

Sebastián, Toa Baja, and Utuado; St. Croix and St. 

John. 

Common names: Puerto Rico: Bandera 

española, Maraca, Maraca morada, Maraca roja. 

Note: The synonyms cited above for Canna 

indica are names thought by Britton & P. Wilson 

(1924) to represent valid species. However, this 

rather short list includes only a small fraction of 

the total synonymy of this species, which was 

characterized by Howard (1979, p. 541) as “a 

nightmare to every taxonomist”. 


Adjuntas: ca. Saltillo, Sintenis 4015 (US). 

Aibonito: Between Aibonito and Cayey, Heller & 

Heller 518 (US). Arecibo: Río Abajo State Forest, 

Acevedo-Rdgz. 10705 (US-2). Cataño: Goll 997 

(US). Guaynabo: Bo. Sonadora, Proctor & 

Thomas 44555 (US-2). Lares: Bo. Callejones, 

Proctor 44463 (US). Maricao: Bo. Montoso, 

Kennedy et al. 4783 (US). Ponce: Prey 83 (US). 

San Juan: Río Piedras, Stevenson 1224 (US). San 

Sebastián: Sargent 277 (US). Utuado: Road from 

Utuado to Lares, Underwood & Griggs 84 (US). 

ST. JOHN: Reef Bay Quarter, Bordeaux Mountain, 

Acevedo-Rdgz. & Siaca 3869 (US). 

3. Canna jaegeriana Urb., Repert. Spec. Nov. 

Regni Veg. 15: 102. 1917. Type: Haiti. Jaeger 

165 (holotype: B, destroyed; lectotype: K; 

isolectotypes: LE, MO), designated by N. 

Tanaka, Makinoa, ser. 1: 46. 2001. 

Canna pertusa Urb., Repert. Spec. Nov. Regni 

Veg. 15: 101. 1917. Type: Puerto Rico; 

Utuado. Sintenis 6494 (holotype: L). 

Rhizomes stout, tuberous; stems 1-5 m tall, 

with the sheaths glabrous or lanuginose. Leaf 

blades narrowly elliptic to narrowly ovate, 50-100 

× 13-38 cm, shortly acuminate at apex, cuneate at 

base, glabrous on upper (adaxial) side, glabrous or 

slightly lanuginose beneath. Inflorescence simple 

or branched, with cincinni 1-or 2-flowered, the 

pedicels up to 0.5 cm long in flower, to 1 cm long 

in fruit. Bracts caducous, the primary bracts 10-30 

cm long; branch bracts 2.5-13 cm long; flower 

bracts broadly ovate to deltate, 0.1-0.18 cm long; 

bracteoles narrowly triangular to deltate, 0.2-2 cm 

long. Flowers orange, 5-6.5 cm long, the tube 0.5- 

2 cm long, the lobes 2.5-4.5 cm long Outer 

staminodes 3 or 4, obovate-elliptic, with apex 

entire or emarginate, 5-6.5 cm long. Inner 

staminode (labellum) slightly reflexed. Capsules 

ellipsoid, 4.5-10 cm long, densely covered by 

pinkish tipped green papillae; seeds shiny black, 

narrowly ellipsoid, 4-7 × 2.5-4 mm. 


Rico, and western South America on forested 

mountain slopes and stream sides at mostly 300- 

2200 m elevation. 

Distribution in Puerto Rico: Very rare; 

recorded from Utuado and Villalba. 

Common name: Puerto Rico: Maraca 

montuna. 


Utuado: Mt. Morales, N.L. Britton & Cowell 827 

(US). Villalba: Vicinity of Ala de la Piedra above 

Villalba, E.G. Britton & Brown 6228 (US). 


Canna edulis Ker Gawl. and Canna lutea 

Mill. were listed by Britton & P. Wilson (1923) as 

cultivated in the Virgin Islands. 

Family 28. MARANTACEAE Arrowroot Family 

Marantaceae Petersen in Engler & Prantl, Nat. Pflanzenfam. 2, Abt. 6: 33. 1888, nom. conserv. 


Perennial herbs with creeping rhizomes (and sometimes also with starchy tubers), acaulescent or with 

more or less elongate stems. Leaves distichous, vaginate at base, differentiated into sheath, petiole proper 

(often missing), pulvinus, and blade; blades with a prominent mid rib and numerous, closely set lateral 

veins that fuse near the blade margins and are interconnected by minute cross veinlets. Inflorescence 

varying from a simple spike to a complex spiciform thyrse, or with aggregates of 1- or 2-flowered cymules


in the axils of spathe-like bracts. Flowers bisexual, epigynous, each arising from between a spathe-like 

unit of 2 bracts; sepals 3, mostly free and equal; petals 3, united below into a tube; usually contorted, 

unequal and often cucullate. Fertile stamen 1, with 1-locular anther; staminodes 2, free or connate with the 

fertile stamen, petaloid. Ovary inferior, 1- to 3-locular, with a single ovule in each locule; style recurved, 

simple. Fruit capsular, nutlike, or fleshy; seeds solitary, hard, arillate. A chiefly Neotropical family of 30 

genera and more than 400 species. The family is easily distinguished from the related Zingiberaceae by the 

swollen pulvinus at the junction of the petiole with the leaf blade. 

TYPE: Maranta L. 

References: Howard, R. A., Fl. Lesser Antill. 3: 542-550. 1979. Kennedy, H., L. Andersson and M. 

Hagberg. 1988. 224. Marantaceae. In: G. Harling & L. Andersson (eds.), Fl. Ecuador 32: 14-136. 


1. Flowers borne in bracteate spikes; ovary and fruit 3-locular ………...................……….. 1. Calathea 

1. Flowers borne in an open, somewhat diffuse inflorescence; ovary and fruit 1-locular. ……………… 2 

2. Flowers white, the terminal one long-pedicellate, erect; exterior staminodes 2; leaves not over 28 

cm long and not more than 10 cm wide ……............................................…………. 2. Maranta 

2. Flowers purplish, sessile, pendent, exterior staminode 1; leaves 30-60 cm long, up to 25 cm wide 

……...........................................................................................................................…. 3. Thalia 

1. CALATHEA 

Calathea G. Mey., Prim. Fl. Esseq. 6. 1818. 

Perennial herbs with rhizomes; leaves all basal or both basal and cauline; aerial stems usually 

unbranched. Leaves ovate to elliptic, rarely obovate, the apex obtuse to rounded, or sometimes acuminate, 

glabrous or pubescent. Inflorescence terminal on a leafy shoot, or sometimes terminating a separate nonleafy 

scape, simple or synflorescence of several spikes, the latter often clustered. Bracts spirally arranged 

or distichous and green, white, or variously colored, usually persistent; bracteoles usually present, 

membranous. Flowers paired, usually sessile, opening or remaining closed, rarely cleistogamous; outer 

staminode 1 (rarely none); corolla tube elongate; ovary with 3 fertile locules, each with 1 ovule. Capsules 

3-locular, 3-seeded; seeds with a basal, usually white aril. A Neotropical genus of more than 300 species, 

only two of which are indigenous to Puerto Rico. Many species are grown for their ornamental foliage, and 

few of these sometimes escape and become naturalized. 

LECTOTYPE: Calathea discolor G. Mey., nom. illeg. ( ≡ Maranta casupo Jacq.), designated by Leman, 

Bull. Sci. Soc. Philom. Paris 1820: 7. 1820. 

Key to the species Calathea 

1. Spikes solitary, ellipsoid or oval, mostly 5-8 cm long, bracts closely spiral; corolla white 

….........................................................................................................................………… 1. C. allouia 

1. Spikes 2-6 in a synflorescence, mostly 8-16 cm long, subcylindric or rectangular. ……….......…….. 2 

2. Spikes flattened-rectangular; bracts 16-34, distichous, light yellow or orange-yellow; corolla 

white ………….................................................................................................…. 2. C. crotalifera 

2. Spikes subcylindric; bracts 8-12, spirally arranged (but often appearing distichous in pressed 

specimens), bronze to reddish brown; corolla yellow ….........................……………. 3. C. lutea 

1. Calathea allouia (Aubl.) Lindl., Bot. Reg. 14: t. 

1210. 1829; Maranta allouia Aubl., Hist. Pl. 

Guiane 3. 1775. Lectotype: Martinique. 

Plumier, manuscript no. 5: t. 35, designated 

by H.A. Kennedy in G. Harling and L. 

Andersson (eds.), Fl. Ecuador 32: 87. 1988.

196 


Plants rather robust, to 1.5 m tall. Leaf blades 

oblong to ovate-oblong, 30-50 × 8-15 cm, pale 

beneath, the apex short-acuminate, the base acute 

to rounded and somewhat inequilateral. Peduncles 

glabrous or pilose, each arising from a leaf-axil; 

bracts ovate, 1.5-3.5 cm long, green, membranous, 

glabrous or appressed-pubescent. Sepals lanceolate, 

obtuse, 7 mm long; corolla tube 2.5 cm long. 

Capsule 8 mm long. 


America, Greater Antilles (except Cuba), Lesser 

Antilles, and Northern South America. Some of 

this distribution may not be natural, as this species 

is cultivated for its edible tubers and easily 

escapes. 


moist, sheltered, disturbed habitats at middle 

elevations (300-500 m). Recorded from Cayey, 

Hatillo, Maricao, and Yabucoa but probably 

occurs more widely. 

Common names: Puerto Rico: Lerenes, 

Llerenes. 


Cayey: Guavate, Liogier et al. 36763 (UPR). 

Hatillo: Bo. Bayaney, Proctor 44086 (US). 

Maricao: Sintenis 492 (US). Yabucoa: Cerro de 

Pandura, Santa Elena, Liogier et al. 30980 (UPR). 

2. Calathea crotalifera S. Watson, Proc. Amer. 

Acad. Arts 24: 86. 1889. Lectotype: 

Guatemala; Izabal. Watson 429 (GH), 

designated by Standley & Steyermark, 

Fieldiana, Bot. 24: 210. 1952. 

Calathea insignis Petersen in Martius, Fl. Bras. 3 

(3): 124. 1890. Syntypes: Central America. 

Oersted s. n. (C, F, photo at XAL), and 

Panama; Canal Zone, Gatun, Wagner s.n. 

(M). 

Calathea quadratispica Woodson, Ann. Missouri 

Bot. Gard. 26: 278. 1939. Type: Panama; 

Bocas del Toro. Woodson et al. 1913 

(holotype: MO). 

Caulescent herb 1.5 m tall or more, bearing 2- 

5 basal leaves and 1 cauline leaf above a stem 

internode; leaf blades ovate, chartaceous, 30-90 × 

15-55 cm, rounded to minutely acuminate at the 

apex, the base rounded or subtruncate, lighter 

beneath, often tinged purple along margins; 

petioles elongate. Inflorescence rarely simple, 

usually a synflorescence of 2-4 spikes, the first one 

terminal, subsequent ones fasciculate in the axil of 

the cauline leaf. Spikes rectangular, laterally 

flattened, 10-25 × 3.5-6.5 cm, on peduncles 11-55 

cm long, these minutely appressed-tomentose at 

the apex. Bracts conduplicately folded, broadly 

reniform, glabrous or minutely tomentulose at 

base and along margins. Sepals glabrous, 12-21 

mm long; corolla tube pilose to glabrous, mostly 

18-25 mm long; staminal tube 3-5 mm long, the 

lobes subequal; outer staminode obovate, 

emarginate, cream to yellow, occasionally tinged 

purple, 9-13 mm long. Ovary glabrous, ca. 3.5 mm 

long. Capsules obovoid, rounded at apex, smooth, 

11-14 mm long, with persistent sepals; seeds dark 

blue, 6-7.5 mm long, with white aril. 

General distribution: widespread and common 

in moist lowlands of Central and tropical South 

America. 


from Río Grande, where it has become naturalized 

in secondary forest. Its time and mode of 

introduction to Puerto Rico are not known. 


Río Grande: El Verde area, secondary rain forest, 

elevation 210-240 m, Proctor 50414 (SJ). 

3. Calathea lutea (Aubl.) Schult., Mant. 1: 8. 

1822; Maranta lutea Aubl., Hist. Pl. Guiane 4. 

1775. Type: Plumier, manuscript no. 5: t. 21, 

22., fide H.A. Kennedy in G. Harling and L. 

Andersson, Fl. Ecuador 32: 87. 1988. 

Fig. 64. H 

Stout caulescent herb mostly 2-3 m tall, with 

3-7 basal leaves and 1 (2) cauline leaves; leaf 

blades elliptic to ovate or sub-rotund, 30-150 × 18- 

60 cm, obtuse, broadly rounded, or apiculate at the 

apex, rounded or very shortly decurrent at the base, 

usually glabrous, the upper surface green, the 

lower surface usually pruinose, sometimes coated 

with a white, wax-like powder that falls off in 

flakes. Inflorescence compound, of 2-5 more or 

less peduncled spikes 8-17 cm long, subcylindrical 

(broadly elliptic in cross-section), 2-5 cm thick or 

more; bracts usually 5-12 (rarely more), spirally 

arranged, broadly elliptic to depressed-elliptic, 

retuse at apex, 3.7-5.8 cm long, glabrous or nearly 

so. Sepals membranous, linear, unequal, 6-11 mm 

long; corolla tube curved, 25-34 mm long, the 

lobes strongly recurved, elliptic to obovate, 14-18 

mm long. Ovary smooth, glabrous, 3-3.5 mm long.


Capsule obovoid, rounded at apex, orange, 14-15 

mm long; seed 1, with orange ail. 

General distribution: Central America, 

Jamaica, Puerto Rico, Lesser Antilles, Trinidad, 

and tropical South America. 


Fajardo, Luquillo, Río Grande, and San Lorenzo. 

Common names: Puerto Rico: Hoja de sal, 

Pámpano. 

Note: The large leaves of Calathea lutea bear 

on their underside a white, flaky material which in 

Brazil is called “cauassu wax”. This is said to have 

considerable commercial value in the making of 

high-quality polishes. 


Luquillo: Río de Mameyes, Eggers 1169 (US). Río 

Grande: Luquillo Mountains, Caribbean National 

Maranta L., Sp. Pl. 2. 1753. 

Forest, Acevedo-Rdgz. et al. 10767 (UPRRP, US); 

Sierra de Luquillo, Sintenis 1327 (US). San Juan: 

Río Piedras, Stevenson 528 (US). 


Calathea zebrina (Sims) Lindl., a native of 

Brazil, is cultivated in Puerto Rico but known only 

to have escaped by only a single specimen record 

made in a ravine in wet forest at Jajome Arriba 

(Liogier & Liogier 30623). 


Calathea ornata (Lindl.) Körn. Britton & P. 

Wilson (1923) cited this species, a native of 

northern South America, as being cultivated in 

Puerto Rico; however, no specimen record has 

been seen by the present writer. 

2. MARANTA 

Perennial herbs with rhizomes. Leaves basal and often cauline; cauline leaves (if present) separated 

by distinct internodes. Spathes long-persistent, usually more or less fibrous. Inflorescence loosely and 

divaricately paniculate, the pedunculate branches consisting of 2-6, 2-flowered cymules, the terminal 

flower of each pair long-pedicelled, the lower flower sessile or nearly so. Sepals 3, equal; corolla tubular 

and usually enlarged or gibbous at the base, the 3 lobes subequal and somewhat hooded at the apex, the 

whole flower very irregular in appearance. Two exterior staminodes petal-like, obovate and conspicuous, 

the single anther free from its petaloid filament. Capsule more or less oblique; seed 3-angled, rugose, 

arillate. A Neotropical genus of about 30 species. 

TYPE: Maranta arundinacea L. 

Reference: Andersson, L. 1986. Revision of Maranta subgen. Maranta (Marantaceae). Nordic J. Bot. 

6: 229-256. 

Key to the species of Maranta 

1. Plants acaulescent, less than 1 m tall, with specialized starch-storing tubers produced on the roots, dying 

back during dry season; leaf blades pubescent on the underside ……...........……. 1. M. arundinacea 

1. Plants caulescent, 1-4 m tall; not producing specialized tubers on the roots, not dying back during dry 

season; leaf blades glabrous beneath (or with hairs only on the mid rib) ….......………… 2. M. gibba 

1. Maranta arundinacea L., Sp. Pl. 2. 1753. 

Lectotype: a specimen (Maranta 1) in the 

Clifford Herbarium (BM), designated by L. 

Andersson, Nordic J. Bot. 6: 739, 740. 1986. 

Maranta indica Tussac, Fl. Antill. 1: 183, t. 26. 

1808. Type: Jamaica. Tussac s.n. (holotype: 

probably at P). If no original material exists, 

this taxon can be typified by the figure cited. 

Acaulescent perennial herb 30-80 (-100) cm 

tall, commonly cultivated and escaping; rhizomes 

producing specialized tuberous branches which 

store starch, these covered with whitish scale-like 

cataphylls; whole plant dying back to the rhizome 

during dry seasons. Leaves basal except for a few 

produced at forks of the elongate stem-like 

inflorescence; petioles more or less hirsute, up to 

20 cm long; pulvinus hirtellous or hirsute above, 

usually glabrous beneath, 0.2-1.5 cm long on basal 

leaves; leaf blades narrowly ovate to ovate, 4-27 ×

198 


2.8-7 cm, acuminate at the apex, rounded or 

truncate at the base, glabrous on upper surface. 

Inflorescence diffusely branched, making up the 

entire stem-like aerial portion of the plant; flowers 

paired; sepals 10-15 mm long; corolla white, the 

tube 10-15 mm long, the lobes 8-10 mm long; 

outer staminode obovate, emarginate, the inner 

one shorter. Ovary glabrous or pubescent on the 

angles. Seed 8-9 mm long, transversely ridged, 

pale red with yellow aril. 

General distribution: Numerous records from 

Mexico, Central America, the Greater and Lesser 

Antilles, and northernmost South America; 

apparently absent from Brazil. An unknown 

number of recorded sites may be based on plants 

escaped or naturalized after cultivation. 


Islands: Naturalized on moist stream banks and 

also sometimes in shaded ditches and along 

roadsides at low to lower middle elevations (10- 

400 m). Recorded from Cataño, Cayey, Fajardo, 

Maricao, Mayagüez, Naguabo, San Juan, and San 

Sebastián; St. Croix. 

Common names: Puerto Rico: Amaranta, 

Maranta, Pitisilén, Yuquilla. 


Cataño: Goll 998, 1065 (US). Cayey: Sintenis 

2208 (US). Maricao: Sintenis 490 (US). Mayagüez: 

Underwood & Griggs 112 (US). San Juan: Río 

Piedras, Stevenson 3027 (US). San Sebastián: 

Sargent 412 (US). ST. CROIX: Big fountain garden, 

Ricksecker 445 (US). 

2. Maranta gibba Sm. in Rees, Cycl. 22. 1819. 

Lectotype: a specimen collected in the 

Liverpool Botanical Garden in 1810 (LIV), 

designated by L. Andersson, Nordic J. Bot. 6: 

749. 1986. 

Maranta divaricata sensu authors, non Roscoe, 

1902. 

Thalia L., Sp. Pl. 1193. 1753. 

3. THALIA 

Caulescent perennial herb with stems 1-4 m 

tall; rhizomes strongly fibrous and woody but 

lacking tuberous starch-filled offshoots. Mature 

plants with only bladeless sheaths at the base; first 

stem internode 1-2 m long, terminated by a node 

bearing either a sheath or a normal leaf blade; 

plants with multiple branches above the first node, 

the branches often long and straggling through 

surrounding vegetation; most branches eventually 

terminating in an inflorescence. Leaves without 

petioles, with hirsute pulvinus 1-6 mm long; leaf 

blades narrowly ovate to ovate, 3.2-20 × 2-9 cm, 

acuminate at the apex, rounded at the apex, 

glabrous on both surfaces except sometimes a few 

coarse hairs on the mid rib. Inflorescences 

terminal on leafy branches, simple or compound. 

Each inflorescence with 1 or 2 spathes; spathes 

glabrous or subglabrous, 2.7-4.6 cm long. 

Inflorescence composed of 1 or 2 cymules, each 

cymule with a peduncle 4-5.5 cm long and two 

unequal pedicels varying from 0.1 to 2 cm long. 

Sepals greenish, glabrous, 12-14 mm long; corolla 

and staminodes white; corolla tube 15-18 mm 

long, the lobes 8-10 mm long; outer staminodes 

subequal, 14-22 mm long; cucullate staminode 9- 

10 mm long; ovary densely sericeous, with 

indument more or less completely covering the 

surface. Capsules nearly ellipsoid to subglobose, 

but very oblique, 11-12 mm long when fresh, 

smaller when dry. Seed rugose, 6-8 mm long. 

General distribution: Southern Mexico to 

Nicaragua, Virgin Islands, St. Vincent, Tobago, 

Trinidad, and northernmost South America from 

Ecuador to the Guianas. 

Distribution in the Virgin Islands: Recorded 

only from St. Thomas by a single collection made 

long ago. The continued existence of this species 

in St. Thomas is in doubt and needs to be 

investigated. 

Erect perennial herbs of marshy or fresh-water aquatic habitats, rhizomes very short, branched. 

Leaves chiefly in a basal cluster, with long petioles. Inflorescence a branched, panicle-like (rarely simple) 

synflorescence terminal on a tall peduncular stem. Flowers in pairs, sessile within 2-valved bract-like 

spathes, these deciduous, shed with the unfertilized flowers or mature fruits, leaving a naked more or less 

zigzag rachis below the currently flowering apical portion of the spike. Sepals 3, equal, very small,


membranous; corolla tube obsolete or nearly so, the lobes membranous, more or less cucullate at the apex; 

outer staminode petaloid, conspicuous, very irregular. Ovary 1-locular with 1 ovule; style helically 

twisted, adnate to the base of the staminal tube; stigma 2-lipped, appendaged on the back. Capsule globose 

or oblong-obovoid, indehiscent, with very thin pericarp which is more or less papery when dry. Seed 

smooth, ellipsoid, with small but distinct aril. A Neotropical genus of about 7 species, with one species 

now widely naturalized in tropical Africa. 

TYPE: Thalia geniculata L. 

Reference: Andersson, L. 1981. Revision of the Thalia geniculata complex (Marantaceae). Nordic J. 

Bot. 1: 48-56. 

1. Thalia geniculata L., Sp. Pl. 1193. 1753; 

Maranta geniculata (L.) Lam., Tabl. Encycl. 

1: 9. 1791. Lectotype: Plumier (Burman ed.), 

Pl. Amer. t. 58, fig. 1. 1755, designated by L. 


Thalia angustifolia C. Wright ex Griseb., Cat. Pl. 

Cub. 256. 1866. Lectotype: Cuba; Nueva 

Filipina. Wright 3279 (BM), designated by L. 


Thalia trichocalyx Gagnep., Bull. Soc. Bot. France 

51: 180. 1904. Lectotype: French Guiana. 

Richard s. n. (P), designated by L. Andersson, 

Nordic J. Bot. 1: 55. 1981. 

Plants 1-3 m tall, usually glabrous or nearly 

so; basal leaves long petiolate, sheathing at base 

and at the apex with a cylindrical pulvinus 1.5-2.4 

cm long; leaf blades ovate to oblong, elliptic or 

lanceolate, mostly 30-60 cm long, up to 25 cm 

wide, acute or acuminate at the apex, rounded at 

the base; cauline leaves (if present) smaller and 

sessile; all leaves often somewhat glaucous 

beneath. Flowers borne in pairs on the zigzag 

rachis of a spike; spikes several to many in a 

compound panicle-like synflorescence; spathes 


glabrous to densely villous, 1-2.4 cm long; sepals 

glabrous or more or less setose outside, 0.5-2 mm 

long; corolla 6-11 mm long; outer staminode 

lavender, 15-20 mm long, 5-10 mm broad. 

Capsule oblong-obovoid, 7-10 mm long; seed 5- 

10 mm long, with a small but evident aril. 


States, Mexico, Central America, Greater Antilles, 

and northern South America; also common and 

widespread in tropical Africa, where it is believed 

to have been introduced. 


Islands: This species is mostly confined to freshwater 

habitats near sea level. It has been 

obliterated from many sites by urban development 

and the draining of wetlands. Recorded from 

Añasco, Arecibo, Carolina, Cataño, Manatí, 

Mayagüez, Moca, Río Grande, and San Juan; St. 

Croix. 


Añasco: Sintenis 5602 (US). Arecibo: Sargent B20 

(US). Manatí: Bo. Tierras Nuevas Saliente, 

Proctor & Díaz 42172 (US-2). Mayagüez: N.L. 

Britton 2365 (US). 

Myrosma cannifolia L. f. is cited by Liogier & Martorell (1982) as cultivated in Puerto Rico. 

Family 29. BROMELIACEAE Pineapple Family 

Bromeliaceae Juss., Gen. Pl. 49. 1789, nom. conserv. 

by. J. A. Cedeño-Maldonado 

Epiphytic, lithophytic or terrestrial perennial herbs; rhizomatous or sometimes stoloniferous; roots 

usually present, but functioning only as support in epiphytic species. Stems well-developed or short,

200 


sometimes wanting, erect to arching or pendent, often bracteate. Leaves simple, sheathing at base, usually 

in rosettes which are often water-impounding; blades linear, lanceolate, or rarely ovate, thick and usually 

stiffened, often with stalked, peltate, water-absorbing scales, margins spinose, serrate or entire, the apex 

sometimes terminated by a thorn. Inflorescences terminal or rarely lateral, of simple or compound 

panicles, racemes, spikes, or heads, rarely with solitary flowers, the axes with distichous or spirallyarranged 

flowers; bracts conspicuous, usually present in compound inflorescences. Flowers 3-merous, 

bisexual or functionally unisexual, actinomorphic or rarely zygomorphic, often showy; sepals 3, free or 

connate; petals 3, free or connate; stamens 6, in 2 series of 3, the filaments free or fused to the petals; 

anthers basifixed or peltate, introrse; ovary superior or inferior, 3-locular; placentation axile; style slender, 

with 3 stigmatic branches or lobes. Fruit a dehiscent septicidal or less often loculicidal capsule or berry. 

Seeds 1-many per locule, winged, caudate, plumose, or unappendaged. A primarily Neotropical family of 

56 genera and ca. 3,000 species distributed from the southeastern United States to southern South America 

with one species native to tropical West Africa. 

Type: Bromelia L. 

References: Holst, B. K. 1997. Bromeliaceae. Pp. 548-676. In: J. A. Steyermark, P.E. Berry, & B. 

Holst (eds.). Flora of the Venezuelan Guayana. Missouri Botanical Garden Press, St. Louis, Missouri. 

Howard, R. A. 1979. Bromeliaceae. Flora of the Lesser Antill. 3: 404-426. Arnold Arboretum of Harvard 

University, Jamaica Plain, MA. Proctor, G. R. & J. A. Cedeno-Maldonado. 1999. New Bromeliaceae from 

Puerto Rico. Harvard Pap. Bot. 4 (1): 111-118. Smith, L. B. & R. J. Downs. 1974. Pitcairnioideae 

(Bromeliaceae). Fl. Neotrop. Monogr. 14 (1): 1-658. Smith, L. B. & R. J. Downs. 1977. Tillandsioideae 

(Bromeliaceae). Fl. Neotrop. Monogr. 14 (2): 663-1492. Smith, L. B. & R. J. Downs. 1979. Bromelioideae 

(Bromeliaceae). Fl. Neotrop. Monogr. 14 (3): 1493-2142. Utley, J. F. 1994. Bromeliaceae. Pp. 89-156. In: 

G. Davidse, M. Sousab, & A. O.Chater (eds.). Flora Mesoamericana. Vol. 6. Utley, J. F., K. Burt-Utley & 

M. J. Huft. 2001. Bromeliaceae. In: W. D. Stevens, C. Ulloa-Ulloa, A. Pool & O. M. Montiel (eds.). Flora 

of Nicaragua, Monographs in Systematic Botany 85 (1): 460-495. Missouri Botanical Garden Press, St. 

Louis, Missouri. Zanoni, T. A., M. M. Mejia P., and R. W. Read. 1986. Notas sobre la flora de la Isla 

Espanola. I. Bromeliaceae. Moscosoa 4: 54-104. 


1. Leaves with spiny margins ……….................................................................................................….. 2 

2. Plants terrestrial; sterile portion of the scape short, stout, to about 18 cm long ……….........…. 3 

3. Plants small to medium sized; leaves to 1 m long; inflorescence simple; gynoecium and bracts 

adnate to axis to form a fleshy compound fruit …………..............................….. 2. Ananas 

3. Plants large; leaves to 2 m long; inflorescence compound; gynoecium free, fruits distinct 

berries ………….............................................................................................… 3. Bromelia 

2. Plants epiphytic, lithophytic or terrestrial; sterile portion of the scape elongate, more than 20 cm 

long ……................................................................................................................................… 4 

4. Leaf blades 1-2 cm wide; fruits capsules …………..............................………… 7. Pitcairnia 

4. Leaf blades more than 2 cm wide; fruits baccate ……….......................................……….. 5 

5. Inflorescence simple or compound; spikes cylindrical; flowers and fruits widely spaced; 

petals without basal appendages ………………...........................……….. 1. Aechmea 

5. Inflorescence compound; spikes cone-like; flowers & fruits congested; petals with basal 

appendages ………......................................................................…….. 6. Hohenbergia 

1. Leaves with entire margins.................................................................................................................... 6 

6. Plants evidently caulescent, with elongate branching stems; leaves to 8 cm long but usually less 

…………….......................................................................................…….. 8. Tillandsia (in part) 

6. Plants acaulescent or with very short, unbranched stems; leaves usually more than 8 cm long 

..................................................................................................................................................... 7 

7. Inflorescences simple …………...........................................................................………… 8 

8. Inflorescences polystichously-flowered or corymbiform ………................…………. 9


9. Leaf sheaths slightly enlarged or indistinct, leaf apex obtuse to rounded and 

apiculate; scape bracts much shorter than internodes; petals free; fruit ovoid 

………….......................................................................................…… 4. Catopsis 

9. Leaf sheaths distinctly enlarged, leaf apex acute to acuminate; scape bracts equaling 

or usually much longer than the internodes; petals connate; fruits cylindricalellipsoid 

or fusiform ………........................................................…… 5. Guzmania 

8. Inflorescences distichously- or secundly-flowered ……….............................………. 10 

10. Plants large; leaves 50-100 cm long; blades 5-8 cm wide; scape to 2 m long; capsule 

4-6 cm long …………......................................................................….. 9. Vriesea 

10. Plants smaller, all other parts smaller than above ………........................…….. 11 

11. Leaf blades usually reddish green, transversely red-banded; petals with basal 

appendages; stigma cupulate (three apical, capitate, cup-shaped untwisted 

lobes without papillae) …………….......................…. 10. Werauhia (in part) 

11. Leaf blades green, concolorous or with white transverse stripes; petals 

unappendaged; stigma never cupulate ………………… 8. Tillandsia (in part) 

7. Inflorescences compound …………….....................................................................…….. 12 

12. Branches of the inflorescence polystichously-flowered; fruits ovoid ……… 4. Catopsis 

12. Branches of the inflorescence not polystichously-flowered; fruits cylindrical-ellipsoid 

or fusiform ……...........................................................................................……….. 13 

13. Petals with basal appendages; stigma cupulate ………….. 10. Werauhia (in part) 

13. Petals unappendaged; stigma never cupulate ………………. 8. Tillandsia (in part) 

1. AECHMEA 

Aechmea Ruiz & Pav., Fl. Peruv. Prodr. 47. 1794, nom. conserv. 

Epiphytic, lithophytic or terrestrial, usually acaulescent herbs. Leaves numerous, in dense rosettes; 

bases sheathing, enlarged; blades linear-attenuate, acute or mostly obtuse to rounded, acuminate to 

apiculate; margins usually spinose-serrate. Inflorescence scapose, simple or compound. Flowers bisexual, 

distichous or polystichous, sessile or stipitate. Sepals free or connate, usually strongly asymmetric and 

mucronate; petals symmetrical, free, bearing 2 basal appendages; stamens shorter than petals, free or the 

second series adnate to the petals, unappendaged; anthers dorsifixed; ovary inferior. Fruit baccate; seeds 

numerous, small, naked, dark colored. A genus of about 170 species from the Neotropics. 

TYPE: Aechmea paniculata Ruiz & Pav.. 

Key to the species of Aechmea 

1. Inflorescence open, evidently compound, the branches distichous, more than 9 cm long 

….........................................................................................................................……… 2. A. lingulata 

1. Inflorescence dense, simple or with polystichous branches, 2-5 cm long. ………...................………. 2 

2. Inflorescence ovoid-pyramidal, flower bracts exceeding the flowers ………….…… 1. A. fasciata 

2. Inflorescence cylindrical, flower bracts shorter than the flowers or wanting …….. 3. A. nudicaulis 

1. Aechmea fasciata (Lindl.) Baker, J. Bot. 17: 

231. 1879; Billbergia fasciata Lindl., Bot. 

Reg. 13: pl. 1130. 1828. Type: Brazil; Rio de 

Janeiro. Lindl., Bot. Reg. 13: pl. 1130. 1828. 

Epiphytic, acaulescent, rhizomatous herbs. 

Leaves numerous in a rosette, pale-appressedlepidote 

on both sides, coriaceous, 30-110 × 3-8 

cm; sheaths large, slightly wider than the blade, 

ovate-elliptic, unarmed; blades linear, attenuate, 

the margins armed with dark spines to 2 mm long; 

apex acute to rounded, apiculate. Scape erect, 50- 

60 cm long, white flocculose; sterile bracteoles 

lanceolate, sheathing at base, acuminate, pungent, 

serrate, 7-9 cm long, the lower loosely arranged 

and erect, the upper congested and subspreading.

202 


Inflorescence simple or with few basal branches, 

densely pyramidal-ovoid, 7-10 cm long; primary 

fertile bracteoles like the sterile ones but smaller, 

exceeding the branches; spicate branches erect, 

ovoid, densely-flowered, 2-5 cm long; bracts rose 

lanceolate or ovate, long acuminate, exceeding the 

sepals, strongly serrulate. Flowers polystichous, 

sessile; sepals asymmetric, mucronulate, or acute, 

1-1.2 cm long; petals ligulate, 2.5-3 cm long, blue, 

purple or red when dry; stamens included, 

filaments of the second series adnate to the petals; 

ovary ellipsoid. 

General distribution: Native to Brazil, one of 

the most popular bromeliads for horticulture. 


cultivated as an ornamental, known to have 

naturalized in a Pterocarpus forest in Humacao. 


Humacao: Proctor 48637 (SJ). 

2. Aechmea lingulata (L.) Baker, J. Bot. 17: 164. 

1879; Bromelia lingulata L., Sp. Pl. 285. 

1753; Chevalliera lingulata (L.) Griseb., Fl. 

Brit. W. I. 591. 1864; Wittmackia lingulata 

(L.) Mez in Martius, Fl. Bras. 3(3): 275. 1891. 

Lectotype: Plate 170 of the Boerhaave set of 

Plumier illustrations at the library of 

Rijksuniversiteit, Groningen, here designated. 

Aechmea plumieri Baker, Handb. Bromel. 50. 

1889. Type: America. Plumier (Burman ed.), 

Pl. Amer. 1: t. 64, f. 1. 1756. 

Fig. 40. A-F 

Epiphytic, terrestrial or lithophytic herb; 

stems to 12 cm long, erect or decumbent. Leaves 

rosulate, rigid, coriaceous, to ca. 1 m × 2-10 cm, 

lepidote throughout; sheaths broader than blade, 

ovate-oblong, unarmed; blade linear, attenuate; 

margins serrate with dark, upwardly curved 

spines, 1-2 mm long; apex acute to rounded, 

apiculate. Scape slender, erect, to 1 m long, 

floccose when young; sterile bracteoles erect, 

lanceolate or oblong, sharply acuminate, 4-8 cm 

long. Inflorescence very large and open, 

distichously branched; primary fertile bracteoles 

like the sterile ones but mostly shorter; branches 

several, ascending, 9-16 cm long, polystichouslyflowered. 

Flowers sessile; bracts deltate-subulate, 

4-10 mm long; sepals 5-6 mm long, asymmetrical, 

subapically cuspidate; petals acute, 7-10 mm long; 

stamens included, those of second series adnate to 

petals; ovary ovoid, ellipsoid, cylindrical or 

clavate. Berries ovoid, 8-10 mm long, purple-red. 

General distribution: Costa Rica, Puerto 

Rico, Virgin Islands, the Lesser Antilles, 

Bahamas, Trinidad, Tobago, Venezuela, Guyana, 

Surinam, French Guiana, and northern Brazil. 


Islands: Woods, thickets and edges of forest at low 

elevations (0-400 m), from xeric to moist 

conditions, usually in rocky situations. A rare 

species not collected in Puerto Rico since 1914. 

Liogier (1982) reported it from Punta Picúa at Río 

Grande but no voucher was located, it is otherwise 

known from Vieques and Culebra; St. John, St. 



Culebra: N.L. Britton & Wheeler 96 (US). San 

Juan: Río Piedras, Stevenson 1805 (US). Vieques: 

vicinity of Isabel Segunda, Shafer 2488 (US). ST. 

JOHN: Vélez 3102a (US); Maho Bay Quarter; 

Center Line Road, Acevedo-Rdgz. et al. 4035 (NY, 

US). ST. THOMAS: E.G. Britton & Marble 1428 

(US); Eggers 1200 (US). 

3. Aechmea nudicaulis (L.) Griseb., Fl. Brit. W. I. 

593. 1864; Bromelia nudicaulis L., Sp. Pl. 

286. 1753. Lectotype: Plate 171 of the 

Boerhaave set of Plumier illustrations at the 

library of Rijksuniversiteit, Groningen, here 

designated. 

Epiphytic, lithophytic or terrestrial, herb; 

stems short. Leaves numerous, rigid, coriaceous, 

30-80 × 2-9 cm, punctate-lepidote throughout or 

glabrescent above, forming a dense fasciculate, 

funnelform rosette, the outer sometimes reduced 

and scale-like; sheaths large, elliptic, slightly 

wider than the blades, unarmed; blades linear, 

attenuate, the margins serrate with dark spines, 2- 

4 mm long; apex obtuse to rounded, apiculate. 

Scape slender, erect or nodding, shorter than the 

leaves, white floccose; sterile bracteoles erect, 

imbricate, the upper congested below the 

inflorescence, 5-10 cm long, elliptic, oblong or 

lanceolate, narrowly acute to long-acuminate, 

chartaceous, weakly and minutely armed or entire, 

red. Inflorescence a simple spike, cylindrical, 

fertile throughout, 5-12 cm long; bracts entire, 

subulate, much shorter than the flowers, 

shortening towards the apex of the inflorescence, 

sometimes wanting. Flowers sessile; sepals


asymmetric, mucronate, 5-10 mm long; petals 

acute, ca. 1.2 cm long, greenish; ovary densely 

pale-lepidote. Berry ovoid to globose, orange to 

brownish red. 

General distribution: Mexico, Nicaragua, 

Costa Rica, Panama, Cuba, Hispaniola, Puerto 

Rico, Venezuela, Trinidad & Tobago, Guyana, 

Ecuador, and Peru. 

Distribution in Puerto Rico: Rare, mostly 

rocky situations on edges of forest, from sea level 

to 730 m. Recorded from Aibonito, Arecibo, 

Barranquitas, Río Grande, and Salinas. 

Note: There has been disagreement with 

respect to the typification of Linnean names that 

were based on Plumier elements (Polhill & Stearn 

in Taxon 25:323-325. 1976.). An example of this 

disagreement involves both A. lingulata, and A. 

nudicaulis. Smith & Downs (1979) typified these 

species with Plumier specimens, collected from 

the West Indies during the Surian expedition in 

1689-1690. These typifications are erroneous, 

since it is well known that Linnaeus never saw the 

Surian material. Howard (1979), on the other 

hand, typified A. lingulata with a Plumier plate (Pl. 

Amer., t. 64, fig. 1. 1756.). This has erroneously 

been accepted as the correct typification even 

Ananas Mill., Gard. Dict. Abr. ed. 4. 1754. 

though this figure was published later than 

Linnaeus’ 1753 Species Plantarum. However, 

there is convincing evidence that Linnaeus studied 

the artist Claude Aubriet’s tracings of Plumier’s 

plates made for Herman Boerhaave at Leyden, and 

used information from these plates to prepare 

diagnoses for the species he described (C. Jarvis & 

D. Nicolson, pers. comm.; Polhill & Stearn in 

Taxon 25: 323-325. 1976). Type material of these 

two taxa are among the Boerhaave set of Plumier 

illustrations (Codex Boerhaavianus) deposited at 

the library of Rijksuniversiteit, Groningen, the 

Netherlands. 


Aibonito: Liogier 37425 (UPR). Arecibo: 

Garrochales, Stevenson 2399 (UPR, US). Río 

Grande: Proctor 42674 (SJ). Salinas: Proctor 

42400 (SJ); Bo. Lapa, Proctor 42501 (SJ). 


Aechmea paniculata Ruiz & Pav., Syst. Veg. 

Fl. Peruv. Chil. 83. 1798. Britton & P. Wilson 

(1923) note that this taxon was reported for St. 

Thomas by Krebs based on an error in 

determination. 

2. ANANAS 

Terrestrial herbs. Leaves numerous, densely rosulate, linear-attenuate, coriaceous, stiff, the margins 

armed with hooked spines, the sheaths slightly enlarged. Inflorescence scapose, densely spicate, usually 

crowned with several, fascicled, imbricate, sterile, foliaceous bracts. Flowers bisexual, sessile; sepals 

free, erect; petals free, ligulate; stamens included; ovary inferior, style filiform. Fruit compound (formed 

by the fusion of the gynoecia, bracts, and axis), fleshy. A genus of seven species native to tropical South 

America. 

NEOTYPE: Bromelia ananas L. (≡ Bromelia comosa L.) (= Ananas comosus (L.) Merr.), designated 

by L. B. Smith, N. Amer. Fl. 19: 214. 1938. 

1. Ananas comosus (L.) Merr., Interpr. Herb. 

Amboin. 133. 1917; Bromelia comosa L., 

Herb. Amboin. 21. 1754. Type. Rumphius, 

Herb. Amboin. 5: t. 81. 1747, designated by 

Merrill, Interpr. Herb. Amboin. 33, 133. 1917. 

Bromelia ananas L., Sp. Pl. 285. 1753; Ananas 

sativus Schult. & Schult. f., Syst. Veg. 7: 

1283. 1830, as a new name; Ananas ananas 

(L.) H. Karst. ex Voss, Vilm. Blumengärtn., 

ed.3, 1: 964. 1895., nom. inadmiss. Type: To 

be sought among several syntypes. 

Ananassa monstrosa Carr., Rev. Hort. 42: 288. 

1870; Ananas monstrosus (Carr.) L. B. Sm., 

Phytologia 8: 12. 1961. Type: Carrière s.n. 

(probably at P). 

Terrestrial, acaulescent plant, highly variable 

with many cultivated varieties. Leaves to 1 m × 2- 

4 cm, linear-lanceolate to deltate-attenuate, 

lepidote on both surfaces, the margins upwardly 

spinose-serrate, cuspidate. Scape short and stout, 

to ca. 18 cm long, shorter than the leaves;

204 


inflorescence simple, conical-ovoid, subtended 

and crowned with spiny, green, leafy bracts; 

flowers numerous, erect, sessile; corolla violet or 

red. Fruits compound, fleshy, varying considerably 

in size, 10-25 cm long, golden yellow, crowned 

with spiny green bracts; seeds typically wanting. 

General distribution: Native of Brazil, widely 

cultivated in the tropics. 

Distribution in Puerto Rico: A major crop in 

Puerto Rico, cultivated for its fruit; persistent after 

Bromelia L., Sp. Pl. 285. 1753. 

3. BROMELIA 

cultivation and naturalizing, especially on the 

northern coastal plain. Reported as naturalized in 

Vega Baja (surroundings of Laguna Tortuguero). 

Common names: Puerto Rico: Piña; Pineapple. 


Bayamón: Sintenis 1033 (US); Stevenson 3925 

(US). Vega Baja: Bo. Algarrobo, white sand area 

just S of Laguna Tortuguero, Proctor et al. 47737 

(US). 

Large, terrestrial herbs, spreading by rhizomes or stolons. Leaves in rosettes, cuspidate, the margins 

with large curved spines. Inflorescence sessile or scapose, mostly paniculate. Flowers bisexual; sepals free 

to connate, obtuse or acute; petals connate at base; stamens included; ovary inferior. Fruit baccate, 

succulent, indehiscent; seeds few to many, flattened, unappendaged. A genus of about 50 species from 

tropical America, extending from Mexico to Argentina, and the West Indies. 

LECTOTYPE: Bromelia karatas L., designated by Regel, Gartenflora 17: 67. 1868. 

1. Bromelia pinguin L., Sp. Pl. 285. 1753. 

Lectotype: Dillenius, Hort. Eltham. 320, t. 

240, f. 311. 1732, designated by R.A. Howard, 

Fl. Lesser Antill. 3: 409. 1979. 

Figs. 40. G-J; 65. A 

Large terrestrial herb, usually in colonies. 

Leaves numerous, in a rosette, much exceeding the 

inflorescence; sheaths broad, dark brown, densely 

lepidote; blades linear, long-attenuate at apex, 

rigid, ca. 2 m × 4 cm; margins armed with stout, 

hooked spines to 10 mm long; apex pungent. Scape 

stout, 8-15 cm long, white floccose; sterile 

bracteoles subfoliaceous, the sheaths subinflated, 

the upper shorter. Inflorescence erect, to ca. 30 cm 

long, white floccose, paniculate, pyramidal, 

many-flowered; fertile bracteoles like the scape 

bracteoles, the upper ones entire; bracts subulate, 

to 2.5 cm long. Flowers pedicellate, whitefloccose, 

3-6 cm long; sepals erect, narrowly 

deltate-subulate; petals pink, whitish towards 

margins and base, to 3 cm long, linear-elliptic, 

white-tomentose at apex; stamens ca. 2 cm long. 

Capsules ovoid, 3-4 cm long, yellow, verrucose, 

white-floccose, beaked. Seeds covered with a 

creamy acid pulp. 


America, Greater Antilles, Virgin Islands, Lesser 

Antilles, Colombia, Venezuela, Surinam, and 

Ecuador. 


Islands: Thickets and hedges, disturbed areas, and 

waste grounds. Of ample occurrence, probably in 

the majority of the municipalities, recorded from 

Aibonito, Arecibo, Cabo Rojo, Cayey, Comerío, 

Corozal, Fajardo, Guánica, Loíza, Maricao, Río 

Grande, Río Piedras, Salinas, San Juan, and 

Vieques; St. Croix, St. John, and St. Thomas. 

Common name: Puerto Rico: Maya. 

Note: Smith & Downs (1979) cited a 

specimen in the Clifford Herbarium at BM as the 

type of Bromelia pinguin. However, we have not 

been able to locate such a collection at BM, 

therefore we are following Howard’s 

lectotypification made a few months earlier than 

Smith & Downs publication. 


Fajardo: Sintenis 950 (US). San Juan: Río Piedras, 

Stevenson 517 (US). Vieques: Isabel Segunda to 

Campo Cielo, Shafer 2337 (US). ST. CROIX: 

Ricksecker 342 (US). Christiansted, Rose et al. 

3546 (US). East End, Ricksecker 284 (US). Ridge 

SW of Lang Peak (Jakobsberg), above Eliza’s 

Retreat, Fosberg & Ogden 55267 (US). ST. JOHN: 

Lind Point area, Acevedo-Rdgz. 2798 (NY, US, 

MO, US, UPR, VINPS). ST. THOMAS: Morrow 93 

(US).


Fig. 40. A-F. Aechmea lingulata. A. Habit. B. Leaf. C. Inflorescence branch. D. Flower, l.s. flower, and detail of stigma. E. 

Stamen adnate to petal, and free stamen. F. Portion of infructescence, and berry. G-J. Bromelia pinguin. G. Habit. H. Section of 

leaf blade. I. Inflorescence branch. J. Flower, and l.s. flower. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York 

Bot. Gard. 78.

206 



Bromelia plumieri (E. Morren) L. B. Sm., 

Phytologia 15: 173. 1967, nom. Illeg. (≡ 

Bromelia karatas L.). Reported from Puerto Rico 

by Bello (Anales Soc. Esp. Hist. Nat. 12: 121. 

1883); this is the only record of the species for the 

island, and may have been based on a specimen in 

cultivation (Britton & P. Wilson, 1923). 

4. CATOPSIS 

Catopsis Griseb., Nachr. Königl. Ges. Wiss. Georg-Augusts-Univ. 1864: 10, 12. 1864. 

Epiphytic acaulescent herbs. Leaves in a rosette, minutely appressed-lepidote, entire, green. 

Inflorescences scapose, spicate or paniculate, exceeding the leaves, the branches polystichous-flowered. 

Flowers small, sessile or subsessile, bisexual or functionally unisexual; sepals free, rounded, strongly 

asymmetric; petals free, naked, white or yellow; stamens included; ovary superior, ovoid or ellipsoid; 

style short or missing; capsule septicidally dehiscent above the middle; seeds thick, the coma apical and 

folded over. A genus of about 20 species from tropical and subtropical America and the West Indies. 

LECTOTYPE: Catopsis nitida (Hook.) Griseb. (≡ Tillandsia nitida Hook.), designated by J. R. Grant & 

Zijlstra, Selbyana 19: 97. 1998. 

Key to the species of Catopsis 

1. Sheaths distinctly dilated, blades deltate-attenuate, the apex acuminate ….....……… 1. C. floribunda 

1. Sheaths slightly dilated to indistinctly so, blades linear, the apex obtuse to rounded and apiculate ... 2 

2. Leaves 10-40 cm long, closely imbricate in a cylindrical rosette; inflorescence always branched; 

sepals 3-5 mm long, strongly asymmetric with one side cuneate at base and the apex much 

expanded into a large wing; capsules 6-11 mm long ………...................………….. 2. C. nitida 

2. Leaves 10-25 cm long, more or less loosely imbricate in a cylindrical-funnelform rosette; 

inflorescence simple or few branched; sepals 6-8 mm long, weakly asymmetric; capsules 12-15 

mm long ………….......................................................................................….. 3. C. sessiliflora 

1. Catopsis floribunda L. B. Sm., Contr. Gray 

Herb. 117: 5. 1937. Holotype: Martinique. 

Pleé s. n. (P, not seen). 

Catopsis nitida sensu Duss, Fl. Phan. Antill. Fr. 

577. 1897, in part, non (Hook.) Griseb., 1864. 

Catopsis nutans sensu Britton & P. Wilson, Bot. 

Porto Rico 5: 137, 1923 and Liogier, 

Phytologia 47(3): 197. 1980, non Grisebach, 

1864. 

Fig. 43. D-F 

Epiphytic, acaulescent herb, commonly 

forming clusters. Leaves numerous, densely 

rosulate, sub-erect , entire, punctulate-lepidote on 

both surfaces, 10-40 cm long; sheaths ovateelliptic, 

to 8 cm wide; blades deltate-attenuate, 

long-acuminate, 1.5-3 cm wide at the base. Scape 

erect, 2-4 mm in diam., glabrous, much longer than 

the leaves; sterile bracteoles erect, lanceolate, the 

lower ones foliaceous, the upper ones longer than 

the internodes. Inflorescence paniculate, many- 

flowered, to 40 cm long; fertile bracteoles 

subspreading, ovate-lanceolate, long acuminate, 

1.5-3 cm long; the branches ascending, 1-2 mm 

wide; bracts subspreading, ovate, acute to 

acuminate, 3-5 mm long. Flowers sessile or 

subsessile, sub-erect ; sepals elliptic, asymmetric, 

4-6 mm long; petals white, elliptic, ca. 7 mm long, 

obtuse. Capsule ovoid, acute, 9-11 mm long. 

General distribution: Southern Florida, 

Central America, Greater Antilles, Virgin Islands, 

Lesser Antilles, Trinidad & Tobago, and 

Venezuela. 


Islands: In moist forest, from near sea level to 900 

m. Recorded from Aibonito, Barranquitas, 

Bayamón, Caguas, Camuy, Cayey, Ciales, 

Guayama, Jayuya, Maricao, Naguabo, Orocovis, 

Río Grande, Salinas, Villalba, and Yauco; St. 

Croix, St. John, St. Thomas, and Virgin Gorda. 



Bayamón: Stevenson 1144 (US). Cayey: Sintenis 

2043 (US). Ciales: E slopes of Los Tres Picachos, 

Axelrod & Axelrod 4830 (US). Guayama: Bo. 

Carmen, along crest (S. side) of Sierra de Jajome 

2-2.8 km. due NE-ENE of village of Carmen, 

Proctor & Rivera 46921 (US); 3 mi S of Guayama 

on Rt. 15, Grant & Rundell 93-02267 (US). 

Jayuya: Sargent 3072 (US). Maricao: Monte 

Alegrillo, N.L. Britton et al. 2560 (US). Naguabo: 

Sierra de Luquillo, Loma La Mina, SE side to E 

Peak, Shafer 3257 (US). Orocovis: Toro Negro 

Forest Reserve, Lago Guineo, Ackerman & 

Meléndez 2662 (US). Río Grande; Caribbean 

National Forest, El Yunque, Acevedo-Rdgz. 2963 

(NY, US). Villalba: Along road 143 (north of 

Villalba) at km 45 hm 8, Stimson 1668 (US). ST. 

JOHN: Bordeaux, N.L. Britton & Shafer 572 (NY). 

ST. THOMAS: Eggers s.n. (US). VIRGIN GORDA: 

Gorda Peak National Park, Acevedo-Rdgz. & 

Clubbe 10899 (US). 

2. Catopsis nitida (Hook.) Griseb., Fl. Brit. W. I. 

599. 1864; Tillandsia nitida Hook., Exotic Fl. 

3. 1826. Type: Jamaica. Wiles s. n. (holotype: 

LIV, not seen). 

Epiphytic, acaulescent herb, producing stolons 

and commonly forming clumps. Leaves few, 

closely imbricate in a cylindrical rosette, suberect, 

entire, 10-40 × 2-4 cm, the outer usually 

smaller; sheaths indistinct; blades linear-attenuate; 

apex obtuse to rounded, apiculate. Scape erect to 

ascending, slender, glabrous; sterile bracteoles 

erect, sheathing, ovate-deltate, acute, 5-10 mm 

long. Inflorescences laxly branched, to 20 cm 

long, exceeding or equaling the leaves, glabrous; 

fertile bracteoles like the sterile bracteoles but 

usually smaller; branches naked below, manyflowered 

above, ascending to subspreading, ca. 1 

mm in diam.; bracts ovate, acute, shorter than the 

sepals. Flowers sessile, bisexual; sepals strongly 

asymmetric, ca. 5 mm long; petals white or cream, 

elliptic, obtuse, 6-8 mm long; style wanting. 

Capsule ovoid, acute, 6-11 mm long. 

General distribution: Southern Mexico, 

Guatemala, Honduras, Nicaragua, Costa Rica, 

Panama, Cuba, Jamaica, Hispaniola, and Puerto 

Rico. 

Distribution in Puerto Rico: Moist to wet 

montane forest above 500 m elev. Recorded from 

Adjuntas, Barranquitas, Ciales, Jayuya, Maricao, 

Naguabo, Orocovis, Río Grande, Utuado, and 

Villalba. 


Adjuntas: vicinity of Pico Guilarte, Liogier 10009 

(US). Barranquitas: Bo. Barrancas, upper NE 

slopes of Monte Torrecilla, Proctor & Concepción 

42308 (US). Jayuya: Los Tres Picachos, Sargent 

3172 (US). Maricao: Sargent 225 (US). Naguabo: 

Bo. Río Blanco, Caribbean National Forest, 

Axelrod & Chavez 4130 (US). Utuado: 7 mi N 

from Adjuntas along road to Arecibo, Grant & 

Rundell 93-02296 (US). Villalba: along border w/ 

Orocovis, W slopes of Cerro Doña Juana, between 

Cerro Doña Juana and Cerro el Bolo, Ackerman & 

Calvo 2467 (US). 

3. Catopsis sessiliflora (Ruiz & Pav.) Mez in C. de 

Candolle, Monogr. Phan. 9: 625. 1896; 

Tillandsia sessiliflora Ruiz & Pav., Fl. Peruv. 

3: 42. 1802. Type: Peru; Huánuco. Ruiz & 

Pavón s. n. (holotype: MA?, isotype: B, photo 

at GH). 

Epiphytic, acaulescent herb, usually in 

clumps. Leaves few in a cylindrical-funnelform 

rosette, 10-25 × 1.5-3.5 cm, linear-attenuate, 

entire, sub-erect ; sheaths indistinct; blades linearattenuate; 

apex acute to mostly rounded or obtuse 

and apiculate. Scape sub-erect , glabrous, 1-2 mm 

in diam., 15-30 cm long; sterile bracteoles erect, 

elliptic-ovate, apiculate, 8-13 mm long. 

Inflorescence a simple spike or with few ascending 

branches, exceeding the leaves; fertile bracteoles 

like the sterile ones; spikes laxly- to denselyflowered, 

subspreading, the axis 1-2 mm wide; 

bracts ovate, acute or obtuse, shorter than the 

sepals. Flowers sessile, sub-erect ; sepals 

asymmetric, 6-8 mm long; petals white, ovatelanceolate, 

about as long as the sepals. Capsule 

ovoid, short beaked, 12-15 mm long. 


Guatemala, Honduras, Costa Rica, Panama, Cuba, 

Hispaniola, Puerto Rico, Guadeloupe, Martinique, 

Grenada, Colombia, Venezuela, Trinidad & 

Tobago, Guyana, Surinam, French Guiana, 

Ecuador, Peru, and Brazil. 

Distribution in Puerto Rico: Rare, known 

only from four specimens. Moist and wet forest, 

from sea level to ca. 600 m. Recorded from 

Maunabo, Naguabo, and San Lorenzo. 


208 


Maunabo: Maunabo Beach Area, Woodbury s. n. 

(UPR). Naguabo: Sierra de Naguabo, Bo. de 

Maizales, N.L. Britton & Cowell 2188 (US); 

5. GUZMANIA 

Guzmania Ruiz & Pav., Fl. Peruv. 3: 37. 1802. 

Caribbean National Forest, Río Sabana Hansen et 

al. 9409 (UPR). San Lorenzo: Hato Grande, 

Monte Gregorio, Sintenis 2466 (US). 

Terrestrial, lithophytic or mostly epiphytic and usually acaulescent herbs. Leaves polystichousrosulate, 

entire, linear-attenuate, green. Inflorescences scapose, simple (or compound outside Flora area), 

shorter than or exceeding the leaves. Flowers bisexual; sepals oblong, symmetrical; petals lacking basal 

appendages, connate proximally and forming a tube; stamens equal, inserted on the corolla throat; ovary 

superior; style elongated. Capsule septicidally dehiscent; seeds basally comose. A genus of about 167 

species from tropical and subtropical America and the West Indies. Named in honor of Anastasio Guzmán, 

a Spanish naturalist. 

TYPE: Guzmania tricolor Ruiz & Pav. 

Key to the species of Guzmania 

1. Inflorescence spicate, elliptic-cylindrical ……….................................................................……… 2 

1. Inflorescence densely corymbiform, torch-like or capitate ….………. 2. G. lingulata var. concolor 

2. Leaves 30-70 cm long; leaf blades just above the sheath 2-5 cm wide; bracts 4-7 cm long; sepals 

2-2.5 cm long; corolla yellow, 5-7 cm long …………….........…….. 1. G. berteroniana 

2. Leaves 20-50 cm long; leaf blades just above the sheath 1.5-3 cm wide; bracts 2-3.5 cm long; 

sepals 1.5-2.0 cm long; corolla white, 2-3 cm long …………..........………. 3. G. monostachia 

1. Guzmania berteroniana (Schult. & Schult.f.) 

Mez in C. de Candolle, Monogr. Phan. 9: 904. 

1896 (as “berteroana”); Caraguata 

berteroniana Schult. & Schult.f., Syst. Veg. 

7(2): 1229. 1830; Tillandsia caraguata D. 

Dietr., Syn. Pl. 2: 1059. 1840; nom. illeg. 

Type. Puerto Rico. Bertero s. n. (holotype: B). 

Caraguata grandiflora Baker, Handb. Bromel. 

145. 1889. Type: Puerto Rico; Maricao. 

Sintenis 471 (holotype: K; isotypes GH, US!). 

Guzmania erythrolepis sensu Britton & P. Wilson, 

Bot. Porto Rico 5: 143. 1923, non Brongniart 

ex Planchon. 

Fig. 65. C 

Terrestrial or epiphytic, usually large, 

acaulescent herb; usually forming large colonies 

when terrestrial. Leaves numerous, fasciculate in a 

dense rosette, 30-70 cm long, ascending to 

arching, entire, obscurely punctulate-lepidote 

toward the base; sheaths inconspicuous to much 

broader than the blades, ovate-elliptic, brown; 

blades linear-attenuate, 2.5-5 cm wide, acute or 

acuminate. Scape erect, glabrous, shorter than the 

leaves; sterile bracteoles erect, densely imbricate, 

the lower ones subfoliaceous, the upper ones 

shorter, red, their sheaths becoming larger and 

broader and their blades reduced toward the apex 

of the scape. Inflorescence a simple, cylindricalelliptic 

spike, 15-30 cm long, the apex usually 

sterile or with aborted flowers; bracts dark red, 

erect, densely imbricate, chartaceous, nerved, 

broadly elliptic, 4-7 × 1.5-3 cm, smaller toward the 

apex of the inflorescence, the lower ones abruptly 

acuminate-apiculate and the upper ones acuteapiculate. 

Flowers bisexual, subsessile, solitary; 

sepals elliptic, acute to obtuse, sometimes bifid, 

connate at base, 1.5-2.5 cm long; corolla yellow, 5- 

7 cm long, the lobes elliptic-oblong, acute to 

obtuse; stamens and stigma inserted. Smith & 

Downs (1979) describe its capsules as subprismatic, 

acute, and 3 cm long, and its seeds as having a 

white coma. 

General distribution: Panama, Dominican 

Republic, and Puerto Rico. 

Distribution in Puerto Rico: Moist and wet 

mountain forests, above 200 m. Recorded from 

Aibonito, Arecibo, Barranquitas, Canóvanas,


Cayey, Ciales, Guayama, Jayuya, Luquillo, 

Maricao, Naguabo, Ponce, Patillas, Río Grande, 

Salinas, Sabana Grande, San Germán, Utuado, 

Yabucoa, and Yauco. 

Common names: Puerto Rico: Parásita, 

Pirigallo. 



10641 (US). Barranquitas: La Torrecilla, trail 

from summit ridge down to forest road, Axelrod 

6993 (US). Guayama: Mt. between Guayama and 

Cayey, N.L. Britton et al. 6573 (US). Maricao: 

Sargent 670 (US); N.L. Britton et al. 2587 (US). 

Río Grande: Caribbean Natl. Forest, along trail to 

Mt. Britton, Thompson & Thompson 3182 (US). 

. 

2. Guzmania lingulata var. concolor Proctor & 

Cedeño-Mald., var. nov. Type: Puerto Rico; 

Sierra de Luquillo, Loma La Mina, J. A. 

Shafer 3279 (holotype: US). 

Fig. 41. A-C 

Folia numerosa in rosula densa, 25-50 cm 

larga, interiora breviora, integra, viridia 

concolora; vaginae plerumque distinctae, ovatae, 

punctulato-lepidotae; laminae lineari-attenuatae, 

ca. 4 cm latae, acutae vel acuminatae. Bracteae 

florales virides vel rubrae, erectae, lanceolatae 

vel lineari-attenuatae, acuminato-acutae, 

cucullatae, sepalis multo longiores. 

Terrestrial or epiphytic, acaulescent herb, 

usually forming colonies at the base of tree trunks. 

Leaves numerous, in a dense rosette, 25-50 cm 

long, the inner ones shorter, entire, green, 

concolorous; sheaths mostly distinct, ovate, 

punctulate-lepidote; blades linear-attenuate, ca. 4 

cm wide, acute to acuminate. Scape erect, stout, 

shorter than the leaves, 12-25 cm long; sterile 

bracteoles erect, densely imbricate, the lower ones 

foliaceous, the uppermost lanceolate, green, red or 

reddish, forming a cup-shaped involucre below the 

inflorescence. Inflorescence apical, simple, 

densely-flowered, corymbiform, with a 

mucilaginous substance within; bracts green or 

red, erect, lanceolate to linear-attenuate, acuminateacute, 

cucullate, much longer than the sepals. 

Flowers bisexual, pedicellate, erect; sepals free, 

linear, obtuse-rounded, 1.5-2 cm long; corolla 

white, the lobes linear, obtuse-rounded, cucullate; 

stamens adnate to the petals. Capsule oblong- 

ellipsoid, short beaked, 2.8-3.5 cm long; seeds 

with reddish brown coma. 

General distribution: Belize, Cuba, 

Hispaniola, Puerto Rico, the Lesser Antilles, 

Trinidad & Tobago, northern South America south 

to Bolivia. 


Islands: Moist and wet forest, ca. 200 m and 

above. Recorded from Adjuntas, Arecibo, 

Bayamón, Caguas, Camuy, Naguabo, San Germán, 

San Lorenzo, San Sebastián, and Yabucoa. 

Note: Several varieties have been attributed 

to Guzmania lingulata (L.) Mez , mostly based on 

the size and color patterns of the leaves and the 

color of the bracts of the inflorescences (see L. B. 

Smith, Fl. Neotrop. Monogr. 14(2): 1349. 1977.). 

The variety with wide and concolorous leaves, 

which occurs in Puerto Rico, has been erroneously 

attributed to G. lingulata var. lingulata. This 

variety does not occur in Jamaica, where Sloan 

collected the type specimen for G. lingulata. On 

the other hand, only one of the varieties has been 

recognized to occur in Jamaica. Instead of being 

considered the typical variety, the Jamaican taxon 

was erroneously attributed to G. lingulata var. 

splendens (Planch.) Mez. Thus, to correct the 

situation, var. splendens, with purplish striped 

leaves, must be considered a synonym of var. 

lingulata. In addition, a new name is hereby 

provided for the taxon previously referred to as 

var. lingulata. 

Guzmania lingulata var. concolor differs 

from G. lingulata var. lingulata (of which G. 

lingulata var. splendens is a synonym) in that the 

former has concolorous leaves and the latter has 

leaves with red-purple longitudinal stripes. 


Arecibo; Río Abajo State Forest, Acevedo-Rdgz. 

10656 (US). Naguabo: Sierra de Luquillo, Loma 

La Mina, SE side to E peak, Shafer 3279 (US). San 

Sebastián: Bo. Cibao, along E side of Río 

Guajataca in gorge S of Lago de Guajataca, 

Proctor et al. 48108 (US). 

3. Guzmania monostachia (L.) Rusby ex Mez in 

C. de Candolle, Monogr. Phan. 9: 905. 1896; 

Renealmia monostachia L., Sp. Pl. 287. 1753; 

Tillandsia monostachia (L.) L., Sp. Pl. ed 2, 

410. 1762. Lectotype: West Indies. A plate of 

the Boerhaave set of Plumier illustrations at 

the library of Rijksuniversiteit, Groningen, 

here designated.

210 


Fig. 41. A-C. Guzmania lingulata var. concolor. A. Fertile plant. B. Fruit. C. Immature seed with tuft of hairs only partially 

expanded. From Mori, S. et al. 1997. Vascular plants of central French Guiana. Mem. NYBG Vol. 76(1). 

Guzmania tricolor Ruiz & Pav., Fl. Peruv. 3: 38. 

1802. Type: Peru. Ruiz & Pavón s. n. 

(holotype: MA, not seen). 

Fig. 65. B 

Epiphytic, or sometimes terrestrial, acaulescent 

herb. Leaves numerous, in a dense rosette, entire, 

20-50 cm long, the inner ones shorter, minutely 

punctulate-lepidote; sheaths conspicuous, broader 

than the blades, ovate-elliptic or ovate-oblong; 

blades linear-attenuate, 1.5-3 cm wide, acute to 

acuminate, green. Scape erect, stout, shorter than


the leaves, glabrous; sterile bracteoles erect, 

imbricate, the lowermost subfoliaceous, sheathing, 

densely imbricate, the upper ones broadly ovate, 

acuminate, pale green, slightly longer than the 

internodes. Inflorescence spicate, densely 

polystichous-flowered, cylindrical-elliptic, shorter 

than leaves, 12-23 cm long, sterile or with aborted 

flowers toward the apex; bracteoles erect, 

imbricate, chartaceous, ca. 2 × 2 cm, broadly ovate 

to quadrangular or obovate, the apex acuminate, 

acute or apiculate, the fertile ones pale green and 

usually with brownish purplish longitudinal 

stripes, the sterile ones pink, red or orange at the 

apex. Flowers bisexual, erect; sepals oblong to 

obovate, obtuse to rounded, connate at base, ca. 

1.5 cm long, smooth; corolla white, 2-2.5 cm long, 

the lobes imbricate, elliptic, obtuse. Capsule 

elliptic-cylindrical, 2-4 cm long, pointed; seeds 

with whitish comose hairs, ca. 2 cm long. 


Nicaragua to Peru, and Brazil, Bahamas, Greater 

Antilles, and Trinidad & Tobago. 


forests, ranging from near sea level to above 900 

m. Recorded from Adjuntas, Arecibo, Bayamón, 

Cayey, Ciales, Guayama, Río Grande, Salinas, 

6. HOHENBERGIA 

Hohenbergia Schult. f. in Roem. & Schult., Syst. Veg. 7(2): lxxi, 1251. 1830. 

San Lorenzo, Utuado, and Yabucoa. 


Adjuntas: Monte La Vega, Sintenis 4403 (US). 


10739 (US). Bayamón: Hato Tejas, Liogier 10371 

(US). Cayey: Bo. Cercadillo, Cerro Avispa, 

Proctor 42434 (US). Ciales: Bo. Toro Negro, 

ravine up N slope of Los Tres Picachos, Axelrod & 

Stenzel 11137 (US). Río Grande: Caribbean 

National Forest, El Yunque, Acevedo-Rdgz. 2962 

(MO, NY, UPR, US). Utuado: 10 mi N from 

Adjuntas along road to Arecibo, Grant & Rundell 

93-02293 (US). 


Guzmania erythrolepis Brongn. ex Planch. 

was erroneously reported for Puerto Rico by 

Britton & P. Wilson, Bot. Porto Rico 5: 143, 1923, 

based Britton et al. 2468 and 2587. Study of these 

collections revealed them to be Guzmania 

berteroniana (Schult. & Schult. f.) Mez. instead. 

The report of G. erythrolepis from the Dominican 

Republic (Smith & Downs, 1977) is also 

erroneous, and similarly based on the 

misidentification of specimens of G. berteroniana 

(T. Zanoni, pers. comm.). 

Medium to large, terrestrial, lithophytic or epiphytic, acaulescent herbs. Leaves in a rosette, 

polystichous; sheaths large, distinct, dark brown; blades linear, coriaceous, spinose-serrate, acuminate. 

Inflorescences scapose, compound (with one Brazilian exception), lanate or glabrous; secondary branches 

spicate, dense, ovoid-strobilate, bracteolate; bracts conspicuous, covering the ovary and often the sepals. 

Flowers bisexual, sessile; sepals asymmetric, distinct or connate at base, mucronulate; petals distinct or 

nearly so, unguiculate, the claw bearing two appendages; stamens included, the second series adnate to the 

petals; ovary inferior. Fruit baccate; seeds curved, unappendaged. A genus of about 48 species from the 

West Indies, northern South America to Brazil. 

LECTOTYPE: Hohenbergia stellata Schult. f., designated by Britton & P. Wilson, Bot. Porto Rico 5: 134. 

1923. 

Key to the Species of Hohenbergia 

1. Leaves 1 m long or longer, 6-13 cm wide; scape erect, stout, 4-20 mm thick; inflorescence densely 

branched; spicate branches 3-10 cm long; bracts 1.2-2.5 cm long, acute-punctate or gradually shortacuminate 

for ca. a third or less of their length ………….......................................….. 1. H. antillana 

1. Leaves usually less than 1 m long, 2-7 cm wide; scape erect, ascending or usually arching, 2-5 mm thick; 

inflorescence laxly branched; spicate branches 2-6 cm long; bracts 1-1.7 cm long, abruptly longacuminate 

(almost caudate) for ca. half or more of their length ……………………. 2. H. portoricensis

212 


1. Hohenbergia antillana Mez in C. de Candolle, 

Monogr. Phan. 9: 137. 1896. Lectotype: 

Puerto Rico; Cayey. Sintenis 2000 (US). here 

designated. 

Hohenbergia tetaensis Proctor & Cedeño-Mald., 

Harvard Pap. Bot. 4: 111. 1999. Type: Puerto 

Rico; Cayey. Proctor 44657 (holotype: US; 

isotypes: IJ, SJ) 

Large, epiphytic, terrestrial or lithophytic, 

acaulescent and rather variable herb, usually 

forming colonies. Leaves numerous, rosulate, 

coriaceous, stiff, arching, to ca. 1.5 m × 6-13 cm, 

minutely appressed-lepidote on both surfaces; 

sheaths slightly wider than the blades, ellipticovate, 

dark brown, the margins entire or spinulose 

near the apex; blades ligulate, the margins armed 

with dark spines 1-2.5 mm long, the apex broadly 

acute to rounded, short-acuminate, terminating in 

a stout, dark mucro. Scape erect, stout, 4-20 mm 

wide, densely creamy-white lanate-floccose; 

bracteoles erect, lanate-floccose, striate, sheathing, 

imbricate, concealing the scape, the margins 

weakly armed or entire, lanceolate, attenuate to 

acuminate, with a dark pungent tip. Inflorescence 

twice-branched, ovoid to ellipsoid or cylindrical, 

erect, exceeding or shorter than the leaves, 

creamy-white lanate or floccose throughout, 10- 

30 cm long, congested, especially toward the apex; 

lowest internodes to 2 cm long; primary bracteoles 

like those of the scape, the lower ones usually 

exceeding the branches, the upper ones usually 

shorter than the branches; spicate branches 15-20, 

strobilate, ovoid or ellipsoid to subcylindric, 

polystichously arranged, usually of simple spikes, 

or the lowermost ones with a small, secondary, 

sessile spike at the base; bracts broadly triangularovate, 

lanate, coriaceous, striate, equaling or 

exceeding the sepals, 1.2-2.5 cm long, acute or 

gradually short-acuminate for ca. a third or less of 

their length, pungent. Flowers sessile, polystichous; 

sepals strongly asymmetric, keeled, mucronate, 

0.5-1.2 cm long; petals white, clavate, exceeding 

the sepals by 4-6 mm; ovary subglobose, enlarged 

in fruit. Fruits baccate, flattened-ovoid to 

pyriform, coriaceous, lanate, two-winged, ca. 1.5 

cm long. 




Islands: Thickets, woodlands and moist forests; 

more common in the northern karst region; from 

sea level to above 800 m. Recorded from Arecibo, 

Cabo Rojo, Cayey, Cidra, Dorado, Isabela, 

Quebradillas, Río Grande, Salinas, San Sebastián, 

Utuado, and Vega Baja; Guana Island, Scrub and 

Great Dog Islands. 

Note: Hohenbergia tetaensis, a species 

published by Proctor & Cedeño-Maldonado 

(1999), is included here as a synonym of H. 

antillana. Additional fieldwork conducted by 

Cedeño-Maldonado between 1999 and 2004 at the 

type locality of H. tetaensis, revealed individuals 

attributable to H. tetaensis and H. antillana 

coexisting at the site, even within the same colony 

or clump. Individuals with intermediate 

morphologies were also found at the same and 

other localities in Puerto Rico. 


Arecibo: 5 km S of Biáfara, Acevedo-Rdgz. & 

Chinea 11646 (US); Acevedo-Rdgz. & Siaca 

11700 (US). 

2. Hohenbergia portoricensis Mez in C. de 

Candolle, Monogr. Phan. 9: 136. 1896. Type: 

Puerto Rico; Sierra de Naguabo. Sintenis 

1321 (holotype: B, destroyed). 

Hohenbergia attenuata Britton in Britton & P. 


Puerto Rico; Sierra de Naguabo. Shafer 3567 

(holotype: NY!). 

Epiphytic, terrestrial and lithophytic, 

acaulescent and rather variable herb, usually 

forming clumps. Leaves numerous, rosulate, 

minutely appressed-lepidote on both surfaces, to 

ca. 1.5 m long but usually less than 1 m long, 2-6 

cm wide; sheaths conspicuous or inconspicuous, 

mostly wider than the blade, elliptic-ovate, entire 

except at the apex; blades ligulate, the margins 

armed with dark, curved spinules 1-1.5 mm long, 

which become smaller closer to the apex; apex 

obtuse to acute or acuminate, apiculate, minutely 

spinulose or entire. Scape erect, ascending or 

usually arching, 2-5 mm thick, covered with lanate 

or floccose creamy-white indumentum at least 

when young; bracteoles erect, chartaceous, striate, 

lanate-glabrescent, spinulose, imbricate, lanceolate, 

attenuate to acuminate, pungent. Inflorescence 

thyrsoid, erect to arching, open, lowest internodes 

ca. 2.5 cm long, shorter toward apex, exceeding or 

shorter than the leaves, 15-30 cm long, sparsely or


densely whitish cream-lanate or floccose 

throughout, the axis 2-3 mm thick; bracteoles like 

the scape ones but spreading and entire, the lower 

ones exceeding the spikes, shortening toward the 

apex of the inflorescence; spicate branches 

subglobose, ovoid or subcylindric, strobilate, 

densely-flowered, 2-6 × 1.5-2 cm, those toward 

the apex of the inflorescence smaller and sessile; 

bracts 1-1.7 cm long, equaling or exceeding the 

sepals, subspreading, subcoriaceous, ovate, striate, 

abruptly long-acuminate for ca. half or more of its 

length, pungent. Sepals strongly asymmetric, 

ovate to deltoid-sagittate, keeled, mucronate, 5-10 

mm long; petals linear-lanceolate, acute, ca. 11 

mm long. Fruit pyriform, striate, glabrescent, 

slightly two-winged, ca. 1.2 cm long; seeds 2-2.5 

mm long, purplish. 

7. PITCAIRNIA 

Pitcairnia L’Hér., Sert. Angl. 7. 1789, nom. conserv. 


Rico. 

Distribution in Puerto Rico: Slopes, ridges 

and peaks, usually in rocky outcrops, of moist and 

wet montane forests above 500 m. Recorded from 

Aibonito, Barranquitas, Cayey, Ciales, Guayama, 

Las Piedras, Luquillo, Maricao, Naguabo, Patillas, 

San Germán, San Lorenzo, and Yabucoa. 


Ciales: Bo. Toro Negro, E slope of Los Tres 

Pichachos, Axelrod & Axelrod 6700 (US). 

Naguabo: Sierra de Naguabo, Barrio de Maizales, 

N.L. Britton & Shafer 2141 (US); Sierra de 

Naguabo, Shafer 3336, 3567 (US). Patillas: Sierra 

de Cayey, Carite Forest Reserve, Proctor & 

Zorilla 49375 (US). 

Small to large, terrestrial, lithophytic or seldom epiphytic herbs with short to elongate stems. Leaves 

numerous, fasciculate or in a dense spiral along the stem, dimorphic; sheaths usually small, sometimes 

bulbose thickened; blades linear-lanceolate, spinulose-margined or entire. Inflorescence usually scapose, 

simple or compound; bracts conspicuous to minute. Flowers bisexual, showy, pedicellate to sessile; sepals 

free convolute; petals free, long and narrow, convergent over the stamens, naked or with a single basal 

appendage; stamens about as long as the petals, 6, with linear anthers; ovary superior to inferior, with a 

long and slender style; ovules numerous. Fruit capsular, three-valved, septicidally dehiscent; seeds 

numerous, variously winged. A genus of about 260 species from tropical America and the West Indies, 

with one African species. Named in honor of Dr. William Pitcairn, 1711-1791. 

TYPE: Pitcairnia bromeliifolia L’Hérit. 

Key to the species of Pitcairnia 

1. Leaf blades 1-2 cm wide; bracts ovate-lanceolate, 5-10 mm long; flowers red; seeds with creamy-white 

appendage, tapering at base; Puerto Rico, Virgin Islands …………….................… 1. P. angustifolia 

1. Leaf blades 0.8-1.2 cm wide; bracts ovate, 3-6 mm long; flowers yellow; seeds with brownish yellow 

appendage, truncate at the base; endemic to Guana Island (B.V.I.) ……..……………… 2. P. jareckii 

1. Pitcairnia angustifolia Aiton, Hort. Kew. 1: 

401. 1789. Type: St. Croix, U. S. Virgin. Is. 

Ryan s. n., (holotype: BM, photo at GH). 

Pitcairnia latifolia Aiton, Hort. Kew. 1: 401. 

1789. Type: West Indies. Anderson s. n. 

(BM). 

Pitcairnia ramosa J. Jacq., Eclog. Pl. 1: 154, 117, 

f. 79. 1816. Lectotype: J. Jacq., Eclog. Pl. 1: 

f. 79. 1816, here designated. 

Pitcairnia gracilis Mez in C. de Candolle, 

Monogr. Phan. 9: 407. 1897. Type: 

Guadeloupe. L’Hérminier s. n. (holotype: G, 

photo at GH). 

Lithophytic or terrestrial herb; acaulescent or 

with stems to 10 cm long. Leaves numerous, in a 

dense fasciculate rosette, coriaceous, fimbriatelepidote 

abaxially, 50-120 cm long, sometimes 

dimorphic with some reduced and forming tufts of 

dark-rigid spines at the base of the rosette; sheaths

214 


conspicuous, triangular-ovate, dark brown, densely 

imbricate and usually forming a pseudobulb; 

blades linear-attenuate, long-acuminate, ca. 1-2 

cm wide, arching, the margins armed with dark 

curved spines ca. 2-3 mm long. Scape erect or 

ascending, to 1.7 m tall, flocculose-glabrescent; 

bracteoles subfoliaceous, serrate, erect, linearlanceolate, 

acuminate, 5-12 cm long, exceeding 

the internodes basally but shorter above. 

Inflorescence few- to many- branched; bracteoles 

like the upper scape ones, much shorter than the 

branches; branches laxly-flowered, ascending to 

spreading; bracts ovate to lanceolate, acuminate, 

5-10 mm long. Flowers erect or sometimes 

pendent, pedicels slender, 4-12 mm long; sepals 

oblong, acuminate to obtuse, 1.3-2.3 cm long; 

petals red, 4-5.5 cm long, linear, acute; stamens 

about as long as the petals. Capsules ovoid, 

trigonous, apiculate, 1.2-2 cm long. Seeds ellipticclavate, 

ca. 2 mm long, narrowly winged along one 

side; the wing creamy white, obliquely-enlarged at 

the apex and tapering at base 

Note: In Puerto Rico this species exhibits two 

different morphologies which are also 

geographically defined. In most of the Island, the 

plants always have compound inflorescences. In 

the Sierra de Luquillo (Caribbean National 

Forest), the vast majority of the plants have simple, 

unbranched racemes. Although Read (Phytologia 

8(5): 222. 1962) and Smith & Downs (Fl. Neotrop. 

Monogr. 14(1): 300. 1974) described the 

inflorescences of P. angustifolia as simple or 

branched, in Puerto Rico these two morphologies 

are evidently and consistently distinct. A new 

varietal name is provided here, for the taxon with 

simple, unbranched inflorescences, which in 

Puerto Rico is limited to the Sierra de Luquillo. 

Key to the varieties of Pitcairnia angustifolia 

1. Inflorescences with numerous usually 

compound branches; pedicels 5-8 mm long, 

sepals 1.4-2 cm long; Puerto Rico and U.S. 

Virgin Islands ………..…. 1. var. angustifolia 

1. Inflorescences usually unbranched or with a 

few basal, short and simple branches; pedicels 

6-15 mm long, sepals 1.6-2.6 cm long; mostly 

limited to the Sierra de Luquillo, eastern Puerto 

Rico ……….....................…. 1. var. simplicior 

1. Pitcairnia angustifolia var. angustifolia 

Fig. 43. A-C 

Inflorescences with numerous mostly 

compound branches; pedicels 5-8 mm long, sepals 

1.4-2 cm long. 

General distribution: Puerto Rico, Virgin 

Islands, Antigua, Saba, St. Eustatius, St. Kitts, 

Montserrat, Guadeloupe, Dominica, Martinique, 

St. Lucia, St. Vincent, the Grenadines, Grenada, 

and Barbados. 


Islands: Moist and wet forests from sea level to 

above 1000 m. Common along stream margins, 

road banks, ledges and exposed rocky outcrops. 

Recorded from Arecibo, Bayamón, Cabo Rojo, 

Coamo, Cayey, Dorado, Guayama, Hatillo, 

Jayuya, Juncos, Maricao, Mayagüez, Naguabo, 

Quebradillas, Río Grande, Salinas, San Germán, 

Toa Baja, Utuado, Vega Baja, Vieques, and 

Yauco; St. Croix, St. John, St. Thomas, and 

Tortola. 

Common names: Puerto Rico: Erizo, Píña 

cortadora, Píña de cuervo. 


Arecibo: Hato Arriba, N.L. Britton & Cowell 1981 

(US). Bayamón: Stevenson & Johnston 911 (US). 

Coamo: Coamo road, Goll et al. 633 (US). Río 

Grande: N Slope of El Yunque Mts., Read 2049b 

(US).Guayama: 3 mi. S of Guayama on Rt. 15, 

Grant & Rundell 93-02268 (US). Juncos: Monte 

Florida, Sintenis 2564 (US). Maricao: Sargent 419 

(US). Naguabo: along Río Cubuy to Meseta Falls, 

Shafer 3162 (US). Río Grande: El Yunque, 

Sargent 557 (US). Salinas: Aguirre & Underwood 

418 (US). San Germán: Sargent 443 (US). Toa 

Baja: Candelaria, Goll et al. 267 (US).Utuado: 

vicinity of Utuado, E.G. Britton 5152 (US).Vega 

Baja: Stevenson 456 (US). Vieques: Cerro 

Ventana, Shafer 2877 (US). Yauco: Underwood 

644 (US). ST. CROIX: Ryan s.n. (Type photo). ST. 

JOHN: Maho Bay Quarter; Waterlemon Bay, 

Acevedo-Rdgz. et al. 1940 (NY, VINPS). ST. 

THOMAS: Cowells Hill, Eggers s.n. (US). Water 

Island, N.L. Britton et al. 135 (US). TORTOLA: Road 

Town to Sea Cow Bay, N.L. Britton & Shafer 696 

(US). 

1. Pitcairnia angustifolia var. simplicior Proctor 

& Cedeño-Mald., var. nov. Type: Puerto 

Rico; Naguabo, Caribbean National Forest, 

Sierra de Luquillo, El Toro trail, Proctor 

47931 (holotype, SJ).


A P. angustifolia var. angustifolia 

inflorescentiis simplicibus differt. 

Inflorescences usually unbranched or with a 

few basal, short and simple branches; pedicels 6- 

15 mm long, sepals 1.6-2.6 cm long. 

General distribution: Similar specimens have 

been recorded from the island of Martinique; 

otherwise, this variety is apparently endemic to 

Puerto Rico. 

Distribution in Puerto Rico: Ridges, slopes, 

streams, banks and edges of forest, usually in 

rocky situations; wet montane forest, above 500 m. 

With the exception of one specimen collected at 

the Carite Forest Reserve, this variety is otherwise 

restricted to the Sierra de Luquillo. Recorded from 

Cayey, Naguabo and Río Grande. 


Naguabo: Sierra de Naguabo, Shafer 3162 (US). 

Río Grande: El Yunque, Sargent 557 (US); Read 

2049a, 2049b (US). 

2. Pitcairnia jareckii Proctor & Cedeño-Mald. sp. 

nov. Type: Guana Island, B.V.I., from ridge 

leading to Palm Point, elev. ca. 450-500 ft., 27 

Oct 1991, Proctor 47242 (holotype: SJ). 

Fig. 42. A-H 

Herba terrestris caulibus redactis. 

Inflorescentia ramosa ampla; rami ascendentes 

divergentes, ad 5 mm diametro, laxiflori, infra 

steriles. Bracteae ovatae acuminatae, 3-6 mm 

longae; flores plerumque erecti; sepala anguste 

deltata vel lineari-attenuata, acuta, 1-1.6 cm 

longa; petala flava, 3-4.5 cm longa, linearioblanceolata, 

acuta vel obtusa, aliquantum 

apiculata. Capsula ovoidea trigona, ca. 1.5 cm 

longa. 

Tillandsia L., Sp. Pl. 286. 1753. 

8. TILLANDSIA 

Terrestrial herb with reduced stems. Leaves 

numerous, in a dense fasciculate rosette, 

ascending, arching, coriaceous, fimbriate-lepidote 

abaxially, to ca. 1 m long; sheaths conspicuous, 

ovate, dark brown in the upper half, densely 

imbricate, forming a pseudobulb; blades linearattenuate, 

long-acuminate, 0.8-1.2 cm wide, the 

margins armed with spines 1-3 mm long. Scape 

erect or ascending, to ca. 1.5 m tall, flocculoseglabrescent; 

bracteoles subfoliaceous, spinulose, 

erect, narrowly deltate to lanceolate, longacuminate, 

exceeding the internodes basally but 

much shorter above. Inflorescence branched, very 

large and open; primary bracteoles like the upper 

scape ones; branches ascending, spreading, to 5 

mm in diam., laxly-flowered, sterile below; bracts 

ovate, acuminate, 3-6 mm long. Flowers mostly 

erect, pedicels slender, 5-8 mm long; sepals 

narrowly-deltoid to linear-attenuate, acute, 1-1.6 

cm long; petals yellow, 3-4.5 cm long, linearoblanceolate, 

acute to obtuse, somewhat apiculate; 

stamens equaling or shorter than the petals. 

Capsule, ovoid, trigonous, ca. 1.5 cm long. Seeds 

elliptic-clavate, ca. 2 m long; narrowly winged 

along one side; the wing brownish yellow, 

obliquely enlarged at the apex, truncate at the base. 

General distribution: Endemic to Guana 

Island 

Note: The main morphological differences 

between P. jareckii and P. angustifolia, the only 

other species of this genus that is found in Puerto 

Rico and the Virgin Islands, are that it has 

narrower leaves (0.8-1.2 cm vs. 1-2 cm), and 

yellow flowers vs. red flowers. 

Distribution in the Virgin Islands: Understory 

of dry forest, on rocky ridge of volcanic origin, 

450-500 ft. Known only from the type locality on 

Guana Island, B.V.I. 

Caulescent or acaulescent herbs with variable habit and size. Leaves fascicled-rosulate or spirally 

distributed along the stem, polystichous or distichous, entire; blades ligulate to narrowly triangular; scales 

peltate, centrally symmetric. Inflorescence various, usually distinctly scapose, of distichous-flowered 

spikes or sometimes reduced to a single polystichous-flowered spike by the reduction of the spikes to 

single flowers or rarely the whole inflorescence reduced to a single flower; bracts conspicuous to minute. 

Flowers bisexual, mostly short-pedicellate; sepals usually symmetric, free or connate; petals free, without 

appendages; stamens included or exserted; ovary superior, glabrous; ovules usually many, caudate. Fruit 

a septicidal capsule; seeds many, erect, narrowly cylindrical or fusiform, the plumose appendage basal,

216 


Fig. 42. A-H. Pitcairnia jareckii. A. Lower portion of habit with detail of leaf margin. B. Upper portion of habit and inflorescence. 

C. Inflorescence branch. D. Flower. E. Stamen. F. Apex of style showing stigma. G. Capsule. H. Seed. (A, photo by Lianna 

Jarecki; B-H, from Proctor 47242).


straight, white. A genus of about 540 species ranging from the southeastern United States to South 

America and the West Indies. Named in honor of Swedish physician and botanist Elias Erici Tillandz 

(1640-1693). 

LECTOTYPE: Tillandsia utriculata L., designated by Britton & Millspaugh, Bahamas Fl. 64. 1920. 

Key to the species of Tillandsia 

1. Plants evidently caulescent, the stems branching; leaves to 8 cm long but usually less ……………. 2 

2. Leaves polystichous; stamens longer than the petals …….....................……….. 12. T. tenuifolia 

2. Leaves distichous, stamens much shorter than the petals ………...................................………. 3 

3. Plants forming long, pendent masses; stems much elongate, up to several meters long, the 

internodes elongate, exposed; inflorescence a single pseudolateral flower ... 13. T. usneoides 

3. Plants forming globose masses; stems up to 10 cm long, the internodes very short, covered by 

the leaf sheaths; inflorescence evidently terminal, 1- to 5- (usually 2-) flowered 

......................................................................................................................... 10. T. recurvata 

1. Plants acaulescent or with very short, unbranched stems; leaves usually more than 8 cm long …....... 4 

4. Leaf sheaths inflated, broadly ovate to suborbicular, much and abruptly contracted into the blades 

.........................................................................................................................................................5 

5. Leaves densely covered with coarse-spreading scales throughout, having a velvety 

appearance; sterile portion of scape very short, infolded by the leaf sheaths …. 9. T. pruinosa 

5. Leaves densely covered with fine, appressed scales throughout, the surface smooth (nonvelvety); 

sterile portion of scape elongated, covered by sterile bracteoles ………………. 6 

6. Leaves and sterile scape bracteoles straight, erect, shorter than the inflorescence 

…………......................................................................................…… 1. T. ariza-juliae 

6. Leaves and sterile scape bracteoles contorted, spreading, longer than the inflorescence 

......................................................................................................................3. T. bulbosa 

4. Leaf sheaths about as thick as the blades, deltate to ovate, gradually attenuating into the blade 

...................................................................................................................................................... 7 

7. Leaf blades linear-subulate, less than 5 mm wide at the base, filiform for about half their 

length or more ….....................................................................................................………. 8 

8. Plants usually forming dense clumps of several plants; leaf blades filiform almost 

throughout, to 2 mm wide but usually narrower; bracts less than 1.5 cm long; sepals to 

1.2 cm long; petals about 2 cm long ……………................…………….. 11. T. setacea 

8. Plants usually solitary or forming clumps of few plants; leaf blades attenuating and 

becoming filiform for about half their length, usually more than 2 mm wide; bracts more 

than 1.5 cm long; sepals about 1.4 cm long or longer; petals about 2.5 cm long or more 

........................................................................................................................................ 9 

9. Leaf sheaths densely covered with subappressed to subspreading scales; 

inflorescence 14-25 cm long, its axis elongated, densely and pinnately branched; 

spikes sub-erect , straight ……………...........................……….. 2. T. borinquensis 

9. Leaf sheaths densely covered with appressed scales, seemingly glabrous and 

smooth; inflorescence, 5-12 cm long, its axis compressed, subdigitately to densely 

digitately branched; spikes divergent-spreading, arching-recurved 

………………….............................................................................. 5. T. festucoides 

7. Leaf blades deltate-attenuate, more than 5 mm wide at the base, filiform only near the apex 

............................................................................................................................................... 10 

10. Leaf blades with whitish transversal stripes, twisting around the scape….. 6. T. flexuosa 

10. Leaf blades concolorous, usually spreading, not twisting around the scape ..........… 11 

11. Leaves papery, chartaceous or subcoriaceous, thin, soft, flexuous; base of the leaf 

sheaths green, concolorous with the blades …………….............… 15. T. variabilis 

11. Leaves coriaceous, thick, stiff; base of the leaf sheaths dark brown or paleferruginous, 

contrasting with the blades. …………...................................…….. 12

218 


12. Inflorescence compound; more or less laxly branched, primary fertile bracteoles not overlapping, 

exposing the internodes of the inflorescence ……….............................................................……… 13 

13. Floral bracts densely imbricate, 2-2.7 cm long, hiding the internodes of the spikes 

….........................................................................................................………… 7. T. lineatispica 

13. Floral bracts not imbricate, 1-1.8 cm long, exposing the internodes of the spikes 

…….............................................................................................................…… 14. T. utriculata 

12. Inflorescence compound or simple; if compound, digitately to subdigitately or densely pinnately 

branched, the sheaths of the primary fertile bracteoles at least slightly overlapping and hiding the 

internodes of the inflorescence ……...............................................................................………….. 14 

14. Floral bracts 3-5 × 1.5-2.5 cm ……............................................................……. 4. T. fasciculata 

14. Floral bracts 1.5-2 × 0.8-1.2 cm …........................................………………… 8. T. polystachia 

1. Tillandsia ariza-juliae L. B. Sm. & Jiménez, 

Phytologia 6: 433. 1959. Type: Dominican 

Republic; La Vega. Ariza Julia s. n. 


Epiphytic, acaulescent herb, usually forming 

clumps. Leaves densely fasciculate, densely 

covered with fine appressed-cinereous scales; 

sheaths conspicuous, suborbicular, abruptly 

contracted into the blades, 3-4 cm long, inflated, 

forming a dense ovoid pseudobulb; blades 

involute, attenuate, erect, nearly straight, somewhat 

rigid, shorter than the inflorescence, 6-11 cm × 2- 

4 mm. Scape erect, hidden by sterile bracteoles; 

sterile bracteoles imbricate, subfoliaceous, with 

lanceolate to ovate sheaths infolding the scape and 

blades like the leaf blades. Inflorescence simple or 

few-branched, equaling to much longer than the 

leaves; primary bracteoles to 3 cm long, deltateovate, 

attenuate, chartaceous; spicate branches 

distichously-flowered, complanate, acute, 8-10 

cm long, appressed-lepidote; bracts erect, ovate to 

ovate-lanceolate, acute, 2-3 cm long, chartaceous, 

nerved, pinkish, densely lepidote on the outside. 

Flowers sessile or subsessile; sepals oblong, acute, 

slightly apiculate, ca. 2 cm long, densely lepidote 

on the outside; petals tubular-involute, erect, 

bluish purple; stamens exserted. Capsule 

cylindrical, about 3 cm long. 

General distribution: Dominican Republic 

and Puerto Rico. 

Distribution in Puerto Rico: Moist forest 

about 700 m. Apparently very rare, if not 

overlooked for its similarity with Tillandsia 

bulbosa Hook. This species is known in Puerto 

Rico from only two specimens collected in 

Maricao. 


Maricao: Maricao Insular Forest, Winters s.n. 

(US); Cantero s. n. (UPR). 

2. Tillandsia borinquensis Cedeño-Mald. & 

Proctor, Harvard Pap. Bot. 4: 113. 1999. 

Type: Puerto Rico; Yauco. R. García 3547 

(holotype: MAPR; isotypes: UPR, UPRRP). 

Tillandsia festucoides sensu Britton & P. Wilson 

(1923) and sensu Liogier & Martorell (1982), 

non Brongniart ex Mez, 1896. 

Epiphytic, acaulescent herb, sometimes 

forming small clusters. Leaves numerous, forming 

a dense fasciculate rosette, about 30-60 cm long; 

sheaths subtriangular-ovate, conspicuous, gradually 

narrowing into the blade, dark brown, densely 

subappressed to subspreading lepidote; blades 

linear-subulate proximally, filiform-attenuate 

distally, infolded or involute, the lower parts also 

dark brown and subappressed-lepidote. Scape 

erect or ascending, the axis 2-4 mm thick; sterile 

bracteoles erect, densely imbricate, chartaceous, 

covered with subappressed scales, green, 

sometimes purplish tinged, the lower ones 

subfoliaceous, the upper ones filiform-laminate. 

Inflorescence twice-branched, generally dense at 

apex, more open below, about 14-25 cm long, from 

almost equaling to much exceeding the leaves; 

primary fertile bracteoles like the upper sterile 

bracteoles but the distal ones without the filiform 

apex, their sheaths about as equal or slightly 

shorter than the axillary branches; branches 

densely distichous, mostly 4- to 6-flowered, about 

3-5 cm long, the lower subspreading, separate 

from each other and exposing the internodes, the 

upper densely clustered; bracts imbricate, ovateoblong, 

acute, glabrous to densely lepidote on the 

outside, coriaceous or subcoriaceous, carinate, 

1.8-2.2 cm long, exceeding the sepals, at least 

three times or more the length of the internodes of


the branches, green, sometimes purplish tinged. 

Flowers sessile to subsessile; sepals lanceolate to 

ovate-oblong, acute, 1.4-1.6 cm long, 

subcoriaceous, glabrous, carinate, connate 

proximally for 6-8 mm; petals 2.5-3.5 cm long, 

purple, stamens and pistil exserted. Capsule 

cylindrical-ellipsoid, short-pointed at apex, 2.4- 

2.8 cm long. 


Rico. 

Distribution in Puerto Rico: From dry scrub 

forest to wet forest, collected from 30-600 m. 

Relatively common. Recorded from Adjuntas, 

Aibonito, Arecibo, Ciales, Florida, Maricao, 

Morovis, Utuado and Yauco. 


Adjuntas: E of Los Rábanos, Rt. 518 near jct with 

Rt. 135, Ackerman & Campbell 2668 (US). Ciales: 

along trail Camino de la Ceiba towards Quebrada 

del Pozo Azul, Acevedo-Rdgz. & Vicens 11845 

(US). Utuado: 10 mi N from Adjuntas along road 

to Arecibo, Grant & Rundell 93-02292 (US). 

3. Tillandsia bulbosa Hook., Exot. Fl. 3: t. 173. 

1825; Platystachys bulbosa (Hook.) Beer, 

Fam. Bromel. 83. 1857. Lectotype: Trinidad. 

Hook., Exot. Fl. 3: t. 173. 1825, designated by 

L. B. Smith & Downs, Fl. Neotrop. Monogr. 

14: 994. 1977. 

Epiphytic, acaulescent herb, variable in size, 

usually forming clumps. Leaves several, densely 

fasciculate, finely appressed cinereous-lepidote; 

sheaths conspicuous, suborbicular, 3-5 cm broad, 

abruptly contracted into the blades, inflated, 

imbricate, forming a dense ovoid pseudobulb; 

blades involute, subterete, attenuate, usually 

contorted and spreading, 7-30 cm × 2-4 mm, 

usually exceeding the inflorescence. Scape erect 

or ascending, straight or slightly curved; sterile 

bracteoles imbricate, subfoliaceous, their blades 

exceeding the inflorescence. Inflorescence erect 

or curved, sometimes pendulous, spicate, simple 

or digitate, green to red; primary bracteoles of the 

inflorescence densely lepidote-cinereous on the 

outside, the lower one subfoliaceous, the sheaths 

ovate, the blades usually longer than the spikes, the 

upper ones subfoliaceous to ovate, acute and 

usually shorter than the spikes. Spikes digitate, 

spreading, lanceolate, acute, complanate, 2-8 cm 

long; rachis slender, lepidote; bracts erect to 

subspreading, conduplicate, carinate, usually red, 

ovate-lanceolate, acute, 1-2 cm long. Flowers 

sessile or subsessile; sepals oblong, apiculate, 

about 12 mm long; petals linear, acute, 2.5-4 cm 

long, blue to bluish purple; stamens exserted. Fruit 

a cylindrical capsule, 3-4 cm long, pointed at the 

apex. 


Belize, Honduras, Nicaragua, Costa Rica, Panama, 

Greater Antilles, Guadeloupe, Martinique, 

Trinidad, Tobago, Colombia, Venezuela, Guyana, 

Surinam, French Guyana, Ecuador, and Brazil. 

Distribution in Puerto Rico: Thickets, slopes, 

ridges and woods, from semi-xeric scrub forest to 

moist forest, from about 100-800 m. Uncommon, 

recorded from: Arecibo, Desecheo Is., Maricao, 

Mayagüez, and San Germán. 


Maricao: Camp. Santana Road, Maricao Insular 

Forest, Schubert et al. 480 (US). 

4. Tillandsia fasciculata Sw., Prodr. 56. 1788. 

Type: Jamaica. Swartz s. n. (lectotype: S-Sw.; 

isolectotype: P), here designated. 

Mostly epiphytic, sometimes terrestrial or 

lithophytic, acaulescent herb, variable in size, 

usually forming clusters. Leaves numerous, 

fasciculate, forming crateriform rosettes, appressed 

lepidote-cinereous on both surfaces, mostly 

ascending, straight to arching, 30-70 cm long; 

sheaths conspicuous, ovate, 3-5 cm broad, dark 

brown; blades narrowly triangular-attenuate, 

about 2-3 cm wide at the base, carinate, involute. 

Scape erect, stout, 20-45 cm long; sterile 

bracteoles subfoliaceous, erect, imbricate, 

coriaceous, lepidote-cinereous, shortening toward 

the apex. Inflorescence variable, simple or 

compound, with spicate branches; primary 

bracteoles ovate, acuminate to long-acuminate, 

coriaceous, shorter than the spikes, red or reddish; 

spikes subdigitate, compressed, erect to spreading, 

sessile or subsessile, usually densely-flowered, 

ellipsoid to cylindrical, acute, 5-17 cm long; bracts 

erect, imbricate, ovate-elliptic, acute, slightly 

uncinate, mostly 3-5 cm long, rigid, subcoriaceous, 

carinate at least toward the apex, lepidote or 

glabrous, green or greenish yellow. Flowers 

sessile to subsessile, erect; sepals lanceolate, 

acute, carinate, shorter than the bracts; petals 

linear, erect, tubular, 4-6 cm long, white to purple;

220 


stamens and pistil exserted. Capsule cylindricalellipsoid, 

3-4 cm long. 

General distribution: Florida, Mexico, 

Guatemala, Honduras, Salvador, Nicaragua, Costa 

Rica, Panama, Bahamas, Greater Antilles, St. 

Martín, Saba, Guadeloupe, La Desirade, Dominica, 

St. Lucia, St. Vincent, the Grenadines, Trinidad & 

Tobago, Venezuela, Colombia, Surinam, French 

Guiana, and Brazil. 


Islands: Variable, from mangrove swamps and 

coastal areas to montane moist forest above 800 m. 

One of the most common bromeliads, especially in 

the limestone areas. Recorded from Adjuntas, 

Arecibo, Aibonito, Bayamón, Camuy, Carolina, 

Cayey, Ciales, Dorado, Guánica, Maricao, 

Mayagüez, Morovis, Ponce, Río Grande, Salinas, 

San Juan, Vega Alta, Vega Baja, Yauco, and 

Vieques; Guana Island, St. John, and St. Thomas. 

Note. This species shows great variation over 

its range. Mez (in Engler, Das Pflanzenr. IV. 32 

(Heft 100): 467-468. 1935) described several 

varieties of this species, of which two have been 

attributed to Puerto Rico: T. fasciculata var. 

fasciculata and T. fasciculata var. venosispica 

Mez (Britton & Wilson, 1923; Smith & Downs, 

1977; Acevedo-Rodríguez et al., 1996). After 

examining all the Puerto Rican material at MAPR, 

UPR, SJ, and US, I have concluded that our 

specimens of T. fasciculata cannot be definitely 

assigned to any of the two varieties. Although in 

some cases weak distinctions could be made, most 

of our material falls intermediate between the 

putative varieties. Therefore, I have decided not to 

recognize the varietal distinctions proposed by 

Mez. A similar situation was described for 

Hispaniola (Zanoni et al, 1986). 


Adjuntas: 10 mi N of Adjuntas, along road to 

Arecibo, Grant & Rundell 93-02291 (US). 

Arecibo: 5 km S of Biáfara, Acevedo-Rdgz. & 

Siaca 11702 (US); Hato Arriba, , N.L. Britton & 

Cowell 1993 (US). Bayamón: Candelaria, Goll et 

al. 278 (US). Carolina: Santa Barbara, Goll et al. 

194 (US). Ciales: along trail Camino de la Ceiba 

towards Quebrada del Pozo Azul, Acevedo-Rdgz. 

& Vicens 11890 (US). Guánica: Mt. Ensenada, 

N.L. Britton & Shafer 1935 (US). Maricao: 

Sintenis 473 (US). Mayagüez: vicinity of 

Mayagüez; Ponce: NE of Ponce, Heller s.n. (US). 

Salinas: Bo. Lapa, E peak, Las Tetas de Cayey, 

Proctor & Díaz 42239 (US). San Juan: Martín 

Peña, Stevenson 1229 (US). Vega Baja: Vega 

Baja, Heller & Heller 637 (US). Vieques: 

Lighthouse Peninsula, Shafer 2804 (US). ST. JOHN: 

Cruz Bay Quarter; Battery Gut, near waterfall, 

Acevedo-Rdgz. & Siaca 4169 (NY, US). 

5. Tillandsia festucoides Brongn. ex Mez in C. de 


to be sought among several syntypes*. 

Epiphytic, acaulescent herb, sometimes 

forming clusters. Leaves numerous, densely 

fasciculate in rosettes, densely and minutely 

appressed-lepidote throughout, 15-40 cm long; 

sheaths narrowly deltate-ovate, conspicuous, dark 

brown; blades subulate-filiform. Scape erect or 

ascending, slender, 1-2 mm wide; sterile 

bracteoles erect, densely imbricate, subfoliaceous, 

sheathing, with long filiform blades reaching the 

inflorescence, chartaceous, pinkish or often bright 

red. Inflorescence bipinnate, sub-digitate to 

densely digitate, 5-12 (-17) cm long; primary 

fertile bracteoles like the upper scape bracteoles, 

their sheaths much shorter than the axillary 

branches; branches, few to several, archingrecurved 

especially after anthesis, linear in 

outline, 3-9 cm long; bracts erect, slightly 

imbricate, lanceolate-ovate, 1.5-2 cm long, 

exceeding the sepals, nerved, pale-appressedlepidote, 

green, pinkish or red. Flowers subsessile; 

sepals lanceolate, acute or obtuse, 1.4-1.7 cm long, 

coriaceous, glabrous or sparsely lepidote, carinate 

and connate proximally for 6-8 mm; petals erect, 

tubular, linear, about 3 cm long, purple; stamens 

and pistil exserted. Capsule narrowly cylindrical, 

acute, 2.5-3 cm long. 


Belize, Honduras, Nicaragua, Costa Rica, and 

Greater Antilles. 

Distribution in Puerto Rico: From dry scrub 

forest to wet forest, collected from 30-600 m. 

Apparently rare, known only from Maricao and 

the Susúa Forest Reserve in Yauco. Specimens of 

T. borinquensis, a rather more widespread and 

common species, have been attributed to this 

species. 


Maricao: Sargent 454 (US). 

Note: * The lectotypification of this name by 

L.B.Smith & Downs, (Fl. Neotrop. Monogr. 14:


914. 1977) is erroneous because the chosen 

lectotype is not part of the original material. 

6. Tillandsia flexuosa Sw., Prodr. 56. 1788. 

Lectotype: Jacquin, Select. Stirp. Amer. Hist. 

t. 63. 1763 (as a new name for T. tenuifolia 

sensu Jacquin 1763, non L., 1762), designated 

by L.B. Smith & Downs, Fl. Neotrp. Mongr. 

14: 977. 1977. 

Tillandsia aloifolia Hook., Exot. Fl. 3: t. 205. 

1826; Vriesea aloifolia (Hook.) Beer, Fam. 

Bromel. 95. 1856. Type: Trinidad. De Schack 

s.n. (K, photo at US). 

Epiphytic acaulescent herb, usually solitary. 

Leaves several, fasciculate, subcoriaceous, curved, 

usually spirally twisted around the scape and 

forming a dense sub-bulbous rosette, densely 

appressed-lepidote, usually with broad white 

transverse stripes, ovate to ovate-lanceolate, longattenuate, 

8-30 (-50) cm long, the outer usually 

shorter and sometimes scale-like; sheaths usually 

undifferentiated from the blade or gradually 

tapering into it. Scape ascending, usually curved 

near the base; sterile bracteoles erect, sheathing, 

the lowest ones foliaceous, the upper ones linearattenuate 

to lanceolate, obtuse or acute, slightly 

apiculate, shorter than the internodes, lepidote. 

Inflorescence simple or laxly twice-branched, 

much exceeding the leaves; primary bracteoles 

like the scape bracteoles, shorter than the sterile 

bases of the branches; the branches few, 

ascending, to 40 cm long, laxly-flowered, the 

rachis slender and flexuous; bracts, spreading, 

elliptic-lanceolate, acute, 2-3 cm long, about as 

long as or slightly shorter than the sepals, 

chartaceous, prominently nerved. Flowers with 

pedicels 3-5 mm long; sepals linear-elliptic, 

obtuse, 2-2.5 cm × 5-7 mm; petals linear, acute, to 

4 cm long, white, rose or purple; stamens exserted. 

Capsule slender, cylindrical, acuminate, to 7 cm 

long. Clusters of plantlets commonly developing 

from seeds germinating in the dried inflorescences. 

General distribution: Florida, Panama, 

Bahamas, Greater Antilles, St. Vincent, the 

Grenadines, Grenada, Colombia, Venezuela, 

Trinidad, Tobago, Guyana, Surinam, and French 

Guiana. 

Distribution in Puerto Rico: Woods in coastal 

moist forest. Rare. Recorded from San Juan and 

recent collections from Laguna Tortuguero in 

Manatí and Susúa forest. 


San Juan: Santurce, Heller & Heller 1394 (NY, 

US). 

7. Tillandsia × lineatispica Mez in C. de 


St. John, U.S. Virgin Is., Eggers 3104 

(holotype: B, photo at US). 

Terrestrial or lithophytic acaulescent herb, 

usually in small clusters. Leaves numerous, 

fasciculate, ascending, the lower spreading, 

forming dense rosettes, densely lepidote 

throughout, about 60-95 cm long; sheaths 

conspicuous, dark brown, ovate-elongate, 5-8 cm 

wide; blades narrowly triangular, attenuate to a 

filiform recurved apex, involute. Scape erect, 

stout, to about 1.5 m long, glabrous; sterile 

bracteoles erect, coriaceous, imbricate, sheathing, 

red, the lower subfoliaceous, shortening toward 

the apex of the scape, the upper ones ovatelanceolate, 

long- to short-acuminate; inflorescence 

a polystichous panicle, 1- to 3-branched, 

exceeding the leaves; fertile bracteoles ovate to 

ovate-lanceolate, acute to acuminate, red, 2.5-5 

cm long; the branches ascending; spikes erect, 

short-stipitate, strict, linear, 10-40 × ca. 1 cm, 

densely-flowered; bracts, erect, imbricate, 

coriaceous, carinate, nerved, ovate, acute, 2-2.7 

cm long, shorter than the sepals, purplish red with 

purple margins, the apices slightly spreading or 

turned outwards. Flowers subsessile, erect; sepals 

narrowly elliptic-lanceolate, acute, 2.5-3 cm long; 

petals white, linear, tubular, acute, to 3.5 cm long; 

stamens and pistil exserted. A sterile hybrid not 

producing fruit (Acevedo-Rdgz. 1996). 




Islands: Rare, known only from a few populations 

in the islands of Vieques, Culebra and St John. 

Note: T. lineatispica seems to be a hybrid 

between T. utriculata and T. fasciculata (Acevedo- 

Rdgz., 1996). 


Culebra: on peak 0.6 km due South of Playa 

Resaca, G. R. Proctor 39984 (SJ). Vieques: Cerro 

Ventana (Mt. Pirata), Shafer 2979 (US). ST. JOHN: 

Eggers 3104 (photo at US-2). Fish Bay, hillside 

overlooking Bay, Mori & Woodbury 17074 (NY);

222 


Half Moon Beach at Sun Bay, Hansen et al. 9460 

(FTG); Adrian, 600’ Eggers 3104-B (photo at US); 

Cruz Bay Quarter, Battery Gut, Acevedo-Rdgz. & 

Siaca 4170 (F, MO, NY, UPR, US). 

8. Tillandsia polystachia (L.) L., Sp. Pl. ed. 2, 

410. 1762; Renealmia polystachia L., Sp. Pl. 

286. 1753. Lectotype: West Indies. A plate of 

the Boerhaave set of Plumier illustrations at 

the library of Rijksuniversiteit, Groningen, 

here designated*. 

Tillandsia angustifolia Sw., Prodr. 57. 1788. 

Type: Jamaica. Swartz s. n. (holotype: S-Sw.; 

isotype: BM, photo at GH, NY). 

Epiphytic or sometimes lithophytic, 

acaulescent, rather variable herb, solitary or in 

clusters. Leaves numerous, in a fasciculate rosette, 

densely pale-appressed-lepidote throughout, about 

20-65 cm long; sheaths conspicuous, ovateelliptic, 

ca. 5-9 × 2-6 cm, dark brown; blades 

deltate-attenuate, gradually tapering into a 

filiform apex, arching to recurving, the margins 

involute. Scape erect or ascending, the sterile base 

shorter than the leaves; scape bracteoles 

foliaceous, reducing toward the apex, with erect, 

densely imbricate sheaths and elongate spreadingrecurved 

blades, green, red or purplish. 

Inflorescence variable, usually subcylindric or 

narrowly fusiform and exceeding the leaves, rarely 

simple to more commonly congested and 

subdigitate, or with a polystichously and densely 

branched axis to 45 cm long with short internodes; 

primary fertile bracteoles longer or shorter than 

the branches, short- or long-acuminate, reduced 

towards apex of the inflorescence (more 

commonly the lower primary fertile bracteoles are 

subfoliaceous with sheaths infolding the bases of 

the branches and involute-filiform blades much 

longer than the branches, the upper ovatelanceolate, 

short- or long-acuminate, and shorter 

than the branches); branches erect, ascending or 

spreading, straight or curved, linear, about 3-14 

cm long, complanate, densely-flowered; bracts 

erect, appressed, rigid, imbricate, ovate-oblong, 

broadly acute or obtuse, mucronulate, 1.5-2 cm 

long, longer than the sepals, much longer than the 

internodes, coriaceous, smooth, glabrous or 

obscurely lepidote, carinate. Flowers subsessile, 

erect; sepals linear-elliptic to ovate-oblong, about 

1.2-1.5 cm long, acute or mostly obtuse, 

coriaceous, even, glabrous, carinate; petals erect, 

tubular, ligulate, about 3 cm long, blue or violet; 

stamens and pistil exserted. Capsule cylindricalellipsoid, 

pointed, 2.5-4 cm long. 

General distribution: Mexico to Bolivia. 

Distribution in Puerto Rico: Variable, from 

dry scrub forest at sea level to moist and wet 

forests about 800 m. Very common and 

widespread, probably in all municipalities; 

recorded from Aguadilla, Arecibo, Cayey, 

Coamo, Comerío, Fajardo, Florida, Jayuya, Loíza, 

Manatí, Maricao, Mayagüez, Mona Island, Ponce, 

Salinas, Utuado, Vega Alta, and Yauco. 

Common Names: Puerto Rico: Parásita, 

Piñón. 

Note. *This lectotypification supersedes a 

neotypification by L.B. Smith & Downs (1977) of 

a Plumier specimen at P (see note under Aechmea 

nudicaulis for an explanation). 



10740 (UPRRP, US). Cayey: 13 km N of Cayey, 

Underwood 364 (US). Maricao: Sargent 711 (US). 

Mayagüez: 10 mi. E of Mayagüez, Grant & 

Rundell 93-02289 (US). Utuado: vicinity of 

Utuado, N.L. Britton & Cowell 1019 (US). 

9. Tillandsia pruinosa Sw., Fl. Ind. Occid. 1: 594. 

1797. Type: Jamaica. Swartz s. n. (holotype: 

S-Sw., R-6148). 

Tillandsia breviscapa A. Rich. in Sagra, Hist. Fis. 

Cuba, Bot. 11: 265. 1850. Type: Cuba. Sagra 

s. n. (holotype: P, photo at GH, NY). 

Small, epiphytic acaulescent herb, usually 

forming clusters. Leaves several to numerous, 

about 10-14 cm long, densely fasciculate, densely 

cinereous-lepidote throughout with coarse 

spreading scales having a velvety appearance; 

sheaths very conspicuous, broadly elliptic-ovate 

to suborbicular, 2-4 × 1.5-2 cm, inflated, forming a 

dense and ovoid pseudobulb, much wider than the 

blades and abruptly contracted into them, the inner 

ones closely infolding the scape, dark brown; 

blades ascending or spreading, recurved or 

contorted, usually exceeding the inflorescence, 

involute, narrowly deltate, gradually attenuate 

from a 4-7 mm wide base to a filiform apex. Scape 

erect, the sterile base short, wholly infolded by leaf 

sheaths; sterile bracteoles foliaceous. Inflorescence 

simple or rarely digitate with 2-5 spikes; primary 

fertile bracteoles subfoliaceous, densely cinereous-


lepidote; spikes densely and distichously 5- to 15flowered, 

to 7 × 4 cm, but usually much smaller, 

ellipsoid-ovoid, complanate; bracts erect, ovate, 

acute, 2-2.6 cm long, much exceeding the sepals, 

subcoriaceous, pink at anthesis, densely lepidote. 

Flowers sessile; sepals broadly elliptic-ovate or 

oblong, obtuse, 1-1.8 cm long, sparsely lepidote or 

glabrous, subcoriaceous, carinate; petals linear, 

obtuse or acute, about 3 cm long, bluish violet; 

stamens exserted. Capsule cylindrical-ellipsoid, 

pointed, about 3-5.5 cm long. 


Guatemala, Honduras, Costa Rica, Greater 

Antilles, Colombia, Venezuela, Ecuador, and 

Brazil. 

Distribution in Puerto Rico: Slopes, ridges 

and river margins of scrubby semi-xeric to moist 

forest, about 130-450 m. Very rare; known only 

from Yauco. 


Yauco: Susúa Forest Reserve, trail from reserve 

office N along Río Loco, Axelrod & Chavez 4038 

(US). 

10. Tillandsia recurvata (L.) L., Sp. Pl. ed. 2, 410. 

1762; Renealmia recurvata L., Sp. Pl. 287. 

1753, excluding var. β. Lectotype: Jamaica. 

Sloane, Voy. Jamaica 1: t. 121, f. 1. 1707, 

designated by L. B. Smith, Proc. Amer. Acad. 

Arts Sci. 70 : 208. 1935. 

Tillandsia uniflora Kunth in Humb., Bonpl. & 

Kunth, Nov. Gen. & Sp. 1 [quarto ed.]: 290. 

1816. Type: Mexico; Guerrero. Humboldt & 

Bonpland s. n. (holotype: P). 

Tillandsia recurvata f. major André, Bromel. 

Andr. 65. 1889. Type: Puerto Rico; Mayagüez. 

Sintenis 265 (BM). 

Epiphytic, lithophytic or sometimes terrestrial 

caulescent herb, also growing on elevated utility 

wires and cables; forming dense, globose or matlike 

clumps. Stems simple or branched, curved, 

typically shorter than the leaves, 1-10 cm long; 

roots few, short, slender. Leaves distichous, 3-17 

cm long, densely lepidote; sheaths almost 

inconspicuous, thinly elliptic-ovate; blades usually 

strongly contorted and recurved, occasionally 

spreading or erect, filiform. Scape to about 13 cm 

long, punctate-lepidote throughout; scape 

bracteoles linear, 1 or sometimes 2 immediately 

below the inflorescence. Inflorescence a simple, 

dense, few-flowered raceme, erect to recurved; 

floral bracteoles lanceolate, 1-1.5 cm long, nerved, 

punctate-lepidote. Flowers 1-5, usually 2, 

subsessile; sepals linear-lanceolate, acute, shorter 

than the bracts; petals linear, narrow, bluish or 

purplish, 2-2.2 cm long, stamens included. 

Capsule cylindric-ellipsoid, acuminate, 2-2.5 cm 

long. Seeds with comose hairs, 2-2.5 cm long. 


(Arizona, Texas, and Florida), Mexico, Guatemala, 

Honduras, El Salvador, Nicaragua, Bahamas, 

Greater Antilles, Anguilla, St. Martín, St. 

Barthelemy, Antigua, Saba, St. Eustatius, St. Kitts, 

Nevis, Redonda, Montserrat, Guadeloupe, 

Martinique, St. Lucia, Colombia, Venezuela, 

Ecuador, Peru, Bolivia, Brazil, Chile, Paraguay, 

Uruguay, and Argentina. 

Distribution in Puerto Rico & the Virgin 

Islands: Variable, probably including all moisture 

regimens and elevations, although probably most 

common in drier areas of the Island. Of ample 

occurrence, probably in every municipality and 

forest reserve; St. Croix and St. John. 

Common Name: Puerto Rico: Nidos de 

gungulén. 



181 (SJ). Cabo Rojo: West of Cabo Rojo, Heller 

4427 (US). Ciales: Casa Blanca, Sargent 3047 

(US). Coamo: Sintenis 3023 (US). Desecheo 

Island, N.L. Britton et al. 1576 (US). Guánica: 

Reserva Forestal Guánica, Boom 9970, 10025 

(US). Guayama: Underwood 427 (US). Isabela: 

Guajataca Gorge, Acevedo-Rdgz. & Cedeño 

10198 (UPRRP, US). Mayagüez: University of 

Puerto Rico, Mayagüez Campus, Strong et al. 435 

(US). Mona Island: N.L. Britton et al. 1813 (US). 

Morovis: Bo. Río Grande, along Rd. 155 ca. 0.9 

km. by road N of River (Río Grande de Manatí), 

Proctor & Concepción 42319 (US). Ponce: Prey 

65 (US). Vieques: 1.5 km N of Punta Carenero, ca. 

6 km E of Camp García, Fosberg 57527 (US). ST. 

CROIX: Bassin, Ricksecker 272 (US). South shore 

near Fanny’s Fancy, Seligson 41 (US). Great 

Pond, Fosberg 53971 (US). St. La Grange, 

Raunkiaer s.n. (US). ST. JOHN: Coral Bay Quarter; 

trail to Fortsberg, Acevedo-Rdgz. 4096 (NY, US). 

11. Tillandsia setacea Sw., Fl. Ind. Occid. 1: 593. 

1797. Type: Jamaica. Swartz s. n. (holotype: 

S-Sw., R-6154, photo at GH).

224 


Tillandsia bromoides Mez, Repert. Spec. Nov. 

Regni Veg. 12: 419. 1913. Type: Dominican 

Republic; Barahona. Fuertes s. n. (holotype: 

B, destroyed). 

Epiphytic, acaulescent herb, usually forming 

clumps of several plants. Leaves numerous, 

densely fasciculate, ascending or spreading, 

straight, curved or arching, 15-30 cm long, usually 

exceeding the inflorescence but sometimes short, 

appressed-lepidote throughout; sheaths triangularovate, 

conspicuous, dark brown-ferruginous, 

often keeled; blades stiff, involute, filiform, to 2 

mm wide but usually less than 1 mm wide for most 

of their length. Scape erect, slender, sparsely 

lepidote; sterile bracteoles erect, densely paleappressed-lepidote, 

imbricate, with lanceolate 

sheaths infolding the scape, the blades filiform, 

chartaceous. Inflorescence erect, bipinnate or 

frequently simple and distichous-flowered, 

slenderly ellipsoid, open at least below, to 12 cm 

long but usually much less; axis slender, lepidote, 

somewhat zigzagging; primary bracteoles like the 

upper scape bracteoles, sub-erect , the sheath 

shorter than the axillary spike but the blade usually 

much exceeding it; spikes subsessile, erect to 

subspreading, very short and dense, few-flowered, 

complanate; bracts erect, concave, imbricate, 

ovate, acute or acuminate, slightly exceeding the 

sepals, subcoriaceous, densely pale-appressedlepidote. 

Flowers subsessile; sepals ellipticoblong, 

acute, 7-12 mm long, carinate; petals 

erect, tubular, linear, acute, dilated at the apex, 

about 2 cm long, violet; stamens and pistil 

exserted. Capsule narrowly cylindrical, about 2.5 

cm long. 


Guatemala, Greater Antilles, and Brazil. 

Distribution in Puerto Rico: Thickets, slopes, 

ridges and woods, from semi-xeric scrub forest to 

moist forest, from sea level to about 500 m. 

Relatively common. Recorded from: Adjuntas, 

Arecibo, Barranquitas, Camuy, Comerío, Hatillo, 

Juana Díaz, Mona Island, Ponce, Utuado, and 

Yauco. 


Arecibo: W. Sabana Hoyos, Acevedo-Rdgz. & 

Chinea 11664 (US). Juana Díaz: Peña de las 

Cuevas, near Juana Díaz, E.G. Britton & Marble 

2295 (US). Ponce: Río San Patricio, Sargent 3130 

(US). Utuado: Road from Utuado to Arecibo, 

Underwood 807 (US). 

12. Tillandsia tenuifolia L., Sp. Pl. 286. 1753. 

Lectotype: without locality. Royen s. n. (L), 

designated by L.B. Smith & Downs, Fl. 

Neotrop. Mongr. 14: 829. 1977. 

Tillandsia pulchella Hook., Exot. Fl. 2: t. 154. 

1825. Type: Trinidad. De Schack s. n. 

(holotype: K). 

Tillandsia pulchra Hook., Exot. Fl. 2: t. 154. 1825. 

Type: Cultivated from Trinidad. 

Tillandsia subulata Vell., Fl. Flumin. 133. 1829 

[“1825”]; Icon. 3: pl. 127. 1831 [“1827”]. 

Type: Brazil; Rio de Janeiro. Vellozo s. n. (not 

located). 

Epiphytic, sometimes lithophytic or terrestrial 

caulescent decumbent herb. The stems slender, 

elongate, branching and spreading, to 25 cm long, 

proximally covered with remnants of old sheaths. 

Leaves densely polystichous along the stem, 4-8 

cm long, densely and minutely appressed-lepidote 

throughout; sheaths small, 4-7 mm wide, deltateovate 

to linear, abruptly constricted into the blade 

of the lower leaves, barely distinct from the blades 

of the upper leaves; blades narrowly deltateattenuate, 

2-4 mm wide at the base, tapering 

gradually and becoming filiform, spreading to 

mostly ascending or erect, straight or curved, the 

margins involute. Scape erect or ascending, 

slender, about 3-7 cm long; scape bracteoles erect, 

slightly imbricate, with ovate-lanceolate sheaths 

8-10 mm long, abruptly constricted into filiformcaudate 

blades 2-3 cm long, membranaceous, 

pinkish. Inflorescence short, its axis about 1-1.5 

cm long, equaling or exceeding the leaves, simple, 

densely polystichously-flowered, ovoid; bracts 

like the scape bracteoles but smaller and with the 

blades much reduced, their laminar sheath slightly 

exceeding the sepals, punctulate-lepidote, nerved. 

Flowers 4-10, sessile, erect; sepals lanceolate, 

acute, glabrous, about 6-10 mm long, 

membranaceous, carinate; petals blue, white, or 

rose, about 20 mm long, the claw linear, the blade 

oblong, broadly obtuse; stamens and style 

exserted. Capsule cylindrical, apiculate, about 1.5- 

2 mm long. 


Martinique, St. Lucia, Venezuela, Guyana, French 

Guiana, Surinam, Bolivia, Brazil, Paraguay, and 

Argentina.



forest, above 500 m. Recorded from Adjuntas, 

Ciales, Cidra, Coamo, Maricao, Salinas, and 

Yauco. 


Coamo: Las Piedras Chiquitas, Liogier & Proctor 

36184 (US).Yauco: Bo Río Prieto, upper W slope 

of Monte Membrillo, Axelrod & Bayman 7364 

(US). 

13. Tillandsia usneoides (L.) L., Sp. Pl. ed. 2, 411. 

1762; Renealmia usneoides L., Sp. Pl. 287. 

1753; Dendropogon usneoides (L.) Raf., Fl. 

Tellur. 4: 25. 1838. Lectotype: “Americas.” 

Collector unknown (BM-Cliff. 558543) 

designated by Gouda in Görts-van Rijn (ed.), 

Fl. Guianas 189: 68, 1987. 

Epiphytic, caulescent, herb, typically forming 

pendent masses to a meter or more in length. Stems 

branched, sympodial, flexuous, recurved, less than 

1 mm in diam., internodes 3-6 cm long, the 

branches bearing 2-3 leaves. Leaves distichous, 2- 

5 cm long, densely cinereous-lepidote; sheaths 

slightly dilated, elliptic-lanceolate, tightly 

enveloping the base of the internodes, 4-8 mm 

long; blades filiform, less than 1 mm diam. No 

reproductive specimens of this species were 

located from Puerto Rico or the Virgin Islands. 

The flowers and fruits have been reported by 

others as follows. Inflorescence reduced to a single 

pseudolateral flower; bract ovate, apiculate or 

caudate, densely lepidote, shorter than the sepals. 

Flowers subsessile; sepals narrowly ovate, acute, 

to 7 mm long, thin, strongly nerved, glabrous, 

short-connate, light green; petals narrow, acute or 

obtuse, 9-11 mm long, pale green or yellowish 

green; stamens included, exceeding the pistil. 

Capsule 2-3 cm long, cylindrical, short-beaked. 

General distribution: From the southeastern 

United States to Chile and Uruguay, including the 

West Indies. 


Islands: From rather open, dry, scrubby forest to 

thickets, woodlands and edges of moist forest; 

ranging from near sea level to about 600 m. 

Arecibo, Cabo Rojo, Coamo, Guánica, Guayama, 

Lajas, Mayagüez, Ponce, Salinas, and Yauco; St. 

Croix. 

Common name: Puerto Rico: Barbas de úcar. 


Arecibo: Bo. Río Arriba, Axelrod et al. 9020 

(UPRRP). Cabo Rojo: Salinas de Cabo Rojo, 

Sintenis 564 (US). Coamo: vicinity of Coamo 

Springs, N.L. Britton & Cowell 1330 (US). 

Guánica: Roadside in and around Guánica, Grant 

& Rundell 93-02269 (US). Guayama: Aguirre, 

Underwood 365 (US). Lajas: La Parguera, near 

km. 2, Schubert & Winters 363 (US). Ponce: 

Between Ponce & Coamo, Heller & Heller 513 

(US). Yauco: Stevenson & Fink 3956 (US). ST. 

CROIX: Caledonia Gut, NW corner of Island, 

Fosberg 55384 (US). Jakobsberg, Lang’s Peak, 

Fosberg 54153 (US). Signal Hill, Ricksecker 267 

(US). 

14. Tillandsia utriculata L., Sp. Pl. 286. 1753. 

Neotype: Jamaica. W. Till 9014 (WU), 

designated by Till in C.E. Jarvis et al. (ed.), 

Regnum Veg. 127 : 94. 1993. 

Tillandsia ramosa Sweet, Hort. Brit. 425. 1826, 

nom. nudum. 

Tillandsia sintenisii Baker, J. Bot. 26: 12. 1888. 

Type: Puerto Rico; Cayey. Sintenis 2134 

(holotype: K; isotypes: GH, US!). 

Figs. 43. G-I; 65. D 

Small to large herbs, 0.5-2 m tall, epiphytic or 

sometimes lithophytic, acaulescent or with short 

stems, solitary or in clusters. Leaves numerous, 

densely fasciculate, rosulate, 20-100 cm long, 

densely pale-appressed-lepidote throughout, 

coriaceous; sheaths conspicuous, their bases dark 

brown, broadly ovate-elliptic, 2-9 cm wide; blades 

narrowly deltate, gradually tapering or longattenuate, 

the inner sub-erect , the outer spreading 

and usually recurving. Scape erect, to 1.5 m long, 

glabrous; sterile bracteoles erect, sheathing, the 

lower subfoliaceous, the upper bladeless, ovate, 

acute, shortening toward the apex. Inflorescence 

variable in length, exceeding the leaves, once to 

thrice-branched, very large and open, glabrous; 

fertile bracteoles like the upper scape bracteoles, 

about 1.5-3.5 cm long, shorter than the sterile base 

of the branches, often dark violet; branches 

ascending; spikes linear in outline, curved, 8-35 

cm long, their bases with several sterile, imbricate 

bracteoles, laxly- and distichously-flowered; 

floral bracts erect, infolding the base of the 

flowers, not imbricate, equaling or shorter than the 

internodes, ovate-lanceolate, acute or obtuse, 1- 

1.8 cm long, shorter than the sepals, nerved,

226 


Fig. 43. A-C. Pitcairnia angustifolia var. angustifolia. A. Inflorescence, and leaf tips. B. Flower. C. Pistil and petal. D-F. Catopsis 

floribunda. D. Habit. E. Infructescence. F. Capsule. G-I. Tillandsia utriculata. G. Habit. H. Inflorescence branch. I. Tepal, 

flower, and detail of stigma. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78.


subcoriaceous, green with a broad, membranaceous, 

often dark purple margin, ecarinate. Flowers erect, 

with pedicels 2-4 mm long; sepals elliptic-oblong 

or oblanceolate, obtuse, 1.4-1.8 cm long, 

subcoriaceous, nerved, with membranaceous 

margins; petals erect, tubular, linear, acute, 3-4 cm 

long, white, creamy or yellowish; stamens and 

pistil exserted. Capsule narrowly cylindricalellipsoid, 

pointed, 3-5 cm long; seed with long 

apical comose hairs. 


States (Georgia and Florida), Mexico to Costa 

Rica, Bahamas, Greater Antilles, Lesser Antilles, 

Trinidad, Tobago, and Venezuela. 


Islands: Common from coastal, dry scrub forests 

to moist and wet forests at about 800 m. Aguadilla, 

Bayamón, Cayey, Coamo, Culebra, Desecheo, 

Fajardo, Icacos, Maricao, Mona Island, Río 

Grande, Sabana Grande, Salinas, San Juan, 

Vieques, and Yauco; Anegada, Guana Island, St. 

Croix, St. John, and St. Thomas. 


Aguadilla: Bo. Caimital Bajo Aibonito, Acevedo- 

Rdgz. et al. 13444 (US). Bayamón: Stevenson 

3926 (US). Cayey: Sintenis 2134 (US). Coamo: 

Coamo Springs, Underwood 588, 589 (US). 

Culebra: N.L. Britton & Wheeler 187 (US). 

Desecheo Island, N.L. Britton et al. 1587 (US). 

Maricao: Sargent 662 (US). Mona Island, Otero & 

Chardon 914 (US). San Juan: Martín Peña, 

Stevenson 3700 (US). Vieques: Cayo Puerto Real, 

Shafer 2769 (US). Yauco: Susúa, Liogier 9903 

(US). ANEGADA: Settlement, N.L. Britton & 

Fishlock 1024 (US); East side of Flamingo Pond, 

Acevedo-Rdgz. 10979 (US). ST. CROIX: Crequis, 

Ricksecker 440 (US). Maroon Ridge, seaward 

slope ca. 1 km SE of Hamms Bluff, Fosberg 60875 

(US). Salt Pond, Ricksecker s.n. (US). Sandy Point 

Peninsula, Fleming 251 (US). Sandy Point, 

Gardner 251 (US). ST. JOHN: Coral Bay Quarter; 

Road to Bordeaux, Acevedo-Rdgz. & Siaca 5065 

(K, MO, NY, UPRRP, US). ST. THOMAS: Bremer’s 

Bay, N.L. Britton & E.G. Britton 255 (US). Univ. 

of the V.I. campus, Fosberg 59410 (US). 

15. Tillandsia variabilis Schltdl., Linnaea 18: 

418. 1844. Type: Mexico; Veracruz. Schiede 

s. n., Jan. 1829, (lectotype: HAL 45636.1; 

isolectotype: HAL 45636. 2), designated by 

W. Weber, Feddes Repert. 95: 597-598. 1984. 

Tillandsia valenzuelana A. Rich in Sagra, Hist. 

Fis. Cuba, Bot. 11: 267. 1850. Type: Cuba. 

Sagra s. n. (holotype: P). 

Tillandsia sublaxa Baker, J. Bot. 25: 280. 1887. 

Type: Jamaica; Westmoreland. Purdie s. n. 


Tillandsia domingensis Mez, Repert. Spec. Nov. 

Regni Veg. 16: 73. 1919. Type: Dominican 

Republic. Tuerckheim 3250 (holotype: B, 

n.v., photo at NY). 

Epiphytic or sometimes lithophytic, 

acaulescent herb, solitary or in clusters. Leaves 

numerous, fascicled, rosulate, 10-45 cm long, the 

outer ones commonly reduced to scale-like 

sheaths, cinereous-lepidote throughout, 

occasionally becoming glabrous above, green, red, 

or reddish green; sheaths conspicuous, ovate, 1.5- 

5 cm long; blades ascending or spreading, 

involute, 7-20 mm wide at the base, narrowly 

triangular-attenuate to an almost filiform apex. 

Scape slender, glabrous, 7-30 cm long, erect or 

ascending, straight or curved; sterile bracteoles 

erect, sheathing, imbricate, the lower ones 

subfoliaceous with spreading almost caudate 

blades, the upper ones ovate-lanceolate, 1.8-2 cm 

long, acute, apiculate or short-acuminate, densely 

lepidote, pinkish or reddish green. Inflorescence 

erect or ascending, simple or pinnately compound, 

shorter or mostly longer than the leaves; primary 

fertile bracteoles like the upper sterile bracteoles; 

spikes few, oblong, ellipsoid or lanceolate, acute, 

complanate, slightly flexuous, laxly- or mostly 

densely-flowered, 5-20 cm long; bracts erect or 

slightly divergent and exposing the rachis, about 

two to three times as long as the internodes, 

elliptic-oblong or ovate-lanceolate, obtuse, acute 

or apiculate, 1.5-2 cm × 6-8 mm, slightly or much 

exceeding the sepals, subcoriaceous, nerved, 

glabrescent, pink or red. Flowers subsessile; 

sepals elliptic-oblong, obtuse or acute, carinate, 

1.3-1.6 cm long; petals linear, acute, 2.5-3 cm 

long, blue or bluish violet; stamens exserted. 

Capsule ellipsoid, acute, 2-3 cm long; seeds with 

white coma about 1.5 cm long. 


Mexico to Colombia, Venezuela, Ecuador, 

Bolivia, and the Greater Antilles. 

Distribution in Puerto Rico: Moist areas; 

thickets, woodlands and moist forest, from near 

sea level to about 830 m. Very common. Recorded

228 


from Adjuntas, Aibonito, Arecibo, Barceloneta, 

Barranquitas, Bayamón, Cabo Rojo, Ciales, Cidra, 

Dorado, Florida, Isabela, Juana Díaz, Quebradillas, 

Maricao, Salinas, Utuado, Vieques, and Yauco. 


Arecibo: Cambalache Forest Reserve, along 

Piletas trail, Axelrod & Ackerman 5801 (US). 

Bayamón: vicinity of Bayamón, Liogier 10720 

(US). Ciales, Bo. Toro Negro, Rt. 147, km 27.7, 

Axelrod & Ackerman 3550 (US). Juana Díaz: Peña 

de las Cuevas, vicinity of Juana Díaz, E.G. Britton 

& Marble 2297 (US). Salinas: Bo. Lapa, E peak, 

Las Tetas de Cayey, Proctor & Díaz 42238 (US). 

Utuado: Cayuco, Sintenis 6414 (US). Vieques: 

Cerro Ventana, Shafer 2869 (US). 


Liogier & Martorell (1982) cite Tillandsia 

juncea (Ruiz & Pav.) Poir. as occurring near 

Adjuntas, Puerto Rico. However, no specimens 

have been seen to confirm this record. 

Doubtful Species 

The name Tillandsia ramosa Bello (Anales 

Soc. Esp. Hist. Nat. 12: 121. 1883) can not be 

applied with certainty to any taxon in Puerto Rico 

because the type was destroyed. Its protolog 

provides a lenghty description which might allow 

it to be associated with a species in Puerto Rico, 

however, since the name has never been in used, it 

would be peferable to propose it for rejection. 

9. VRIESEA 

Vriesea Lindl., Edward’s Bot. Reg. 29: 1843 [“Vriesia”], nom. et orth. conserv. 

Acaulescent usually epiphytic herbs. Leaves in a rosette, polystichous, entire; blades predominantly 

ligulate and inconspicuously lepidote; leaf scales centrally symmetric. Scape usually conspicuous. 

Inflorescence various, usually of distichous-flowered spikes, rarely of one or more polystichous-flowered 

spikes, the distichous flowers becoming secund in many species; bracts generally conspicuous. Flowers 

bisexual, mostly short-pedicellate; sepals convolute, free, or nearly so, symmetric or subsymmetric; petals 

free or connate into a tube, much shorter than the sepals, with 2 ligulate appendages at the base, firm and 

erect or soon flaccid and drooping; stamens included or exserted; ovary superior, or nearly so; stigma 

lobulate, papillose; ovules many, usually caudate. Fruit capsular. Seeds fusiform with a basal coma and an 

apical awn. A genus of ca. 193 species from tropical and subtropical South America and the West Indies. 

Named in honor of Dutch botanist and physician Willem Hendrick de Vriese (1806-1862), of Leyden, The 

Netherlands. 

TYPE: Vriesea psittacina (Hook.) Lindl. (≡ Tillandsia psittacina Hook.). 

1. Vriesea macrostachya (Bello) Mez in C. de 

Candolle, Monogr. Phan. 9: 601. 1896; 

Caraguata macrostachya Bello, Anales Soc. 

Esp. Hist. Nat. 12: 122. 1883; Neovriesea 

macrostachya (Bello) Britton in Britton & P. 


Neotype: Puerto Rico; Orocovis. Axelrod 

6935, (US; isoneotypes, B, MARY, NY, SEL, 

UPRRP), designated by J.R. Grant, Nordic J. 

Bot. 17(2): 158. 1997. 

Large, epiphytic or sometimes terrestrial or 

lithophytic herb. Leaves several, fascicled, 

rosulate, entire, mostly erect, 50-100 cm long, 

glabrous above, lepidote below; sheaths 

conspicuous, dark brown; blades linear-attenuate, 

5-8 cm wide, rounded to acute, apiculate, 

concolorous. Scape very stout, erect or slightly 

curved at the base, to 2 m long; sterile bracteoles 

ovate-elliptic, acute, densely imbricate. 

Inflorescence simple, erect, distichous, 45-80 cm 

long, many-flowered, the axis mostly over 1 cm 

thick; bracts ovate, navicular, 3-6 cm long, 

coriaceous, acute or obtuse, yellowish. Flowers 

spreading, not secund, nocturnal; pedicels stout, 1- 

1.5 cm long; sepals imbricate, coriaceous, ovateelliptic 

or obovate, obtuse, 2.5-4.6 × ca. 1.5 cm; 

petals imbricate, linear-obovate, obtuse or acute, 

4-4.5 × 1.5-2 cm, yellowish white, with a pair of 

linear appendages at the base; stamens included. 

Capsule ellipsoid-ovoid, pointed, 4-6 cm long; 

seeds with whitish coma. 

General distribution: Cuba, Hispaniola, and 

Puerto Rico.



forests, about 500 m and above. Recorded from 

Adjuntas, Barranquitas, Luquillo, Maricao, 

Naguabo, Orocovis, Ponce, Río Grande, San 

Germán, Salinas, and Utuado. 

Common names: Puerto Rico: Parásita, 

Pirigallo. 


Barranquitas: Bo. Barrancas, upper NE slopes of 

10. WERAUHIA 

Werauhia J. R. Grant, Trop. Subtrop. Pflanzenwelt 91: 28. 1995. 

Mt. Torrecilla, Proctor 42300 (SJ). Maricao: Río 

Maricao, N.L. Britton & Cowell 4187 (US). 

Naguabo: Sierra de Luquillo, Caribbean Natl. 

Forest, Proctor 47980 (SJ). Orocovis: Bo. Ala de 

la Piedra, Toro Negro Forest Reserve, Axelrod 

6935 (US). Río Grande: Sierra de Luquillo, 

Sintenis 2792 (US). Utuado: 7 mi N from Adjuntas 

along road to Arecibo, Grant & Rundell 93-02294 

(US). 

Epiphytic or terrestrial, acaulescent or caulescent herbs. Leaves numerous, rosulate or in a spiral 

along the stem, inconspicuously lepidote; sheaths conspicuous to non-differentiated from blades; blades 

ligulate to linear-triangular, entire. Inflorescence scapose, distichous, simple or compound. Scape 

bracteoles, primary bracteoles, and floral bracts usually dull, green or brown, or rarely red. Scape 

bracteoles transitional in form between the leaves and primary bracteoles of inflorescence; bracts 

conspicuous, fleshy and secund. Flowers bisexual, pedicellate to sessile; sepals free, ecarinate, imbricate, 

coriaceous; petals free, imbricate in bud, elliptical, with dactyloid appendages at the base, fleshy, dull in 

color, white, greenish, yellowish or light orange; petal appendages two, dactyloid; stamens shorter than 

petals, asymmetric; anthers forming a hood over the pistil; ovary superior to inferior; stigma lobes 

cupulate, without papillae. Fruit a septicidal capsule; seeds numerous, with basal coma. A genus of about 

66 species from tropical America and the West Indies. Named in honor of German botanist Dr. Werner 

Rauh, born in 1913. 

TYPE: Werauhia gladioflora (H. Wendl.) J. R. Grant (≡ Tillandsia gladioflora H. Wendl.). 

Key to the species of Werauhia. 

1. Leaves usually with transverse dark reddish bands; inflorescence simple or laxly branched, with suberect 

, few-flowered secund branches …………..........................................................…. 2. W. ringens 

1. Leaves never with transverse reddish bands; inflorescence with very short branches, wholly aborted, 

each subtending a pair of flowers at the apex …………….............................................…………… 2 

2. Sterile bracteoles erect, involute; inflorescence axis about 5-10 (-12) cm long, somewhat dense 

and congested at the apex; primary fertile bracteoles 3 to 6, 4-6 cm long, the apical ones slightly 

shorter than the basal ones, sub-erect…………………………................……….. 1. W. proctorii 

2. Sterile bracteoles subspreading to spreading, foliaceous; inflorescence axis 15 to 30 cm long, 

laxly- to densely-flowered; primary fertile bracteoles usually more than 10, 5-15 cm long, the 

apical ones much shorter than the basal ones, spreading ……………….………… 3. W. sintenisii 

1. Werauhia proctorii Cedeño-Mald., Harvard 

Pap. Bot. 4: 115. 1999. Type: Puerto Rico; 

Sierra de Luquillo. Proctor 50061, (holotype: 

US; isotypes: IJ, JBSD, NY, UPR). 

Epiphytic and terrestrial, acaulescent herb. 

Leaves numerous, densely and polystichously 

fascicled, forming crateriform rosettes, erect to 

sub-erect , to 30 cm long but usually shorter, 

typically red in exposed situations, but sometimes 

tinged with green especially near the base, or 

mostly green when in shade, dark brown lepidotepunctulate 

throughout on both surfaces; sheaths 

elliptic, mostly wider than the blades; blades 

linear-attenuate, 2-3 cm at the middle, acute to 

mostly acuminate. Scape erect, exceeding the 

leaves, axis yellowish green, glabrous, punctulate, 

somewhat verrucose above the nodes; sterile

230 


bracteoles erect and involute, exceeding the nodes, 

elliptic-ovate, acute to short-acuminate, lepidotepunctulate 

on both surfaces, chartaceous, bright 

red with yellowish green bases. Inflorescence fewflowered, 

polystichously branched, mostly dense 

and congested at the apex of the scape, ellipticcylindrical 

in general outline, 6-12 cm long; 

primary fertile bracteoles like the scape bracteoles, 

erect to sub-erect , 3.5-6 cm long, equaling or 

mostly exceeding the flowers; branches almost 

wholly aborted, punctate-verrucose, subtending a 

pair of flowers; bracts broadly ovate, obtuse to 

acute, sometimes bifid, 1.3-1.5 cm × 6-8 mm, 

infolding the bases of the flowers, lepidotepunctulate 

on the outside, prominently nerved and 

punctulate within. Flowers erect; pedicels stout, 3- 

5 mm long, punctulate-verrucose; sepals 

oblanceolate-spatulate to oblong, obtuse, 1.8-2.5 

cm × 4-6 mm, punctate, rigid-coriaceous, 

sometimes bifid; petals ligulate, sub-elliptic, acute 

to obtuse, 2.6-3 cm long, yellow; stamens 

included. Capsule ellipsoid-cylindrical, short 

pointed at apex, 3-3.5 cm long. 


Rico. 

Distribution in Puerto Rico: Wet montane 

forest to cloud forest above 900 m. Recorded from 

Cayey, Jayuya, Naguabo, Río Grande, and San 

Lorenzo. 


Cayey: Sierra de Cayey, Carite Forest Reserve, 

Proctor et al. 47101 (US). Jayuya: Cordillera 

Central, Toro Negro State Forest, Proctor & 

Rivera 47079 (US). Naguabo: Sierra de Luquillo, 

Caribbean Natl. Forest, Bo. Río Blanco, Proctor 

47980 (US). Río Grande: Caribbean Natl. Forest, 

Roca el Yunque, Proctor 42077 (US). 

2. Werauhia ringens (Griseb.) J. R. Grant, Trop. 

Subtrop. Pflanzenwelt 91: 35. 1995; Tillandsia 

ringens Griseb., Cat. Pl. Cub. 255. 1866; 

Vriesea ringens (Griseb.) Harms, Notizbl. 

Bot. Gart. Berlin-Dahlem 10: 801. 1829. 

Type: Cuba; Oriente. Wright 1518, in part 

(holotype: GOET; isotypes: GH, MO). 

Vriesea guadalupensis sensu Bold., Fl. Dutch W. 

Ind. Is. 34. 1909, non (Baker) Mez 

Epiphytic or occasionally lithophytic herb, 

variable in size. Leaves numerous, ascending, to 

32 (-90) cm long, punctate-lepidote beneath; 

sheaths ovate-elliptic, mostly indistinct; blades 

linear-attenuate, acute or acuminate, 3-6 cm wide, 

usually reddish green, red-banded. Scape erect, 

stout; sterile bracteoles erect, densely imbricate, 

sheathing, elliptic-lanceolate, acute to acuminate, 

pale green or tinged with purple. Inflorescence 

erect, simple or less often laxly branched, to 50 cm 

long; primary fertile bracteoles like the scape 

bracteoles but thinner, much shorter than the 

axillary branches; branches sub-erect , with dense 

to open, secund, few-flowered clusters, bearing 

several imbricate sterile bracteoles at the base; 

rachis usually slender, verrucose just below the 

nodes; bracts broadly ovate, acute-acuminate, 3- 

6.5 cm long, infolding the flowers, exceeding the 

sepals (at least in lower flowers), straight, carinate 

toward apex, green or brownish. Flowers erect, to 

8 cm long, glabrous; sepals elliptic, acuminate, 

2.5-3.5 cm long, subcoriaceous, lustrous; petals 

white, yellow or greenish, sometimes finely redtinged, 

recurved. Capsule to 4 × 1 cm, ellipsoidcylindrical, 

apiculate; seeds with entire, minute 

coma. 

General distribution: Costa Rica, Panama, 

Jamaica, Hispaniola, Puerto Rico, Lesser Antilles 

and Trinidad. 

Distribution in Puerto Rico: Wet mountain 

forest, ca. 500-700 m. Known from three 

specimens collected in the Luquillo Mts. 

Note: This is a very rare species in Puerto Rico 

that has only been vouchered with three specimens 

from the Sierra de Luquillo. Although undoubtedly 

identifiable, the specimens are relatively 

depauperate. The description provided is mostly 

based on Smith and Downs (1977), and Howard 

(1979). 


Luquillo: Río Mameyes, in the Luquillo forests in 

the mountains, Roehrs s.n. (US); Luquillo Mts., 

Woodbury s.n. (UPRRP). Río Grande: El Verde, 

Woodbury s. n. (UPRRP). 

3. Werauhia sintenisii (Baker) J. R. Grant, Trop. 

Subtrop. Pflanzenwelt 91: 48. 1995; Caraguata 

sintenisii Baker, Handb. Bromel. 145, 1889; 

as [“sintenesii”]; Guzmania sintenisii (Baker) 

Mez in C. de Candolle, Monogr. Phan., 9: 920. 

1896; Thecophyllum sintenisii (Baker) Mez, 

Bull. Herb. Boissier, Ser. 2, 3: 131. 1903; 

Vriesea sintenisii (Baker) L. B. Sm. & 

Pittendr., J. Wash. Acad. Sci. 43: 403. 1953.


Type: Puerto Rico; Sierra de Luquillo. 

Sintenis 1589 (holotype: K, US-fragment). 

Fig. 65. E 

Epiphytic or sometimes terrestrial, acaulescent 

herb. Leaves numerous forming a dense crateriform 

rosette, erect or sub-erect , mostly ca. 30 cm long, 

but sometimes up to 56 cm long, chartaceous, 

green, reddish or dark red; sheaths elliptic-ovate, 

from very distinct to slightly wider than the blade, 

densely punctate-lepidote; blades linear-attenuate, 

acute or obtuse, apiculate, ca. 2-4 cm wide, flat, 

glabrous above, very obscurely punctulatelepidote 

beneath. Scape erect or ascending, 

slender, ca. 5 mm thick, glabrous; sterile 

bracteoles erect, sheathing, imbricate, 

subfoliaceous, exceeding the internodes, deep red, 

the upper ones sometimes subspreading. 

Inflorescence erect, exceeding the leaves, 

polystichously branched, laxly- to denselyflowered, 

15-30 cm long; primary fertile 

bracteoles sub-erect to mostly spreading, ellipticovate, 

acute or acuminate, exceeding the flowers, 

deep red; branches almost wholly aborted, each 

subtending a pair of flowers; bracts ovate to 

broadly ovate or almost suborbicular, acute or 

obtuse, 8-15 mm long, punctulate, infolding the 

bases of the flowers. Flowers subsessile or with 

pedicels to ca. 5 mm long; sepals elliptic-oblong, 

acute to mostly obtuse, 1.5-2.5 cm long, 

coriaceous, glabrous; petals yellowish, linear, 

obtuse, 3 cm long; stamens included. Capsule 

ellipsoid-subcylindric, pointed, ca. 3 cm long. 

General distribution: Greater Antilles. 

Distribution in Puerto Rico: Moist, wet 

montane and cloud forests above 700 m. Recorded 

from Adjuntas, Barranquitas, Cayey, Ceiba, 

Jayuya, Las Piedras, Maricao, Naguabo, Patillas, 

Ponce, Río Grande, and Utuado. 


Adjuntas: Arroyo de los Corchos, between 

Adjuntas and Jayuya, N.L. Britton & Cowell 5292 

(US). Barranquitas: Bo. Barrancas, Upper NE 

slopes of Monte Torrecilla, Proctor & Concepción 

42300 (US). Jayuya: Toro Negro State Forest, 

upper W summit of ridge of Cerro de Punta, 

Proctor & Rivera 47080 (US). Las Piedras: El 

Toro, Agric. Exp. Station, Woodbury s.n. (US). 

Maricao: Buena Vista Camp, Maricao Insular 

Forest, Schubert et al. 481 (US). Patillas: Carite 

Forest Reserve, Axelrod & Weisel 11129 (US). Río 

Grande: Luquillo Mountains, Caribbean National 

Forest, El Yunque area, Acevedo-Rdgz. 10518 

(US). 


Billbergia pyramidalis (Sims) Lindl. is grown 

as an ornamental in Puerto Rico. Although listed in 

Liogier and Martorell (1982), there are no records 

of this species indicating its naturalized status in 

Puerto Rico. Dyckia sulphurea K. Koch was noted 

by Britton & P. Wilson (1923) as cultivated and 

established in St. Thomas. Several species of 

Neoregelia are cultivated extensively as 

ornamentals in Puerto Rico. Neoregelia carolinae 

(Beer) L. B. Sm. and N. marmorata (Baker) L. B. 

Sm., both native to Brazil, were collected by 

Proctor (43730-SJ and 43729-SJ respectively) as 

naturalized. However, further visits to the site 

show that the species did not persist. 

Family 30. XYRIDACEAE Yellow-eyed grass Family 

Xyridaceae C. Agardh, Aphor. Bot. 158. 1823, nom. conserv. 

by G. R. Proctor & M. T. Strong 

Annual or perennial, more or less rosulate, scapose terrestrial herbs of marshy or hydric soils; 

rhizomes absent but base of plant often somewhat hard or bulbous. Culms terete, ribbed, smooth, or 

sometimes wing-margined. Leaves basal or several lower cauline, tufted or closely alternate and 

distichous or equitant, the blades linear and flat to terete-filiform, sheathing at base, glabrous or minutely 

pubescent. Inflorescence usually terminal on a naked culm, consisting of a compact, bracteate, cone-like 

spike, the imbricate bracts spirally arranged. Flowers bisexual, zygomorphic, solitary in the axils of the 

bracts. Calyx of 3 sepals, the outer two chaffy, boat-like, and persistent, the inner one membranous and

232 


covering the flower in bud. Corolla ephemeral, of 3 subequal clawed petals, the outer expanded portion 

usually yellow (rarely white), the claws concealed by the bracts. Stamens 3, adnate to the petals by their 

filament bases, the anthers divided into 2 thecae by a broad connective; staminodes 3, alternating with the 

petals, long-stalked, bifid at apex. Ovary superior, 1-locular to incompletely 3-locular, the placentation 

marginal or parietal, style terminal, hollow, 3-branched towards the apex; stigmas 3, glandular-hairy. Fruit 

a capsule with valvate or irregular dehiscence; seeds usually numerous, small, striate or striate-reticulate, 

ovoid, ellipsoid, or fusiform. A pantropical to warm-temperate family of 4 genera and nearly 300 species. 

Only the genus Xyris occurs in Puerto Rico. There is no modern monographic treatment of the family, only 

a few regional surveys and inclusion in local or regional floristic studies. 

TYPE: Xyris L. 

Reference: Kral, R. 1966. Xyris (Xyridaceae) of the continental United States and Canada. Sida 2 (3): 

177-260. 

Xyris L., Sp. Pl. 42. 1753. 

1. XYRIS 

Characters as given for the family, but does not include special features that apply to genera other than 

Xyris. A genus of about 275 species with its center of distribution in the Americas, particularly tropical and 

warm temperate regions. United States, southeastern Mexico, Central America, West Indies, South 

America, Asia, Africa, and South Pacific Islands. 

TYPE: Xyris indica L. 

Key to the species of Xyris 

1. Leaves narrowly linear to filiform, their bases lustrous deep brown or tan; keel of lateral sepals firm, 

ragged or ascending-fimbriate toward the apex ………...............................................… 2. X. elliottii 

1. Leaves linear, their bases pale, stramineous, or pinkish; keel of lateral sepals scarious, lacerate. ….. 2 

2. Plant annual; culms solitary or 2-3 together; leaves 10-60 cm long, 5-10 mm wide; mature spikes 

7-15 (-25) mm long, obtuse ……………............................................................….. 3. X. jupicai 

2. Plant perennial; culms tufted; leaves 1-8 (-12) cm long, 1-4 mm wide; mature spikes 4-7 (-8) mm 

long, acute …………………….......................................................................……. 1. X. curtissii 

1. Xyris curtissii Malme, Ark. Bot. 13: 24. 1913; 

Xyris serotina var. curtissii (Malme) Kral, 

Rhodora 62: 310. 1960; Xyris difformis var. 

curtissii (Malme) Kral, Sida 2: 255. 1966; 

Xyris neglecta Small, Bull. Torrey Bot. Club 

21: 30. 1894, non Nilsson, 1892.Type: 

Florida; Duval County. Curtiss 4136 

(holotype: NY). 

Caespitose perennial. Culms 7-26 (-30) cm 

tall, linear-filiform, angulate-terete, not much 

widened distally, several-ribbed, ribs equally 

prominent, somewhat scabrous, the sheath at base 

rich deep red-brown, lustrous. Leaves spreadingascending 

in broad fans, 1-8 (-12) cm ×1-4 mm, 

linear-sword-shaped, dull green, often papillate, 

margins somewhat scabrous; sheaths reddish or 

purplish, finely rugulose-papillate. Mature spikes 

ovoid, 4-7 (-8) mm long; lateral sepals included, 

slightly curved, 5-7 mm long, the keel brown, 

scarious, lacerate; petals with obtriangular blades, 

4 mm long, unfolding in the morning; staminodes 

bearded. Seeds translucent, ovoid to ellipsoid, 0.5 

mm long, finely lined longitudinally with small 

papillae. 

General distribution: United States 

(southeastern and gulf coastal plains), Central 

America (Belize), and Puerto Rico. 

Distribution in Puerto Rico: This species is 

confined to moist white silica sand habitats near 

sea level along the north side of Puerto Rico from 

Dorado west to the area of Laguna Tortuguero. 

Recorded from Dorado, Manatí, Vega Alta, and 

Vega Baja.



Vega Baja: Algarrobo, Tortuguero Lagoon 

Natural Reserve, Vélez-Gavilán et al. 374 

(MAPR). 

2. Xyris elliottii Chap., Fl. South. U. S. 500. 1860. 

Type: Florida; Franklin County. Chapman s. 

n. (lectotype: NY; isolectotype: NY), 

designated by Kral, Sida 2: 224. 1966. 

Xyris brevifolia sensu Elliott, Sketch Bot. S. 

Carolina 1: 52. 1816, non Michaux, 1803. 

Perennial (and spreading) by means of pale, 

fleshy, elongate, lateral buds, often forming tufts 

with brownish bases. Culms usually 35-60 cm tall 

or more, often slightly twisted. Leaves 8-35 cm × 

1-2 (-2.5) mm, linear to narrowly linear (rarely 

filiform), flat or slightly twisted, the margins pale 

and minutely tuberculate toward the base, the apex 

sharply acute or acuminate, the base abruptly 

dilated, lustrous, the surfaces above the equitant 

base green or reddish and smooth. Mature 

flowering spikes ovoid to broadly ellipsoid, 6-15 

mm long, acute at apex, bearing several closely 

imbricate bracts; fertile bracts 5-6 mm long, 

obovate, the surfaces pale to dark brown; lateral 

sepals scarious, included or slightly exserted, the 

keels ragged or ascending-fimbriate toward the 

apex; blades of petals obovate, yellow, ca. 5 mm 

long, opening in the morning. Seeds 0.5-0.6 mm 

long, ellipsoid, translucent, with ca. 12 distinct 

straight longitudinal lines. 


States, Cuba, and Puerto Rico. 

Distribution in Puerto Rico: In moist white 

sands at sea level. Recorded from Dorado, Manatí, 

and Vega Alta. 


Dorado: Stevenson 854 (US); Sintenis 6844 (US). 

Manatí: Laguna Tortuguero, track at E end of 

mogotes (haystack hills) to S shore of lake, 

Axelrod & Sastre 5241 (US). Vega Alta: Bo. 

Sabana, silica-sand area NE of Regadera, Proctor 

45810 (US). 

3. Xyris jupicai Rich., Actes Soc. Hist. Nat. Paris 

1: 106. 1792. Type: French Guiana. LeBlond, 

s. n. (holotype: P). 

Xyris communis Kunth, Enum. Pl. 4: 12. 1843. 

Syntypes: French Guiana. Desfontaines s.n. 

(B, destroyed); Venezuela. Otto s.n. (B, 

destroyed). 

Xyris caroliniana sensu authors, non Walter, 

1788. 

Short-lived perennial, solitary or in small 

tufts, dying after one year or persisting by means of 

small bulbous offshoots. Culms 20-70 cm tall or 

more, terete and many ridged toward base, 

flattened and 1- or 2-edged above. Leaves 10-60 

cm × 5-10 mm, linear, ascending, often yellowgreen 

and somewhat lustrous long; sheath of the 

culm shorter than the principal leaves, somewhat 

loose toward the oblique apex, their bases usually 

stramineous, rarely pinkish. Mature spikes ovoid, 

ellipsoid, or oblong, 7-15 (-25) mm long, with 

rather many loosely imbricate bracts; fertile bracts 

obovate to oval, 5-7 mm long, the outer surface 

pale to dark brown; lateral sepals scarious, 

included, about the same length as the bracts, 

linear and slightly curvate, lacerate; petals with 

yellow cuneate blades ca. 3 mm long, opening in 

the morning. Seeds broadly ellipsoid, 4-5 mm 

long, with numerous faint longitudinal lines. 

General distribution: Southeastern coastal 

plain of the United States from New Jersey to 

Florida and west to Texas, Mexico and Central 

America, Greater Antilles, Trinidad, and South 

America. 

Distribution in Puerto Rico: Occurs near sea 

level especially in moist white sand but also in 

other soils near the north coast of Puerto Rico, and 

also in wet ditches and open banks in the Luquillo 

Mountains at 600-1000 m. Recorded from 

Bayamón, Carolina, Cataño, Manatí, Río Grande, 

San Juan, Vega Alta, and Vega Baja. 


Bayamón: Sintenis 934 (US). Carolina: Sabana 

Abajo, vicinity of Carolina, N.L. Britton & Brown 

5717 (US). Cataño: Heller & Heller 1379 (US). 

Las Piedras/Río Grande: Caribbean Natl. Forest, 

trail to El Toro Peak, Axelrod & Herscoviz 5161 

(US). Manatí: Laguna Tortuguero, N.L. Britton et 

al. 3842 (US); Bo. Tierras Nuevas Saliente, 

Proctor & Díaz 42162 (US); Proctor & Thomas 

43812 (US). Río Grande: El Yunque, Fosberg 

520, 44200, 44201, 44202 (US); Sargent 520 

(US). San Juan: Santurce, Heller & Heller 586 

(US); Martín Peña, Stevenson 497 (US). Vega 

Alta: Bo. Sabana, silica-sand area NE of Regadera, 

Proctor 45809 (US).

234 


Family 31. JUNCACEAE Rush Family 

Juncaceae Juss., Gen. Pl. 43. 1789, nom. conserv. 

by P. Lewis 

Grass-like herbs (one species woody) with mostly short, erect or horizontal rhizomes. Leaves basal, 

tufted, linear or filiform, cylindrical or flat, sheathing at the base. Flowers single or in panicles, corymbs, 

or heads, usually small, bisexual or unisexual and dioecious, actinomorphic. Perianth segments 6 in two 

series, or rarely 3 segments only, mostly glumaceous. Stamens (male part of flower), 6 or 3, opposite to 

the perianth-segments; anthers 2-locular, basifixed, opening lengthwise; pollen in groups of 4. Ovary 

superior, 1-locular or divided by 3 septa or 3-locular; styles 1 or 3; stigmas 3; ovules ascending or parietal. 

Fruit a loculicidally dehiscent capsule. Seeds sometimes tailed with a small straight central embryo. Eight 

genera with ca. 300 species, generally distributed but more numerous in temperate and cold regions. One 

genus, Juncus occurs in Puerto Rico. 

TYPE: Juncus L. Sp Pl. 325. 1753. 

Reference: Balslev, H. 1996. Juncaceae, Fl. Neotrop. Monogr. 68: 1-167. 

Juncus L., Sp. Pl. 325. 1753. 

1. JUNCUS 

Annual or perennial, rhizomatous, glabrous herbs. Culms usually erect, rarely procumbent or 

ascending. Leaves alternate, cataphyllous (i. e., with reduced blades) and/or foliar at the base of the culm; 

sheaths open, usually conspicuously auriculate at junction with the blade; blades linear, varying in cross 

section: flat with raised margins and slightly channeled above, canaliculate, round to elliptic, or flattened. 

Inflorescence compound and often with elongate branches; involucral bracts decreasing in size from the 

base of the inflorescence upwards, the lowermost one often conspicuously different from the upper ones. 

Flowers bisexual, sometimes clasped by 2 bracteoles on the pedicel; tepals equal or subequal, lanceolate, 

entire, persistent, stramineous or dark brown, sometimes light green. Stamens 3 or 6, the filaments filiform 

or flat, sometimes widened at the base, the anthers linear or oblong, obtuse; style 3-branched, the stigmas 

filiform, twining, papillose; ovary sessile. Capsule 1-locular, 3-septate or 3-locular; seeds many, 

ellipsoidal, oblong, or ovoid, smooth, rugose or sometimes with a reticulate pattern, brown, yellowish, or 

dark brown. Distribution: Cosmopolitan, with ca. 220 species, the center of distribution in north temperate 

regions. 

LECTOTYPE: Juncus acutus L., designated by Coville in Britton & A. Brown, Ill. Fl. N. U.S. ed. 2, 1: 

465. 1913; Hitchock & M.L. Green, Prop. Brit. Bot. 147. 1930. 

1. Juncus tenuis Willd., Sp. Pl. 2: 214. 1799. 

Type: United States; “America boreali”, 

collector unknown (lectotype: B-W6888, 

sheet 1; isolectotypes: B-W6888, sheets 2, 3, 

and 4, HBG; photo at AAU; fragment: NY), 

designated by Balslev, Fl. Neotrop. Monogr. 

68: 79. 1996. 

Fig. 44. A-E 

Perennial, caespitose herb, 15-70 cm tall; 

rhizome densely branched, 1.5-2 mm diam. 

Culms erect, terete, smooth or longitudinally 

ridged, 0.8-1.5 mm diam. Cataphylls 0-3 per 

culm, inconspicuous, to 5 cm long, margin 

scarious, rudimentary blade acicular, to 15 mm 

long. Leaves all basal, erect, 1 /3 of to equaling the 

length of the culm; sheaths 1.5-9 cm long, with 

scarious margins terminating in 2 scarious or 

cartilaginous auricles: blade linear, flat or terete 

and canaliculate in cross section, 0.5-1.5 mm wide. 

Inflorescence terminal on the culm, occupying less 

than ¼ of the total plant height, compound-cymose, 

consisting of several unilateral cymes (drepania). 

Lower inflorescence bract resembling basal leaves, 

herbaceous, 2-20 cm long, distal ones progressively 

shorter, ultimate ones scarious, to 5 mm long. 

Tepals unequal, lanceolate, acute to acuminate, at 

first green with distinct scarious margin, turning


Fig. 44. A-E. Juncus tenuis. A. Habit. B. Inflorescence branch. C. Flower. D. Capsule. E. Seed. (from Proctor 50694).

236 


brown to stramineous, outer tepals 3.5-5 mm long, 

concave, inner tepals 3-4.5 mm long, flat. Stamens 

6, 1.3-1.8 mm long, the anthers linear to oblong, 

0.4-0.8 mm long. Capsule ellipsoidal, acute to 

obtuse, apiculate, 2.5-3.5 × 2-2.5 mm, light brown 

at maturity, 3-septate; seeds ellipsoidal, apiculate, 

often curved, 0.4-0.6 × 0.2-0.3 mm, smooth to 

slightly rugose, yellow-brown. 

General distribution: Widespread and 

common species in North America, introduced in 

the 19 th century to western Europe where it is now 

widespread and still extending its range. Scattered 

introductions are found in Hawaii, Japan, the 

Azores, and New Zealand. In the Neotropical 

region, this species is found throughout the 

highlands. 

Distribution in Puerto Rico: Occurs in several 

varieties of which two are found in Puerto Rico 

while none are found in the Virgin Islands. 

Key to the varieties of Juncus tenuis in 

Puerto Rico: 

1. Plants not over 60 cm tall; auricles 0.5-1.5 mm 

long, usually longer than wide; leaf blades flat 

or channeled, adaxial side with a band of 

hyaline cells occupying ¾ or more width of 

blade. …….........……. J. tenuis var. tenuis 1a 

1. Plants up to 90 cm tall; auricles up to 0.5 mm 

long, wider than long; leaf blades terete or 

channeled, when channeled, the central part of 

the x. s. is thicker than the edges, adaxial side 

with or without a narrow band of hyaline cells 

occupying less than 1 /3 width of blade 

………….......….. J. tenuis var. dichotomus 1b 

1a. Juncus tenuis Willd. var. tenuis 

(See above for type and general distribution). 

Distribution in Puerto Rico: Occurs in open 

disturbed ground at high elevations (900-1300 m), 

Family 32. CYPERACEAE Sedge family 

Cyperaceae Juss., Gen. Pl. 26. 1789, nom. conserv. 

by M. T. Strong and P. Acevedo-Rodríguez 

locally frequent. Recorded from Jayuya, Naguabo, 

Ponce, Río Grande, and Villalba. 


Jayuya: Cerro de Maravilla to Cerro de Punta, 

Liogier & Martorell 35893 (UPR, US); Cerro de 

Punta, Woodbury s.n. (US); vicinity of Cerro 

Maravilla, Liogier et al. 28233 (UPR, US). 

Naguabo: Caribbean National Forest, near Pico 

del Este, alongside road, Axelrod & Wen 9160 

(US). Ponce: Toro Negro Forest, Vivaldi 71-49 

(UPR). Río Grande: Sierra de Luquillo, Caribbean 

Natl. Forest, Summit of El Yunque, Proctor 45543 

(US). 

1b. Juncus tenuis var. dichotomus (Elliott) A. 

W. Wood, Class-book Bot. 726. 1861; Juncus 

dichotomus Elliott, Sketch Bot. S. Carolina 1: 

406. 1817. Type: United States; South 

Carolina or Georgia. Elliott s. n. (CHARL, not 

seen, photo at KANU). 

General distribution: Eastern North America 

from Massachusetts to Texas, Mexico, Central 

America and South America at elevations of up to 

2500 m, with a previous solitary outlying record 

from Jamaica; recorded here from Puerto Rico for 

the first time. 


from Naguabo. 


Naguabo: Sierra de Luquillo, Caribbean National 

Forest, vicinity of Pico del Este, ca. 1020 m, 

Proctor 47993 (US). 


Liogier & Martorell (1982) cite Juncus 

effusus L. as occurring in the Toro Negro area of 

Puerto Rico. However, no specimens have been 

seen to confirm this record nor has it been recently 

collected there. 

Perennials with rhizomes or stolons, or annuals, grass-like or rush-like herbs, of aquatic or terrestrial 

habitats; roots fibrous. Culms (stems) solitary or caespitose, triquetrous to trigonous or obtusely so,


rarely 4- or 5-angled, sometimes terete or flattened, solid or hollow, ribbed or entire, smooth or scabrous, 

glabrous or sometimes pubescent. Leaves well-developed or reduced to bladeless sheaths, borne at base 

or both basal and cauline; blades flattened, pleated, folded, inrolled, or cylindrical, linear to lanceolate, 

glabrous or sometimes pubescent, often antrorsely scabrous on margins and primary veins or occasionally 

smooth; sheaths closed, the inner band membranous with a concave to convex orifice, or sometimes 

entirely herbaceous with the summit frequently prolonged as a triangular or tongue-like contraligule; 

ligule absent or short, sometimes a narrow band of hairs. Inflorescences diverse, typically umbelliform, 

anthelate, corymbose, spicate, paniculate, racemose, or capitulate, sessile or branching, bearing 1 to 

many spikelets or reduced spikes; involucral bracts leaf-like, well-developed or reduced, or the lowest 

appearing as a continuation of the culm, rarely absent; spikelets 1- to many-flowered, often with 1 to 

several empty (sterile) scales at base. Flowers (florets) spirally imbricate or distichous, bisexual or 

unisexual (the plant then monoecious or rarely dioecious), borne singly, each from the axil of a spikelet 

scale; perianth, when present, of 3 to many, smooth or barbed, hypogynous bristles or scaly segments, 

usually persistent at base of mature achene; stamens 1-3, the anthers elliptic to linear, basifixed, bilocular, 

with theca parallel, longitudinally dehiscent, apiculate or appendaged at apex; ovary 2- or 3-carpellate, 

unilocular, the style 2- or 3-branched, the unbranched portion capillary, flattened and strap-like, or 

sometimes 3-angled. Fruit an achene, trigonous (sometimes dorsally compressed), biconvex, globose, 

cylindrical, ellipsoid, or obovoid, usually apiculate at apex, smooth, reticulate, puncticulate, papillose, 

verrucose, or transversely rugulose, the style-base deciduous or persistent at apex. A large cosmopolitan 

family of approximately 3,500 species in 114 genera. As an aid to using the keys and descriptions, 

features of the Cyperaceae and their terminology are given in Fig. 45. 

TYPE: Cyperus L. 

References: Adams, C. D. 1992. Cyperaceae. Fl. Trinidad and Tobago 3(5): 383-561. Clarke, C. B. 

1900. Cyperaceae. In: I. Urban, Symb. Antill. 2: 8-169. González-Más, A. 1964. Cyperaceae of Puerto 

Rico. Ph.D. dissertation, Louisiana State University, Baton Rouge. Kearns, D.M., Wm. W. Thomas, 

G.C. Tucker, R. Kral, K. Camelbeke, D.A. Simpson, A.A. Reznicek, M. Socorro González-Elizondo, 

M.T. Strong, and P. Goetghebeur. 1998. Cyperaceae. Pp. 486-663. In: J. A. Steyermark, P.E. Berry, & B. 

Holst (eds.). Flora of the Venezuelan Guayana. Missouri Bot. Gard. Press, St. Louis, Missouri. Steudel, 

E.G. 1855. Synopsis Plantarum Cyperacearum. J. B. Metzler, Stuttgart, Germany. 348 pp. Strong, M.T. 

1996. Cyperaceae. Pp. 478-492. In: P. Acevedo-Rdgz., Flora of St. John, U.S. Virgin Islands, Mem. New 

York Bot. Gard. 78. Tucker, G. C. 1987. The genera of Cyperaceae in the southeastern United States. J. 

Arnold Arbor. 68: 361-445. 


1. Inflorescence contracted without evident lateral branches or rays, unispiculate (a solitary spikelet at 

the summit of the culm) or a terminal or pseudolateral, solitary, glomerate or hemispherical capitate 

cluster of 2-many spikes or spikelets at the apex of the culm. ........................................... 2 

2. Spikelets laterally compressed or subcompressed; scales distinctly distichous, at least towards 

base of spikelet. ...................................................................................................................... 3 

3. Elongate leaf blades and involucral bracts wanting, the summit of the leaf sheaths merely 

apiculate or entire; inflorescence unispiculate; achene usually bearing persistent bristles at 

base............................................................................................................ 6. Eleocharis 

3. Elongate leaf blades and/or involucral bracts present; inflorescence multispiculate or 

unispiculate; achene without bristles at base. .................................................................. 4 

4. Lowest involucral bract shorter than the subtending inflorescence; unbranched portion 

of style fimbriate towards apex, expanded at base; spikelets 1 (-2); achene coarsely 

papillate ............................................................................... 1. Abildgaardia 

4. Lowest involucral bract longer than the subtending inflorescence; unbranched portion 

of style smooth, not expanded at base; spikelets 2-many; achene finely papillose, 

punctate, puncticulate, reticulate, or smooth. ........................................... 5

238 


5. Spikelets 1- to 2-flowered; rachilla short; scales conduplicate, conspicuously keeled 

................................................................................................ 9. Kyllinga 

5. Spikelets 2- to many-flowered; rachilla elongate; scales navicular...... 5. Cyperus 

2. Spikelets rounded in cross section; scales or bracts spirally imbricate. ................................... 6 

6. Leaf blades and involucral bracts wanting; inflorescence unispiculate.......... 6. Eleocharis 

6. Leaf blades and/or involucral bracts well-developed; involucral bracts present; inflorescence 

multispiculate or unispiculate. .................................................................. 7 

7. Summit of leaf sheath long-pilose; inflorescence unispiculate............... 2. Bulbostylis 

7. Summit of leaf sheath glabrous, or at most short-pubescent; inflorescence a cluster of 

2-many spikes or spikelets. .................................................................................. 8 

8. Culm leafy from base to apex, the sheaths closely overlapping; leaf blades stiff 

and strongly recurved; plants strictly of coastal beaches and dunes. 13. Remirea 

8. Culm leafy only at base or lower cauline, or the leaves reduced to bladeless sheaths; 

plants of diverse habitats, not strictly of coastal beaches and 

dunes....................................................................................................... 9 

9. Inflorescence a terminal, sessile head of 3-several, well-developed spikelets 

subtended by foliaceous involucral bracts which are whitened basally in 

several species; achenes biconvex, transversely rugose or rugulose, bearing 

the persistent base of the style at apex .............................14. Rhynchospora 

9. Inflorescence a pseudolateral head of spikelets or small spikes, overtopped by 

a single, erect involucral bract appearing as a continuation of the culm; achenes 

plano-convex or subterete, smooth or minutely papillate or punctate, merely 

apiculate or short-beaked, not bearing the persistent base of the style at apex. 

....................................................................................................... 10 

10. Spikelets reduced, 1-flowered, each subtended and hidden by an outer 

herbaceous, obovate to spatulate, scale-like bract; spikelet scales 1-2 

(often wanting), hyaline or membranous; achenes terete or subterete, 

ellipsoid, ellipsoid-obovoid, or narrowly subcylindrical, merely apiculate 

at apex, minutely papillate or punctate; bristles at base of achene 

wanting.............................................................. 11. Lipocarpha 

10. Spikelets well-developed, many-flowered, the capitula of spikelets 

subtended by an elongate bract appearing as a continuation of the culm; 

spikelet scales numerous, ovate, herbaceous; achenes plano-convex, 

smooth, short-beaked at apex; bristles 1-6, down-curved or retrorsely 

barbed, persistent at base......................... 15. Schoenoplectus 

1. Inflorescence open with evident lateral branches or rays, terminal or with 2-several lateral partial 

inflorescences from the upper leaf-like bracts, umbelliform, corymbose, paniculate, or racemose, 

rarely interruptedly fasciculate-spicate, the 2-many spikelets solitary or in fascicles, digitate or 

glomerate clusters, or spikes at branch or ray tips......................................................................... 11 

11. Summit of leaf sheath long-pilose; style base persistent as a bulbous tubercle at the apex of the 

achene ........................................................................................................... 2. Bulbostylis 

11. Summit of leaf sheath glabrous or at most short-pubescent; style base, if persistent at apex of 

achene, triangular or conical, not bulbous. ....................................................................... 12 

12. Achenes globose or globose-trigonous with white bony or crustaceous pericarp, often 

smooth and shiny or sometimes verrucose, reticulate, or tuberculate, the base often with a 

persistent 3-lobed hypogynium ......................................................................16. Scleria 

12. Achenes trigonous or biconvex, or a subterete fructification, with stramineous to dark 

brown, reddish brown, or brown-black pericarp, transversely rugulose, puncticulate, 

papillate, or smooth, the base without a hypogynium, either naked or bearing persistent 

bristles, bladed-bristles, or minute scales. ..................................................................... 13


13. Fruit an achene bearing bladed or both bladed and spinulose bristles at 

base............................................................................................................ 8. Fuirena 

13. Fruit an achene or fructification, naked, or bearing (1-) 6 spinulose or smooth bristles or minute 

ciliolate scales at base................................................................................................................. 14 

14. Spikelet scales distinctly distichous ....................................................................... 5. Cyperus 

14. Spikelet scales imbricate or obscurely distichous................................................................ 15 

15. Fruit an achene bearing persistent, barbed, spine-like or needle-like bristles at base.. 16 

16. Leaves reduced to bladeless sheaths; culms terete, soft, easily compressed, with internal 

air cavities; inflorescence pseudolateral, the lowest involucral bract erect and appearing 

like a continuation of the culm; achenes plano-convex....................... 15. Schoenoplectus 

16. Leaves with well-developed blades; culms trigonous or lenticular, rarely terete, firm 

or hardened, without internal air cavities; inflorescence terminal or a series of partial 

inflorescences from the upper leaf-like bracts; achenes biconvex or with 3 wingangles..................................................................................................................... 

17 

17. Culm lenticular in cross section; leaves distichous; leaf blades unifacial, lacking 

distinct midvein; spikelet scales dark purple-black or black; style 3-branched; 

achene with 3 wing-angles......................................... 12. Machaerina 

17. Culm trigonous or subterete; leaves spirally 3-ranked; leaf blades with distinct 

midvein; spikelet scales stramineous, brown, or reddish brown; style 2-branched; 

achene biconvex .....................................................14. Rhynchospora 

15. Fruit an achene or fructification without persistent bristles at base. ............................. 18 

18. Fruit a herbaceous prominently veined sac (perigynium), loosely or tightly enclosing 

an achene, bidentate at its apex..................................................... 3. Carex 

18. Fruit an achene or fructification, not surrounded by a perigynium, naked or with 3 

minute scales at base. ............................................................................................. 19 

19. Fruit a biconvex or sometimes obtusely trigonous achene bearing the persistent 

style base or merely apiculate or short-beaked at apex, transversely rugose or 

rugulose, cancellate, or warty.......................................................................... 20 

20. Unbranched portion of style often flattened and fimbriate or slender and 

papillate distally; style base disarticulating from the apiculate or short-beaked 

achene apex; achene surface often cancellate or warty, usually of vertical 

rows of isodiametric to horizontally rectangular 

cells........................................................................................ 7. Fimbristylis 

20. Unbranched portion of style slender and smooth, expanded at base; style 

base persistent at apex of achene, triangular, triangular-lanceolate, or discoid; 

achene surface transversely rugose or rugulose.. 14. Rhynchospora 

19. Fruit a terete or subterete drupe-like achene or bottle-shaped fructification, conical 

or acuminate at apex, smooth, longitudinally wrinkled, or minutely rugulosepapillate. 

.......................................................................................... 21 

21. Culms rounded-trigonous to terete proximally; ventral apex of sheath with a 

deeply U-shaped orifice; fruit an achene, drupe-like, ovoid-globose, acuminate 

at apex, longitudinally wrinkled when dry, minutely granular, naked at base 

...................................................................................................... 4. Cladium 

21. Culms subtriquetrous to trigonous proximally; ventral apex of sheath with a 

stiff, rounded-deltate contraligule; fruit a fructification, bottle-shaped, terete 

or obscurely trigonous, conical at apex with a truncate tip, smooth, minutely 

rugulose-papillate, with a dark brown patch on each side, often trisulcate on 

margins, with 3, minute, rounded-obdeltate, ciliate or sparsely ciliate, 

sometimes shortly bristle-tipped, reddish scales basally ....... 10. Lagenocarpus

240 


Fig. 45. CYPERACEAE. Terminology used in description of Cyperaceae. From Mori, S. et al. 1997. Vascular plants of central 



1. ABILDGAARDIA 

Abildgaardia Vahl, Enum. Pl. 2: 296. 1805. 

Caespitose perennials or annuals. Culms erect or ascending to deflexed, trigonous, obscurely angled, 

or subterete, ribbed. Leaves basal, blade bearing or reduced to bladeless sheaths; blade-bearing sheaths 

closed, the inner band membranous, splitting with age, lacking cilia or fimbriae at orifice; ligule absent; 

blades narrowly linear to filiform, thickened, flattened to involute, antrorsely scabrous on margins. 

Inflorescence a single spikelet or cluster of several spikelets at the summit of the culm, or a simple 

anthela, subtended by a single involucral bract which rarely exceeds the inflorescence; spikelets ovate 

or linear, subcompressed or spirally twisted, many-flowered, the lower 1-2 scales sterile; scales loosely 

imbricate, distichous or subdistichous, at least towards the base, dorsally keeled, smooth, carina 3- to 5nerved. 

Flowers bisexual, or sometimes staminate only in terminal scales of spikelet; hypogynous 

squamellae or bristles absent; stamens 2 or 3, the anthers narrowly oblong; style 3-branched, the 

unbranched portion narrowly wing-angled, smooth or glandular-puberulent at junction with the angled, 

glandular-hairy stigma branches, smooth or remotely fimbriate on margins below, expanded at base, 

disarticulating from the apiculate achene apex. Achene subglobose to obovoid, obscurely trigonous 

with convex faces, strongly tricostate, abruptly narrowed at apex to a pyramidal or truncate-pyramidal 

apiculum, short-stipitate at base, the surface verrucose or coarsely papillate with large, low, domeshaped 

papillae. Approximately 15 species in the New and Old World tropics. 

LECTOTYPE: Abildgaardia monostachyos (L.) Vahl (≡ Cyperus monostachyos L.), designated by 

Svenson, N. Amer. Fl. 18: 556. 1957. 

References: Kral, R. 1971. A treatment of Abildgaardia, Bulbostylis, and Fimbristylis (Cyperaceae) 

for North America. Sida 4: 57-227. Kral, R. & M.T. Strong. 1999. Eight novelties in Abildgaardia and 

Bulbostylis (Cyperaceae) from South America. Sida 18(3): 837-859. 

1. Abildgaardia ovata (Burm.f.) Kral, Sida 4: 72. 

1971; Carex ovata Burm.f., Fl. Ind. 194. 1768; 

Fimbristylis ovata (Burm.f.) J. Kern, Blumea 

15: 126. 1967. Type: Java. Collector unknown 

(holotype: G). 

Cyperus monostachyos L., Mant. Pl. 180. 1771; 

Abildgaardia monostachyos (L.) Vahl, Enum. 

Pl. 2: 296. 1805; Fimbristylis monostachyos 

(L.) Hassk., Pl. Jav. Rar. 61.1848; Iria 

monostachyos (L.) Kuntze, Revis. Gen. Pl. 2: 

751. 1891. Lectotype: India. Koenig s.n. 

(LINN-70.3), designated by Gordon-Gray, J. 

S. African Bot. 32 : 144. 1966. 

Cyperus caribaeus Pers., Syn. Pl. 1: 65. 1805. 

Type: West Indies. Collector unknown 

(holotype: probably at L). 

Fig. 46. A-D 

Caespitose perennial, (5-) 10-35 (-50) cm tall; 

rhizomes short, knotty. Culms obtusely trigonous 

to subterete, 0.4-0.8 mm wide, 6-10-ribbed, 

glabrous, light green to stramineous, smooth or 

antrorsely scabrous on angles just below 

subflattened apex. Leaves several; sheaths 

glabrous, prominently veined abaxially, the ventral 

band membranous, open at summit, prolonged on 

either side into an auricle; blades 3-30 cm × 0.3- 

1.1 mm, flattened to broadly U-shaped or involute, 

prominently veined on abaxial surface, essentially 

smooth adaxially, light green to stramineous, 

glabrous, smooth to antrorsely scabrous on 

margins, at least distally, abruptly acuminate to 

an often slightly curved tip. Inflorescence of 

1(-2) spikelets at the summit of the culm, 

ascending, subtended by a single involucral bract; 

rays when present, short, ribbed; spikelets ovate, 

6-14 × 3.3-6 mm, acute to acuminate, broadly 

acute to sub-rounded at base; scales ovate, 4-6 mm 

long, ca. as broad as long, thickly herbaceous, 

curvate-carinate, glabrous, light green to 

stramineous, carina faintly 5-nerved, greenish, 

prolonged beyond the acute apex as a stiff mucro. 

Stamens 3, the anthers 1.5-2.5 mm long, apiculate; 

style branches minutely fimbriate-scaly. Achene 

obscurely trigonous with convex sides, obovoid, 

2-3 × 1.5-2 mm, rounded to truncate at apex, 

apiculate, abruptly attenuate at base, stipitate, 

verrucose-tuberculate, stramineous to bone-white. 



Islands: In coastal and inland, often successional 

habitats; sea cliffs, ledges, rocky slopes, scrub

242 


forest and thickets, grassy slopes and clearings, 

lake shores and margins, open trails, pastures, 

savannas, lawns, and disturbed areas. Aguas 

Buenas, Arecibo, Bayamón, Cabo Rojo, Cataño, 

Ceiba, Dorado, Fajardo, Guajataca, Isabela, 

Manatí, Mayagüez, Quebradillas, Rincón, Río 

Grande, Sabana Grande, San Juan, San Sebastián, 

Trujillo Alto, Vega Alta, Vega Baja, and Yauco; 

Anegada, St. Croix, and St. John. 


2. BULBOSTYLIS 

Bulbostylis Kunth, Enum. Pl. 2: 205. 1837, nom. conserv. 

Aguas Buenas: Proctor 43297 (US). Bayamón: 

Sintenis 1210 (US). Cabo Rojo: González-Más 

1765 (US). Cataño: Bo. Palmas, Axelrod & Díaz 

12618 (UPRRP). Fajardo: Bo. Cabezas de San 

Juan, Axelrod 7154 (US). Guajataca: Guajataca 

State Forest, Liogier 9937(US). Manatí: Sintenis 

6615 (US). Sabana Grande, Sargent 679 (US). San 

Juan: Río Piedras, Stevenson 5655 (US). ST. CROIX: 

Parasol Hill, Thompson 438 (US). ST. JOHN: Dittlif 

Point, Woodbury 648a/7069 (VINPS). 

Annuals or perennials, tufted, rarely solitary; culms ascending to deflexed, often wiry, obtusely 

trigonous to subterete, coarsely ribbed. Leaves basal; sheaths closed at summit, splitting with age, coarsely 

nerved medially, usually scaberulous, pubescent or glabrous, the orifice oblique or truncate, long-ciliate 

or fimbriate along margins, rarely glabrous; ligule absent or present; blades capillary, setaceous, or 

narrowly linear, often canaliculate or crescent-shaped in cross section, antrorsely scabrous or scabrid on 

margins, strongly nerved abaxially, pubescent or glabrous. Inflorescence a simple or compound 

umbelliform cyme with elongate rays, a cluster of few to several spikelets at culm tips, or a solitary 

spikelet; involucral bracts 1-3 (-5), leaf-like, or wanting; rays filiform, ribbed, prophyllate, frequently 

with intraprophyllar buds; spikelets ovate to oblong-lanceolate, cylindrical or angled, sometimes 

compressed, many-flowered; scales spirally arranged, rarely somewhat 2-ranked, carinate, curvate-keeled 

or boat-shaped, dorsally obtuse to acute, pubescent or glabrous. Flowers bisexual; hypogynous squamellae 

or bristles absent; stamens 1-3, the anthers oblong; styles 3-branched, with minutely papillate or scaly 

branches, the unbranched portion cylindrical or 3-angled, smooth or rarely ciliate distally, typically 

disarticulating above the swollen, bulbous base. Achene trigonous to obtusely so, rarely plano-convex, 

obovoid, or sometimes oblong, with rounded angles, smooth, papillate, reticulate, or transversely rugulose. 

Approximately 80 species distributed in warm-temperate and tropical regions of the world. 

TYPE: Bulbostylis capillaris (L.) C. B. Clarke (≡ Scirpus capillaris L.), typ. conserv. 

Reference: Kral, R. 1971. A treatment of Abildgaardia, Bulbostylis, and Fimbristylis (Cyperaceae) 

for North America. Sida 4: 57-227. 

Key to the species of Bulbostylis 

1. Inflorescence a single spikelet at the summit of the culm, appearing lateral due to the erect lowest 

involucral bract which exceeds the spikelet and appears as a continuation of the culm................. 2 

2. Diminutive annual, 2-15 cm tall; spikelets often forming at culm bases; spikelet scales 2-2.6 

mm long, stramineous with a pale greenish carina, conspicuously and coarsely reddish dotted 

and lineolate over entire surface ........................................................ 3. B. curassavica 

2. Densely tufted perennial, 7-30 cm tall; spikelets not forming at culm bases; spikelet scales 3-3.5 

mm long, light brown with a dark olive-brown carina, finely reddish lineolate on sides 

.......................................................................................................................... 4. B. pauciflora 

1. Inflorescence with 2-many spikelets and at least some evident branching, terminal, anthelate or 

corymbose, open or contracted........................................................................................................ 3 

3. Culms and leaves hirsute ............................................................................. 7. B. vestita 

3. Culms and leaves glabrous, at most antrorsely scabrous on margins. ........................... 4 

4. Base of plant bulbous-thickened; spikelets 7-15 × 1.8-2.5 mm; anthers 2-3 mm long 

.................................................................................................................. 6. B. subaphylla


4. Base of plant not bulbous-thickened; spikelets 3-7 × 1-1.8 mm; anthers 0.5-1.2 mm 

long.................................................................................................................................. 5 

5. Coarse rhizomatous perennial with knotty rhizome, 47-100 (-135) cm tall; culms 0.8- 

2.7 (-3) mm wide; achenes somewhat plano-convex, narrowly obovate to oblong, finely 

lustrous-papillose .......................................................5. B. stenocarpa 

5. Slender caespitose perennials, 6-40 cm tall; culms 0.3-0.6 mm wide, achenes obovoid 

to broadly so, rugulose or finely papillose-rugulose.................................. 6 

6. Inflorescence open to subdense; carina of spikelet scales distinctly 3-nerved, not 

thickened, stramineous; achenes broadly obovoid, rugulose-papillose on sides, 

papillose on angles and at base, brown to often silvery-gray with pale margins at 

maturity ...........................................................2. B. capillaris subsp. insulana 

6. Inflorescence dense, turbinate to subcapitate; carina of spikelet scales 1-nerved or 

indistinctly 3-nerved, thickened, whitish, sharply contrasting with the brown-black 

sides of the scales; achenes narrowly obovoid, finely rugulose-papillose on sides 

only, light brown to brown at maturity ............1.B.antillana 

1. Bulbostylis antillana (Britton) Fernald, 

Rhodora 40: 392. 1938. Stenophyllus 

antillanus Britton, Bull. Torrey Bot. Club 43: 

447. 1916; Bulbostylis capillaris subsp. 

antillana (Britton) T. Koyama in R.A. 

Howard, J. Arnold Arbor. 60: 322. 1979. Type: 

Dominica; Grand Savanna. F. E. Lloyd 822 


Slender, densely tufted, short-lived perennial, 

9-35 (-40) cm tall; rhizome short, inconspicuous. 

Culms ascending, wiry, obtusely trigonous, 7- or 

8-ribbed, glabrous, smooth proximally, antrorsely 

scabrous on ribs distally, 0.4-0.6 mm wide. Leaves 

3-5 per culm; sheaths glabrous, with cinnamoncolored 

broad membranous margins, pale brown 

medially, the orifice oblique, densely ciliate with 

long crisped whitish trichomes; ligule absent; 

blades linear-filiform, 3-25 (-30) cm × 0.3-0.5 mm, 

canaliculate adaxially, substiffened, 5-costate 

abaxially, essentially smooth to spinulose-ciliate 

on margins and ribs with tubercle-based hairs, 

subacute to acuminate at apex. Inflorescence 

simple, corymbose, with short ascending branches 

or contracted into a turbinate or narrowly 

hemispherical aggregate without evident 

branching, 0.5-1.5 × 0.5-1.7 cm; involucral bracts 

1-3, ciliate proximally, the lowermost one 

elongate, leaf-like, exceeding the inflorescence; 

corymb branches to 1 cm long or wanting, 

terminated by small cymules of (1) 2-4 spikelets; 

spikelets lanceolate or oblong-lanceolate, 3-6(-7) 

×1.2-1.5 mm, acute, short-cuneate to sub-rounded 

at base; scales ovate, straight- to slightly curvate- 

keeled, dorsally acute, puberulent on sides, reddish 

brown with brown-black sides distally, margins 

narrowly scarious, ciliate, carina obscurely 3nerved, 

thickened, whitish, sharply contrasting 

with the brown-black sides of the scale, prolonged 

beyond the acute apex as a mucro. Stamens 2, the 

anthers 0.6-0.9 mm long, apiculate. Achene 

narrowly obovoid or obovoid, trigonous, 0.8-1.1 

× 0.6-0.8 mm, truncate and shallowly 3-lobed at 

apex, short-cuneate to short-attenuate at base, with 

a finely and closely papillose rugosity, 

stramineous, pale brown, or often yellowish brown 

at maturity; style base depressed-globose, 2-3 mm 

wide. 

General distribution: Dominican Republic, 

Puerto Rico, and the Lesser Antilles. 

Distribution in Puerto Rico: On clay or sandy 

soils on rocky slopes and savannas. Cabo Rojo, 

Sabana Grande, Vega Baja, and Yabucoa. 


Cabo Rojo: Proctor 44020 (SJ). Sabana Grande: 

Susúa Forest Reserve, Delgado Montano & 

González-Más 53 (MAPR); González-Más 2035 

(MAPR). 

2. Bulbostylis capillaris subsp. insulana M.T. 

Strong, subsp. nov. Type: Puerto Rico; Bo. 

Tierras Nuevas Salientes, just S of Laguna 

Tortuguero, near sea level, 15 May 1977, 

Woodbury s.n. (holotype: SJ; photo at US). 

B. capillaris subsp. capillaris spicis ovoidlanceoloidis 

vel anguste lanceoloidis, 

gracilibus; acheniis rugulose-papillosis 

differt.

244 


Caespitose perennial, 6-40 cm tall; rhizome 

short, thickened. Culms slender, wiry, trigonous 

to subterete, 9-ribbed, ribs smooth proximally, 

antrorsely scabrid distally, 0.3-0.6 mm wide. 

Leaves 2-3 per culm; sheaths elongate, brown to 

stramineous, essentially smooth, with broad 

scarious margins, the orifice oblique, longfimbriate 

apically with tawny hairs; ligule absent; 

blades filiform, 2-12 cm × 0.2-0.5 mm, 

canaliculate adaxially, coarsely 5-ribbed 

(including ribbed margins) abaxially, these often 

smooth or antrorsely scabridulous (at least 

distally), often spirally twisted distally. 

Inflorescence a simple to compound, open or 

contracted anthela, 0.8-3 × 0.4-2 cm; involucral 

bracts 3-4, shorter than the inflorescence, linearfiliform, 

with broad membranous bases, ciliate on 

margins; primary rays filiform to capillary, to 20 

mm long; spikelets ovoid-lanceoloid or narrowly 

lanceoloid, 3-7 × 1-1.7 mm, acute, short-cuneate 

to sub-rounded at base; fertile scales ovate to 

broadly lanceolate, 1.4-2.2×1.3-1.8 mm, curvate 

keeled or only slightly so, dorsally acute, often 

ciliate on the erose margins, deep brown or redbrown, 

with black sides distally, puberulent or 

sometimes glabrous, carina with 3 distinct nerves, 

equaling or rarely prolonged beyond the obtuse to 

acute apex. Stamens 3, the anthers 0.5-0.6 mm, 

long, blunt to apiculate. Achene trigonous, 

obovoid, 0.9-1.2 × 0.5-0.7 mm, truncate and 

slightly 3-lobed at apex, attenuated at base, finely 

transversely rugulose-papillose at maturity, 

papillose on angles, nearly smooth when 

immature, brown to often silvery-gray with pale 

margins at maturity; style base globuliform, 0.2- 

0.3 mm wide. 


Puerto Rico. 

Distribution in Puerto Rico: Sandy savannas 

at Laguna Tortuguero and vicinity. Manatí and 

Vega Baja. 

Note: This taxon differs from the typical 

subspecies in having slender ovoid-lanceoloid or 

narrowly lanceoloid spikelets and a rugulosepapillose 

achene surface vs. ovoid spikelets and 

transversely rugose achene surface. 


Manatí: Bo. Tierras Nuevas Saliente, just S of 

Laguna Tortuguero, Woodbury s.n. (SJ). Vega 

Baja: Laguna Tortuguero, Delgado 40, 47, 

(MAPR); 105 (MAPR-3, UPR, UPRRP); Bo. 

Algarrobo, just S of Laguna Tortuguero, Proctor 

et al. 47745 (SJ); Laguna Tortuguero, González- 

Más & Woodbury 277b (MAPR); Tortuguero area, 

Liogier et al. 33574 (UPR); Laguna Tortuguero, 

Sep 1960, Woodbury s.n. (MAPR, UPRRP); Feb 

1962, Woodbury s.n. (UPR); Feb 1968, Woodbury 

s.n. (NY); Sep 1969, Woodbury s.n. (NY, UPR); 

Jun 1970, Woodbury s.n. (UPR). 

3. Bulbostylis curassavica (Britton) Kük. ex 

Ekman., Ark. Bot. 22A(16): 8. 1929. 

Stenophyllus curassavicus Britton, Bull. 

Torrey Bot. Club 43: 445. 1916, nom. et stat. 

nov. for Bulbostylis floccosa var. β “(?)” 

pumilo C. B. Clarke in Urban, Symb. Antill. 

5: 290. 1907; Fimbristylis curassavica 

(Britton) Alain, Bull. Torrey Bot. Club 92: 

290. 1965. Type: Curaçao. Suringar s.n. 


Bulbostylis curassavica var. pallescens Kük. & 

Ekman ex Urb., Ark. Bot. 22A(17): 6. 1929. 

Type: Haiti. Ekman 8749 (lectotype: US; 

isolectotype: US), here designated. 

Diminutive, tufted annual, 2-15 cm tall. 

Culms ascending to erect, filiform to 

subcompressed, 5- to 7-ribbed, antrorsely 

scabrous-hispidulous, 0.3-0.5 mm wide. Leaves 

1-3 per culm; sheaths short, sparsely antrorsely 

scabrous-hispidulous, tan, the orifice truncate, 

long whitish fimbriate; ligule absent; blades linearfiliform 

or subflattened, 0.6-8 cm × 0.3-0.5 mm, 

soft, flexuous, canaliculate adaxially, ribbed 

abaxially with antrorsely scabrous-hispidulous 

ribs, acuminate at apex. Inflorescence a solitary, 

pseudolateral spikelet at the summit of the culm, 

often with spikelets forming at the base of culm 

as well; involucral bracts 1-3, the lowermost one 

erect, appearing as a continuation of the culm; 

spikelets ovoid to oblong-ovoid, 4-7 × 1.4-2 mm, 

spreading to 3 mm wide with mature achenes, 

subcompressed, spirally twisted with distichous 

scales, acute, cuneate at base; scales ovate, 

curvate-keeled, dorsally acute, 1.4-2.6 × 1-2 mm, 

glabrous, stramineous, coarsely reddish dotted and 

lineolate, margins scarious, ciliate, carina 3nerved, 

pale greenish, slightly prolonged beyond 

and excurved at the acute apex. Stamens 2, the 

anthers 0.6-0.9 mm long, apiculate. Achene 

broadly obovoid or sub-rounded, obtusely 

trigonous, 0.6-0.9 × 0.4-0.6 mm, broadly rounded 

to subtruncate at apex, obtuse at base, foveate, 

lustrous, pale brown to reddish brown; style base


depressed-globose, 0.2-0.3 mm wide, brownish 

black. 

General distribution: Haiti, Puerto Rico, 

Virgin Islands, and Curacao. 


Islands: In soil pockets on limestone rocks. Caja 

de Muertos, Guánica, Maricao, Mona Island, and 

Yauco; Anegada. 

Note: The name (new combination) 

Bulbostylis curassavica (Britton) Kük. ex Ekman 

was inadvertently validated by an indirect 

reference to the basionym when the varietal name 

Bulbostylis curassavica (Britton) Kük. var. 

pallescens Kük. & Ekman ex Urb. was listed in 

an enumeration of plants on Navassa Island by 

Ekman. 


Guánica: San Jacinto Beach, González Más 990 

(US). Yauco: at end of Rt. 333 in Guánica Reserve, 

Taylor et al. 9504 (UPRRP). 

4. Bulbostylis pauciflora (Liebm.) C. B. Clarke 

in Urban, Symb. Antill. 5: 290. 1907, nom. 

conserv.; Oncostylis pauciflora Liebm., 

Kongel. Danske Vidensk. Selsk. Skr. 

Naturvidensk. Math. Afh., Ser. 5. 2: 241. 1851. 

Type: St. Croix, U.S. Virgin Islands. Oersted 

s.n. (holotype: C; isotype: NY!). 

Stenophyllus portoricensis Britton, Torreya 13: 

216. 1913; Bulbostylis portoricensis (Britton) 

Fernald, Rhodora 40: 392. 1938; Fimbristylis 

portoricensis (Britton) Alain, Bull. Torrey 


Guánica, Britton & Shafer 1916 (holotype: 


Bulbostylis ekmanii Kük., Repert. Spec. Nov. 

Regni Veg. 23: 197. 1926. Lectotype: Cuba; 

Nipe Bay. Ekman 7341 (US!), here 

designated. 

Fig. 46. E-H 

Densely tufted perennial, 7-30 cm tall; 

rhizomes short; roots filiform. Culms 

subtrigonous, wiry, 4- or 5-ribbed, soft, flexuous, 

reddish or reddish brown, glabrous, antrorsely 

scabrid on ribs distally, 0.2-0.3 mm wide. Leaves 

1-3 per culm; sheaths glabrous, prominently 3veined 

abaxially, membranous and finely veined 

on margins, reddish brown, the orifice oblique, 

long whitish fimbriate at apex; ligule absent; 

blades wiry, linear-filiform, 2-16 cm × 0.2-0.3 mm, 

canaliculate adaxially, prominently 3- to 5-ribbed, 

abaxially, reddish or chestnut-colored, glabrous, 

antrorsely scabrid on margins and ribs, at least 

distally, the apex acuminate. Inflorescence a single, 

often pseudolateral, spikelet at the summit of the 

culm; involucral bracts 3, the lowermost with 

sheathing base and apical fimbriae, elongate, often 

overtopping the spikelet and appearing as a 

continuation of the culm, the upper two typically 

scale-like, shorter than the spikelet; spikelets 

ovate-lanceolate, subcompressed, 4.5-10 (-12) × 

1.3-2 mm, often falcate; scales narrowly-ovate, 

2.7-3.5 ×1-2 mm, glabrous, light brown to reddish 

brown, straight- to slightly curvate-keeled, 

dorsally acute to obtuse, finely reddish lineolate 

on sides, carina 3-nerved, dark reddish brown, 

ending at the acute apex or prolonged and shortmucronate 

on basal scales of spikelet. Stamens 3, 

the anthers 1-1.3 mm long, with a triangular 

prickly apiculum. Achene ellipsoid-obovoid, 

trigonous, 1.3-1.6 × 0.6-0.9 mm, obtuse at apex 

and base, transversely rugulose, yellowish brown 

to brown, the angles often lighter, yellowish; style 

base depressed-pyramidal, 0.2-0.3 mm wide, 

sometimes deciduous. 


Puerto Rico, Virgin Islands, and the Lesser 

Antilles. 


Islands: On limestone or serpentine substrate, 

often in dry, rocky or sandy areas of scrub forest 

and thickets along the coast. Cabo Rojo, Guánica, 

Ponce, Sabana Grande, and Vieques; Anegada, St. 

Croix, St. John, and St. Thomas. 


Cabo Rojo: Morrillos de Cabo Rojo, N.L. Britton 

et al. 4710 (US). Ponce: Bo. Cañas, N off Rt. 2 

just before prison, Axelrod & Chávez 7102 (US). 

Vieques: Punta Este, Proctor 48570 (US). ST. 

CROIX: Sandy Point, Raunkiaer s.n. (US); Acevedo- 

Rdgz. et al. 5304 (US). 

5. Bulbostylis stenocarpa Kük., Bot. Jahrb. Syst. 

56(Beibl. 125): 15. 1921. Type: Brazil; 

Amazonas. Ule 8069 (holotype: B, destroyed; 

isotypes: K!, US!). 

Bulbostylis papillosa Kük., Repert. Spec. Nov. 

Regni Veg. 23: 198. 1926; Fimbristylis 

papillosa (Kük.) Alain, Bull. Torrey Bot. Club 

92: 290. 1965. Lectotype: Cuba. Ekman 2690 

(US!), here designated.

246 


Fig. 46. A-D. Abildgaardia ovata. A. Habit, detail of leaf blade apex, and detail of culm showing apex of sheaths. B. Spikelet. C. 

Flower. D. Achene. E-H. Bulbostylis pauciflora. E. Habit, and detail of culm showing apex of sheaths. F. Spikelet. G. Flower. H. 

Achene. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78. 

Caespitose perennial, 47-100 (-135) cm tall; 

rhizome short, thickened, knotty. Culms erect, stiff, 

breaking easily when bent, obtusely trigonous to 

subterete, low-ribbed, shallowly grooved, 

essentially smooth, 0.8-2.7 (-3) mm wide. Leaves 

numerous; sheaths herbaceous, low-ribbed 

dorsally, sub-scarious and finely veined along 

reddish margins, scabrid on veins, brownish 

dorsally, gradually narrowing distally, longfimbriate 

with pale fimbriae apically at junction 

with blade; ligule a short, ciliate, prolongation of 

the adaxial apex of sheath; blades mostly 

ascending, 7-77 cm × (0.5-) 0.8-1.8 mm, 

subflattened or slightly involute, abaxially convex 

and closely ribbed, smooth or antrorsely 

scaberulous, margins slightly thickened, adaxial 

surface smooth, the apex sub abruptly acuminate. 

Inflorescence a terminal, compound, narrowly to 

broadly turbinate, often dense and broom-like 

corymb of densely fascicled spikelets at tips of 

short, stiff, primary branches, 1.5-8 × 1-5 cm; 

involucral bracts 2-3, shorter than the


inflorescence or the lowermost one sometimes 

slightly exceeding it; spikelets lanceoloid, 3.5-5 

(-6) × 1-1.3 mm, acute, short-cuneate at base; 

scales loosely imbricate, ovate, 1.7-2.2 × 1-1.6 

mm, strongly curvate-keeled, dorsally acute, 

scabridulous, dark red-brown to brown-black, 

carina dark green, obscurely 3-nerved, prolonged 

beyond the acute apex as a slightly excurved 

mucro. Stamens 3, the anthers 0.8-1.2 mm long, 

with a lanceolate apiculum. Achene somewhat 

plano-convex, oblong-obovate, 1-1.3 × 0.5-0.7 

mm, truncate at apex, short-cuneate at base, 

indistinctly tricostate, often with pale angles, graybrown 

at maturity, finely lustrous-papillose; style 

base bulbiform, 0.1 mm diam. 


America, West Indies, and South America. 

Distribution in Puerto Rico: On grassy slopes 

and road banks. Aguada and Moca. 

Note: Bulbostylis stenocarpa has been 

included in the synonymy of B. junciformis 

(Kunth) C. B. Clarke by some authors, but its 

coarser habit, wider leaf blades, and somewhat 

plano-convex, oblong-obovate, lustrous-papillose 

achene (vs. obovoid and honeycomb-reticulate 

achene of B. junciformis) is distinct from the latter. 


Aguada: Sargent 613 (US). 

6. Bulbostylis subaphylla C. B. Clarke in Urban, 

Symb. Antill. 2: 86. 1900; Stenophyllus 

subaphyllus (C. B. Clarke) Britton, Bull. Dept. 

Agric. Jamaica 5 (Suppl. 1): 12. 1907. Type: 

Cuba. C. Wright 1533 (holotype: K). 

Stenophyllus harrisii Britton, Torreya 20: 83. 1920; 

Fimbristylis harrisii (Britton) C.D. Adams, 

Phytologia 21: 66. 1971. Type: Jamaica; Old 

England Falls. Harris 12980 (holotype: NY!; 


Densely caespitose, rhizomatous perennial, 

22-60 (-75) cm tall; rhizome short, thickened; 

Culms erect, obtusely trigonous to subterete or 

subcompressed, firm but flexuous, many-ribbed, 

smooth, often antrorsely scabrid on margins near 

apex, 0.5-0.8 mm wide. Leaves 5-12 per culm, 

crowded at base; sheaths short, glabrous, pale, 

reddish brown proximally, wooly-fimbriate at the 

oblique orifice, with pale crisped hairs; ligule 

absent or partially ligulate with two dense tufts of 

hair on adaxial margins at junction of sheath and 

blade; blades linear-filiform, 4-35 (-53) cm × 0.6- 

1.2 (-1.5) mm, thickly herbaceous, substiffened, 

crescent-shaped, subinvolute, or subflattened, 5to 

7-nerved abaxially, antrorsely scabrid on 

margins, the apex acute to acuminate, often with 

one margin curving to tip. Inflorescence a simple, 

open anthela with short branches or contracted 

with little evident branching, 2-4 × 1-2.5 (-3) cm; 

involucral bracts 1-3, leaf-like, but reduced, ciliate 

with crisped hairs on margins of sheathing base; 

branches, when present, to 2.5 cm long; spikelets 

borne singly at branch tips, oblong-lanceolate, 7- 

15 × 1.8-2.5 mm, subcompressed, acute to short 

acuminate at apex, cuneate at base, often 

proliferous; scales narrowly ovate, 3-5 × 1.4-2.3 

mm, slightly curvate- to straight-keeled, dorsally 

acute to widely obtuse, sparsely hirtellous to 

glabrous, reddish brown, margins narrowly 

scarious, ciliate-fimbriate with crisped hairs, 

carina 3-nerved, pale green, prolonged beyond the 

acute apex as a short mucro. Stamens 3, the anthers 

1.5-2.7 mm long, bluntly apiculate. Achene 

obovoid, trigonous, subdorsiventrallycompressed, 

1.2-1.5 × 0.8-1.1 mm, rugulose, shiny, 

pale brown to brown or yellowish brown, often 

with lighter-colored angles; style base obtusely 3angled, 

depressed, minutely 3-lobed above each 

angle, with a minute central apiculum, 0.3-0.4 mm 

wide. 

General distribution: Cuba, Haiti, Jamaica, 

and Puerto Rico. 

Distribution in Puerto Rico: In dry, rocky 

areas of scrub forests and thickets. Guánica and 

Ponce. 


Guánica: Guánica State Forest, Proctor & Rivera 

47162 (US). Ponce: Bo. Cañas, off Rt. 2, behind 

the Holiday Inn Hotel, Axelrod et al. 4724 (US). 

7. Bulbostylis vestita (Kunth) C. B. Clarke in 

Urban, Symb. Antill. 2: 87. 1900; Isolepis 

vestita Kunth, Enum. Pl. 2: 210. 1837; 

Oncostylis vestita (Kunth) Nees in Martius, 

Fl. Bras. 2(1): 88. 1842; Scirpus vestitus 

(Kunth) Rchb. ex Boeck., Linnaea 36: 753. 

1870; Fimbristylis vestita (Kunth) Hemsl., 

Biol. Cent.-Amer., Bot. 3: 460. 1885; 

Stenophyllus vestitus (Kunth) Britton, Bull. 

Torrey Bot. Club 43: 446. 1916. Type: 

Surinam. Weigelt s.n. (holotype: B, destroyed; 

isotype: K!).

248 


Scirpus hirtus Griseb., Cat. Pl. Cub. 241. 1866. 

Type: Cuba. C. Wright 3383 (holotype: 

GOET; isotype: GH). 

Bulbostylis hirtella sensu Urban, Symb. Antill. 4: 

120. 1903, non (Schrader) Nees von 

Esenbeck, 1843. 

Caespitose perennial, (7-) 12-62 cm tall; 

rhizome short, bearing very dense clumps of 

culms, seemingly arising from a common point. 

Culms stiffly ascending to erect, obtusely 

trigonous to subterete, 8- or 9-ribbed, hirsute, 0.4- 

0.8 mm wide. Leaves 5-10 or more per culm; 

sheaths hirsute, brownish with broad, scarious, 

reddish brown, finely veined margins, fimbriate 

at the oblique orifice; ligule absent; blades 2-30 

cm × 3-5 mm, ascending to spreading, often 

curling or recurved, filiform or subflattened, 

convex and 3-5 veined abaxially, spreadinghirsute, 

canaliculate and glabrous adaxially. 

Inflorescence a terminal, simple, anthela with short 

rays bearing small clusters or fascicles of spikelets 

at ray tips, or sometimes a single lobed cluster of 

spikelets at the summit of the culm, 4-30 mm in 

diam.; involucral bracts 2-4, leaf-like but reduced, 

shorter than the inflorescence or sometimes the 

lowermost one exceeding it; spikelets ovoid to 

ovoid-lanceoloid, 3.4-6 ×1.4-2 mm; scales broadly 

ovate to sub-rounded, curvate-keeled, dorsally 

acute, 1.8-2.2 × 2-2.2 mm, rugulose and hirtellous, 

deep red-brown, carina 3-nerved, or obscurely so, 

thickened, prolonged as a prominent, slightly 

excurved mucro. Stamens 3, the anthers 0.8-1 mm 

long, with a triangular apiculum bearing a tuft of 

crystalline prickles at tip. Achene obovoid, 

obtusely trigonous with convex faces, 0.7-0.9 × 

0.6-0.7 mm, truncate at apex, short-cuneate at base, 

3. CAREX 

Carex L., Sp. Pl. 972. 1753, nom. conserv. prop. 

often lustrous, finely transversely rugulosepapillate 

to nearly smooth, dark brown or gray 

brown with pale angles, whitish when immature; 

style base obtusely trigonous, depressed, with a 

central apiculum, 0.2-0.3 mm wide. 


America, West Indies, Trinidad, and South 

America 

Distribution in Puerto Rico: In dry, red, 

lateritic soils or sandy soils of clearings, grassy 

areas, slopes, pastures, and roadsides. Maricao, 

Mayagüez, Moca, San Sebastián, and Vega Baja. 


Mayagüez: Cerro Las Mesas, Bo., Proctor 42186 

(US); Las Mesas, Holm 111 (US). San Sebastián: 

Sargent 347 (US). Moca: Rd. 110, km 92, 

González-Más 1806 (US). 

Excluded species 

Bulbostylis junciformis (Kunth) C. B. Clarke, 

Trans. Linn. Soc. London, Bot. 4: 512. 1895. 

Reported by Liogier & Martorell (1982: 212; 

2000: 238) for Puerto Rico based on a Woodbury 

collection from Laguna Tortuguero. No specimens 

have been seen of this species from the flora area. 

This name has been misapplied to B. stenocarpa 

Kük. by some authors. 

Bulbostylis langsdorffiana (Kunth), C.B.Clarke in 

Urban, Symb. Antill. 2: 89. 1900. 

Cited by C. B. Clarke for Puerto Rico (in 

Urban, Symb. Antill. 2: 89. 1900), based on a 

misidentification of Sintenis 1208 which is 

Bulbostylis vestita (Kunth) C. B. Clarke. 

Grass-like perennial herbs. Tufted or with short or elongate creeping rhizomes, sometimes 

stoloniferous, monoecious, rarely dioecious. Culms triquetrous or trigonous, rarely subterete, solid or 

sometimes hollow. Leaves 3-ranked, basal and cauline or primarily basal, the lowermost often bladeless; 

sheaths finely veined, often with a membranous inner band which is closed at the concave to truncate 

orifice; ligule often present; blades flat, involute, V-shaped , or folded, narrowly linear, rarely lanceolate 

or elliptic and subpetiolate, finely veined. Inflorescence a single terminal spike or a terminal and series 

of 1-several subcontiguous or remote spikes or panicles from the upper leaf-like or scale-like bracts; 

spikes sessile or peduncled with spirally-arranged perigynia, unisexual, androgynous, or gynecandrous, 

rarely mixed, inflorescences bearing unisexual spikes often with the 1-several staminate spikes terminal


and pistillate spikes below. Flowers unisexual; hypogynous squamellae or bristles absent; staminate 

flowers with 3 stamens subtended by a single scale; pistillate flowers a single ovary contained within a 

tight or inflated, 2-sided, trigonous or rounded, often bottle-shaped perigynium, subtended by a single 

scale; style 2- or 3-branched, continuous with the ovary and persistent or articulated with it and deciduous, 

straight or flexuous, often thickened at base, the tips of the stigmas exserted from the apex or often 

bidentate beak of the perigynium; perigynium membranous, chartaceous, or coriaceous, often ribbed or 

nerved with raised or impressed nerves, or smooth, sometimes winged, glabrous, pubescent, hispid, 

papillose, puncticulate, or smooth, sometimes spongy at base, the apex often beaked, bidentate or bifurcate. 

Fruit an achene, lenticular, biconvex, or trigonous, generally green, puncticulate or sometimes smooth. 

Approximately 2,000 species, cosmopolitan, widespread in temperate zones and montane tropics. 

LECTOTYPE: Carex hirta L., typ. conserv. prop. 

Note: Britton, in “The sedges of Jamaica”, Bull. Dept. Agric. Jamaica 5, Suppl. 1. 1907, designated 

Carex pulicaris L. as the lectotype of Carex. However, Jarvis (Taxon 41: 559. 1992) has proposed C. 

hirta L. as the lectotype with a conserved type following the proposal of Hitchcock & Green (1929) in 

International Botanical Congress Cambridge (England), Nomenclature Proposals by British Botanists, 

London. The spermatophyte committee has accepted this proposal but at this juncture it is still in limbo 

awaiting decision by the general committee. 

References: Kern, J.H. & H.P. Noteboom. 1979. Cyperaceae-II, 28. Carex. Pp. 107-183. In: C.G.G.J. 

van Stennis, ed., Flora Malesiana, Vol. 9, Part 1, Noordhoff International Publishing, Leyden, The 

Netherlands. Reznicek, A.A. 1993. Carex (Cyperaceae). Pp. 243-267. In: R. McVaugh, Flora Novo- 

Galiciana, Vol. 13: Limnocharitaceae to Typhaceae, The University of Michigan Herbarium, Ann Arbor. 

Key to the species of Carex 

1. Inflorescence moniliform with setaceous bracts; perigynia plano-convex, wing-margined, densely 

disposed in gynecandrous spikes; style 2-branched; achenes biconvex ..........................1. C. longii 

1. Inflorescence a series of 3-11 narrowly pyramidal panicles from the upper leaf-like bracts; perigynia 

trigonous, not wing-margined, laxly disposed in androgynous spikes; style 3-branched; achenes 

trigonous................................................................................................... 2. C. polystachya 

1. Carex longii Mack., Bull. Torrey Bot. Club 49: 

373. 1922. Type: United States; New Jersey, 

Cape May County, Long s.n. (holotype: PH). 

Caespitose, forming small to medium-sized 

clumps, (15-) 30-80 (-120) cm tall; rhizome short. 

Culms erect, slender, 1.3-3 mm wide, trigonous 

to sharply so, stiff and hardened, smooth 

proximally, scabrous on angles at apex. Leaves 4- 

6, lower cauline, the basal ones often bladeless; 

sheaths elongate, tight, proximal ones pale brown, 

cauline ones green, the inner band herbaceous, 

prominently veined, with concave orifice; ligule 

rounded, 2-6 mm long; blades linear, 6-40 cm × 

1.5-4 (-4.5) mm, plicate, glabrous, long-acuminate. 

Inflorescence moniliform, a series of (2-) 3-10 

(-11), subcontiguous, sessile, ellipsoid to obovoid 

spikes at the summit of the culm, (1.4-) 2.2-6 cm 

long; inflorescence bracts setaceous, reduced 

distally; spikes gynecandrous, 6-17 × 4-6 mm, 

obtuse, the scales appressed-ascending or slightly 

spreading; pistillate scales ovate, 2.2-3.7 × 1.1- 

1.8 mm, obtuse to acute, lustrous, silvery white to 

pale brown, midcosta 3-nerved, green, the 

midnerve prolonged as a short awn or mucro on 

proximal scales, ending short of the tip on distal 

scales. Anthers 1-2.3 mm long; style 2-branched; 

perigynia plano-convex, broadly obovate or 

suborbicular, broadest near the middle, 3-4.5 × 

1.6-2.8 mm, light silvery green or silvery brown, 

3- to 9-nerved adaxially, 5- to 13-nerved abaxially, 

the margins thinly winged, abruptly contracted into 

a scabrous-margined, bidentate beak, 0.7-1.3 mm 

long, with teeth ca. 0.2-0.4 mm long; achenes 

biconvex, ovate to oblong-ovate, 1.4-1.7 × 0.9- 

1.1 mm, short-stipitate, pale brown. 

General distribution: North America, Mexico, 

Central America, West Indies and South America;

250 


introduced and naturalized in Hawaii and New 

Zealand. 

Distribution in Puerto Rico: Only recently 

collected in an open area along roadside of Hwy 

184, 902 m, Guayama. Because the center of 

distribution of Carex longii is in the United States 

and it is not known historically from Puerto Rico, 

this occurrence likely represents a recent 

introduction. 

Specimens examined: PUERTO RICO: Guayama, 

Sierra de Cayey, along Hwy 184 E of jct. Hwy 

179 at Communication Tower facility, Worthington 

31243 (photo at US, UTEP). 

2. Carex polystachya Sw. ex Wahlenb., Kongl. 

Vetensk. Acad. Nya Handl. 24: 149. 1803; 

Carex cladostachya f. polystachya (Sw. ex 

Wahlenb.) C. B. Clarke ex Lindm., Bih. 

Kongl. Svenska Vetensk.-Akad. Handl. 26, 

III(9): 36. 1900. Type: Jamaica. Swartz s.n. 

(holotype: S-Sw. R-925). 

Carex cladostachya Wahlenb., Kongl. Vetensk. 

Acad. Nya Handl. 24: 149. 1803. Type: 

Jamaica. Swartz s.n. (holotype: S-Sw. R-912). 

Carex dussiana Boeck., Beitr. Cyper. 2: 42. 1890. 

Type: Martinique. Duss 763b (holotype: B, 

destroyed). 

Fig. 47. A-F 

Caespitose, 15-70 (-100) cm tall; rhizome 

short, woody. Culms erect, slender, trigonous, 

often channeled along one side distally, smooth, 

sometimes antrorsely scabrous on margins distally, 

otherwise glabrous, 0.8-1.8 mm wide. Leaves 5-9 

per culm, primarily basal, 1-2 cauline; sheaths 

tight, elongate, finely veined, green to stramineous 

or tinged with yellowish brown proximally, the 

basal sheaths often fibrillose at base, the inner band 

with concave orifice; ligule a membranous, 

inverted widely V-shaped band, 0.3-0.5 mm long; 

blades narrowly linear, 12-60 (-80) cm × (1.2-) 

1.5-6 (-8) mm, flattened-plicate, antrorsely 

scabrous on margins and abaxial midvein, graygreen, 

attenuate to triquetrous apex. Inflorescence 

a terminal and series of 3-11 narrowly pyramidal, 

long-pedunculate partial panicles from the upper 

3-4 leaf-like bracts, 1-3 from each node, 1.5-5 × 

0.6-1.7 (-2.2) cm; spikes 1-25, androgynous, 

simple or sometimes compound, ovate to oblongovate, 

4-16 × 3-5 mm, staminate portion 1.5-6 × 

0.6-1.2 mm with 2-12 flowers, pistillate portion 

3-10 × 3-5 mm with 6-18 flowers; scales ovate 

to ovate-lanceolate, narrowly scarious on margins, 

glabrous to antrorsely scabrous distally, carina 1to 

3-nerved, prolonged beyond the scarious, acute 

to emarginate apex as a straight to slightly 

excurved, antrorsely scabrous awn; staminate 

scales 1.4-3 × 0.9-1.4 mm, pale reddish brown; 

pistillate scales 0.8-2 × 0.8-1.6 mm (excluding 

awn), green to stramineous or reddish brown, often 

red lineolate distally along margins. Perigynia 

ellipsoid- to obovoid-fusiform, 2.5-4 × 0.7-1.2 mm 

(including beak), trigonous, with plane to concave 

sides, spreading, short-stipitate, membranous, 

distinctly 12- to 15-nerved, green, glabrous to 

antrorsely scabrous distally, the subulate beak 

straight to slightly recurved, shallowly to deeply 

bidentate. Anthers 1-1.7 mm long, with a prickly 

apiculum; style 3-branched. Achene elliptic, 

trigonous, with concave sides, 1.5-2.3 × 0.7-1.1 

mm, obtuse at apex, short-apiculate, shortattenuate 

at base, stipitate or subsessile, 

puncticulate, tightly enveloped by the perigynium, 

pale brown. 



Distribution in Puerto Rico: Shaded, wet, 

montane forest on rocky slopes and roadbanks, 

cliffs, along trails, thickets, and stream banks from 

480-1250 m. Adjuntas, Arecibo, Cayey, Ceiba, 

Jayuya, Luquillo, Maricao, Naguabo, Orocovis, 

Ponce, Río Grande, Salinas, and Villalba. 

Selected specimens examined: PUERTO 

RICO: Luquillo: Luquillo Forest, road towards El 

Yunque Mt., González-Más 1549 (US). Jayuya: 

Los Tres Picachos, Sargent 3063 (US); near Salto 

Doña Juana, González-Más 2189 (US). Naguabo: 

Sierra de Naguabo, Monte El Duque, Shafer 2254 

(US); Loma Icaco, Shafer 3434 (US). Río Grande: 

Sierra de Luquillo, Hioram 365 (US); El Yunque, 

McKee 10642 (US). Salinas: Bo. Lapa, summit 

area of East Peak, Las Tetas de Cayey, Proctor 

43523 (US).


4. CLADIUM 

Cladium P. Browne, Civ. Nat. Hist. Jamaica 114. 1756. 

Large to medium-sized, often emergent perennials; rhizomes horizontal, scaly, stolons sometimes 

present below culm bases; roots coarse. Culms leafy, often proliferous at lower nodes. Leaves numerous, 

basal and cauline; sheaths elongate; ligule absent; blades broadly linear, flattened, coarsely antrorsely 

serrulate-scabrous on margins. Inflorescence paniculate, consisting of a terminal and series of 1-many 

lateral partial compound corymbs from the upper leaf-like bracts; branches often elongate, the spikelets 

in small heads or fascicles at branch tips; spikelets ovoid, with irregularly spirally imbricated scales 

borne on a flexuose rachilla; scales 6-8, the basal 4-6 sterile, or sometimes staminate, often smaller than 

the 2 upper fertile bisexual ones, only the terminal floret producing an achene, the lower floret with an 

abortive pistil. Flowers bisexual; hypogynous squamellae or bristles absent; stamens 2; style 3-branched. 

Achene ovoid, subterete, smooth, the pericarp thick and corky; bristles absent. As recognized in the 

strict sense here excluding Machaerina and Baumea, there are four species in warm-temperate and 

tropical regions of Europe, Asia, Pacific Islands, North America, Central America, and South America. 

TYPE: Cladium jamaicense Crantz [as “iamaicense”]. 

1. Cladium jamaicense Crantz, Inst. Rei Herb. 

1: 362. 1766; Schoenus cladium Sw., Prodr. 

Veg. Ind. Occ. 19. 1788, nom. illeg.; Mariscus 

jamaicensis (Crantz) Britton ex Small, Fl. 

Miami 31. 1913; Cladium mariscus subsp. 

jamaicense (Crantz) Kük., Repert. Spec. Nov. 

Regni Veg. Beih. 40(1): 523. 1938. Lectotype: 

Jamaica. Browne s.n. (LINN-68.2), 

designated by McVaugh, Fl. Novo-Galiciana 

13: 268. 1993. 

Cladium occidentale sensu Grisebach, Fl. Brit. W. 

I. 573. 1864, non Schrader, 1806. 

Fig. 47. G-L; 65. G 

Coarse, emergent, rhizomatous perennial, 1- 

3 m tall; rhizome horizontal, 1-2 cm thick, often 

emitting stout stolons; sheathing base of culm 2-5 

cm wide. Culms solitary, sometimes proliferous 

from lower nodes, rounded-trigonous to terete, 

stiff, hardened, finely striate, glabrous, 4-15 mm 

wide. Leaves numerous; sheaths slightly spongythickened, 

adaxially septate-nodulose, basal 

sheaths brown, the inner band longitudinally 

yellow-brown medially, deeply U-shaped at 

orifice; ligule absent; blades broadly linear, 

flattened, 60-130 cm × 7-15 mm, thickly 

coriaceous, antrorsely serrulate-scabrous on 

margins and abaxial midvein, attenuate to 

triquetrous, caudate apex. Inflorescence a terminal 

and series of 3-7 lateral corymbs from the upper 

leaf-like bracts, these remote proximally, 

subcontiguous towards apex; corymbs dense, (2-) 

3.5-9 (-14) cm diam., branches flattened, the 

spikelets congested in globose or fasciculate heads 

of 3-12 at branch tips, 4-9 mm in diam; spikelets 

ovoid-ellipsoid to broadly so, 3-4.7 × 1-2 mm, with 

6-8 scales; fertile scales broadly ovate or ovateelliptic, 

membranous, rusty brown, midcosta 1nerved, 

pale, inconspicuous, not prolonged or only 

very shortly so beyond the obtuse to subacute apex; 

sterile scales ovate to broadly ovate, otherwise like 

the fertile, reducing in size towards base of 

spikelet. Anthers 2-3.5 mm long, with a subulate, 

black apiculum. Achene ovoid-globose, 2.5-3.2 × 

1.2-1.7 mm, longitudinally wrinkled when dry, 

long-beaked at apex, sub-rounded at base, 

estipitate, brown. 

General distribution: Southern United States, 

Mexico, Central America, West Indies, and South 

America. 

Distribution in Puerto Rico: Primarily in 

coastal areas; growing in lakes, swales, and 

marshes; also collected in a quarry near Santurce 

(San Juan). Arecibo, Cataño, Cidra, Humacao, 

Manatí, Mayagüez, Río Grande, San Juan, Vega 

Baja, and Yauco. 

Common names: Puerto Rico: Cortadora de 

ciénaga, Serrucho. 


Arecibo: Bo. Islote, Caño Tiburones, Axelrod et 

al. 10514 (UPRRP, US). Cataño: in paludosis, 

Sintenis 1214 (US); Puerto Salinas, González-Más 

2127 (US). Cidra: Pueblo Viejo, Hioram 101 (US). 

Humacao: Playa of Humacao: “Pozal”, Eggers 681 

(US). Manatí: circa Lagunam Tortuguero, Sintenis 

6814 (US). San Juan: Santurce, 2 mi. E of stone 

quarry, Heller & Heller 1274 (US); Martín Peña, 

Stevenson 1876 (US). Vega Baja: Bo. Algarrobo, 

along open shore at E end of Laguna Tortuguero, 

Proctor 45651 (US). Yauco: Sargent 681 (US).

252 


Fig. 47. A-F. Carex polystachya. A. Habit. B. Inflorescence spike. C. Spikelet scales, dorsal view and lateral view. D. Terminal 

staminate portion of spike. E. Perigynia. F. Achene. G-L. Cladium jamaicense. G. Upper portion of habit with inflorescence. H. 

Detail of leaf blade. I. Inflorescence branch. J. Flower. K. Spikelet scale. L. Achene. (A-F, from Proctor 48897; G, from Proctor 

45651; H, from Proctor 45651; I-L, from Proctor 43811).

Cyperus L., Sp. Pl. 44. 1753. 


5. CYPERUS 

Perennial or annual glabrous herbs; rhizomes, when present, short or stoloniferous, rarely elongate 

and horizontally creeping. Culms trigonous or sometimes terete, smooth or scabrous, green. Leaves 

primarily basal with several often lower cauline; sheaths, at least the uppermost, rarely bladeless, finely 

veined, sometimes cross-veined, usually glabrous, the ventral (inner) band membranous, often reddish 

dotted, with a truncate to U- or V-shaped orifice; ligule absent or short; blades herbaceous or stiffened, 

flattened, V-shaped, plicate, inrolled, terete, or crescentiform, finely veined abaxially, finely cellularreticulate 

and semi-glossy adaxially, sometimes septate or cross-veined, the margins, abaxial midvein, 

and adaxial lateral veins, when present, usually antrorsely scabrous, green. Inflorescence a simple or 

compound, terminal, umbel-like corymb, rarely congested or pseudolateral and head-like; involucral 

bracts generally leaf-like, approximate, spreading in a radius, the lowermost one longest, the upper 

successively shorter, or the lowest erect and appearing as a continuation of the culm, green; rays generally 

unequal in length, finely ribbed, trigonous, compressed-trigonous, or obscurely so, glabrous, rarely 

scabrous, prophyllate at base, the lowermost each borne from the axil of a single involucral bract, the 

uppermost borne just below the base of the central spike, these frequently flexuous, becoming divergent 

to reflexed at maturity; spikes solitary or subumbellate at ray tips, the central spike usually sessile or 

subsessile; spikelets ovate, ovate-lanceolate, or linear in shape, flattened, compressed or subcompressed, 

subcylindrical, or 4- angled in cross section, palmately, pinnately, or imbricately arranged on the rachis, 

many-flowered, the lowermost scale and subtending bract empty; scales 2-ranked, ovate, oblong-ovate 

or elliptic, acutely keeled, boat-shaped, or sub-rounded in cross section, sometimes 2-keeled, often 

longitudinally veined on sides, with a 1- to 5-nerved often greenish carina, usually glabrous, the apex 

acute, obtuse, or cuspidate, often mucronulate; rachilla winged from the persistent, decurrent base of 

the scale, or wingless, the wings hyaline or sometimes corky or thickened, disarticulating at base from 

the rachis, the whole spikelet falling entire, or disarticulating at the nodes, the internodes, scales, and 

achenes falling as 1-fruited segments, or rachilla persistent on the rachis, the scales and achenes deciduous. 

Flowers bisexual; hypogynous squamellae or bristles absent; stamens 1-3, the anthers elliptic, oblongelliptic, 

lanceolate or linear, apiculate or sometimes minutely appendaged at apex; styles 2- or 3-branched, 

slender, uniform, the branches minutely scaly, shorter than to exceeding the smooth unbranched portion. 

Achene trigonous or lenticular, ovoid, obovoid, oblong-obovoid, ellipsoid, or narrowly-ellipsoid, usually 

short-apiculate at apex, sometimes shortly stipitate at base, the surface puncticulate, reticulate, or smooth, 

rarely transversely wrinkled. A cosmopolitan genus with approximately 650 species, in temperate, 

subtropical, and tropical regions. 

LECTOTYPE: Cyperus esculentus L., designated by Britton, Bull. Dept. Agric. Jamaica 5, Suppl. 1: 6. 

1907. 

References: Kükenthal, G. 1935. Cyperaceae-Scirpoideae-Cypereae. In: A. Engler, Das Pflanzenreich 

IV. 20 (Heft 101): 1-671. McLaughlin, A. D. 1944. The genus Cyperus in the West Indies. Catholic 

Univ. Amer. Biol. Stud. 5: i-viii, 1-108. Tucker, G.C. 1994. Revision of the Mexican species of Cyperus 

(Cyperaceae). Syst. Bot. Monogr. 43: 1-213. 

Key to the species of Cyperus 

1. Style 2-branched; achenes lenticular or biconvex, generally borne with an edge, or in C. laevigatus a 

face, against the rachilla............................................................................................................... 2 

2. Inflorescence a pseudolateral capitate head of loosely subdigitate spikelets at the summit of the 

culm; involucral bracts 1, culm-like, erect, appearing as a continuation of the culm; spikelets 1 

mm thick; achene borne with a face against the rachilla ..............23. C. laevigatus 

2. Inflorescence a terminal, open or contracted, umbel-like corymb with ascending rays; involucral 

bracts (1-) 2-7, leaf-like, ascending to spreading; spikelets 0.5-0.6 mm thick; achene borne 

with an edge against the rachilla...................................................................... 3

254 


3. Achene pericarp with rectangular, longitudinally elongate cells forming transverse 

undulations.............................................................................................. 14. C. flavescens 

3. Achene pericarp with isodiametric cells, not forming transverse undulations.................. 4 

4. Spikelets linear-lanceolate to linear, 1-1.6 (-2) mm wide; scales (1.2-) 1.4-1.8 (-2) × 

0.8-1.5 mm; anthers 0.4-0.6 mm long; achenes 0.8-1 (-1.2) × 0.4-0.6 mm 

......................................................................................................31. C. polystachyos 

4. Spikelets lanceolate to ovate-lanceolate or oblong-lanceolate, 3-5 mm wide; scales 3- 

5 × 2-2.6 mm; anthers 2-2.8 mm long; achenes 1.1-1.3 × 0.8-1.1 mm 

..........................................................................................................41. C. unioloides 

1. Style 3-branched; achenes trigonous or sometimes dorsiventrally compressed, borne with a face against 

the rachilla.......................................................................................................................... 5 

5. Leaves reduced to bladeless sheaths........................................................................................ 6 

6. Culm with transverse septa ...................................................................... 2. C. articulatus 

6. Culm without transverse septa.......................................................................................... 7 

7. Culms triquetrous to nearly 3-winged, soft and friable, easily compressed, (1-) 1.8-5 

(-5.5) mm wide, sheathing bases 3-8 mm wide; involucral bracts 2 (-3) 

.............................................................................................................. 19. C. haspan 

7. Culms trigonous to obtusely trigonous or subterete, firm, 3-30 mm wide, sheathing 

bases 7-50 mm wide; involucral bracts 6 or more.................................................... 8 

8. Primary involucral bracts 6-8, up to 10 cm long; rays 100-200............ 29. C. papyrus 

8. Primary involucral bracts (6-) 8-25, 15-45 cm long; rays 35 or fewer............... 9 

9. Culms (6-) 8-30 mm wide; involucral bracts of varying lengths, gradually and 

narrowly acuminate at apex; spikelets linear-lanceolate, 0.8-1.2 mm wide; 

scales ovate-elliptic, carina 5- to 7-nerved; achenes 0.9-1.2 mm long; rachilla 

unwinged (wings deciduous) ................................ 18. C. giganteus 

9. Culms 3-9 mm wide; involucral bracts subequal in length, abruptly acute to 

acuminate at apex; spikelets ovate to lanceolate or oblong-lanceolate, 1.8- 

2.2 mm wide; scales ovate-deltate; carina 3-nerved, achene 0.6-0.8 mm long; 

rachilla hyaline-winged ....................................... 21. C. involucratus 

5. Leaves, at least the uppermost on culm, with well-developed blades. .................................. 10 

10. Spikelet scales dorsally 2-keeled or invaginated proximally. ....................................... 11 

11. Sheaths, leaves, and involucral bracts conspicuously cross-veined ...... 43. C. virens 

11. Sheaths, leaves, and involucral bracts not conspicuously cross-veined................. 12 

12. Culms retrorsely scabrid, at least distally; spikelets 1.5-2 mm wide; scales 1.2- 

1.5 × 0.7-0.9 mm; achenes 0.8-0.9 × 0.3 mm .................... 37. C. surinamensis 

12. Culms smooth; spikelets 2-3 mm wide; scales (1.2-) 1.5-2 × 1.2-2 mm; achenes 

1-1.5 × 0.4-0.7 mm .................................................... 28. C. ochraceus 

10. Spikelet scales dorsally 1-keeled from apex to base, not invaginated proximally........ 13 

13. Rachilla disarticulating into 1-fruited segments (the rachilla node and achene falling 

together), the wings becoming thick and corky at maturity, closely enveloping the 

achene; spikelets cylindrical and often flexuose............................ 14 

14. Leaf blades 1-2.3 mm wide; inflorescence a terminal corymb or reduced to 2-6 

subfastigate spikelets, 1.2-5 × 0.7-3 cm; involucral bracts 1-3, the lowest erect, 

leaf-like but appearing as a continuation of the culm ............... 13. C. filiformis 

14. Leaf blades 3.5-17 mm wide; inflorescence an umbel-like corymb with ascending 

rays, (1.5-) 2.5-32 (-40) cm in diam.; involucral bracts 4-15, ascending to spreading. 

.................................................................................. 15 

15. Inflorescence of densely congested, lobate heads of spikes at ray tips or a 

single lobate head at summit of the culm .............................. 10. C. eggersii 

15. Inflorescence of oblong or widely ovoid to subglobose spikes with laxly or 

sublaxly disposed spikelets at ray tips...................................................... 16


16. Internodes of the spike rachis between spikelets less than 1 mm long; spikelets spicate or 

sometimes radiate, somewhat congested, the lower ones reflexed at maturity; scales reddish 

brown to dark brown on sides 15........................................................................... C. flexuosus 

16. Internodes of the spike rachis between spikelets 1-5 mm wide; spikelets never radiate, rarely 

reflexed; scales light yellowish brown on sides................................................ 28. C. odoratus 

13. Rachilla persistent (the scales and achenes deciduous) or disarticulating at base of spikelet (the 

spikelet falling entire), the wings hyaline and not becoming thick and corky nor closely enveloping 

achene, or absent; spikelets laterally compressed or subquadrate. ............................. 17 

17. Inflorescence rays wanting or not evident; spikes sessile or essentially so, subtended by leaflike 

involucral bracts, solitary or arranged in lobate or digitate clusters at the summit of the 

culm and bearing 2-many, subloosely, densely congested, or radiately spreading spikelets..... 

.....................................................................................................................................................18 

18. Plants often with polymorphic inflorescences, these ranging from (2-) 4-15 (-24) cm diam., 

grading from contracted and head-like (rays not evident) to open corymbs with welldeveloped 

rays. ...................................................................................................... 19 

19. Spikes cylindrical with densely disposed spikelets, the rachis not clearly visible between 

the spikelets, not flattening when pressed and dried; florets 1-2 (-4) per spikelet 

............................................................................................. 1. C. aggregatus 

19. Spikes ovoid to oblong-ovoid with subloosely disposed spikelets, the rachis clearly 

visible between the spikelets, flattening when pressed and dried; florets (2-) 4-38 

(-44) per spikelet. ................................................................................................... 20 

20. Spikelets 0.7-1.1 mm wide; anthers 0.3-0.5 mm long; achenes linear to 

narrowly oblong, 0.4-0.5 mm wide ......................................... 39. C. tenuis 

20. Spikelets 1.4-3 mm wide; anthers 0.6-1.3 mm long; achenes obovoid, broadly 

obovoid, or ellipsoid-obovoid to oblong-obovoid, 0.6-1.1 mm 

wide.......................................................................................................... 21 

21. Leaf blades 1.5-3 mm wide; rachilla of spikelet without persistent hyalinewings 

(wings caducous); carina at scale apex prolonged as a short, straight 

to slightly divergent awn, 0.5-1 mm long 

.................................................................................... 4. C. compressus 

21. Leaf blades 2-12 (-15) mm wide; rachilla of spikelet with persistent 

hyaline-wings; carina at scale apex scarcely prolonged, merely shortmucronate......................................................................................... 

22 

22. Scales brownish, 2-2.8 mm wide, the nerves coarse, evenly spaced, 

extending down to the scarious margins; achene obovoid to ellipsoidobovoid, 

plane on the adaxial side, 0.8-1.1 mm wide 

................................................................................. 3. C. brunneus 

22. Scales reddish, 1.4-1.8 (-2) mm wide, the nerves, crowded towards 

the carina, not extending down to scarious margins; achene ellipsoidobovoid 

to oblong-obovoid, slightly curvate on the adaxial side, 0.6- 

0.9 mm wide ......................................................... 30. C. planifolius 

18. Plants with small, uniform inflorescences, 0.3-1.5 cm in diam. ................................... 23 

23. Spikelet scales concavely curvate-keeled.............................................................. 24 

24. Culms scabid at apex; inflorescence globose or hemispherical; rachilla strongly 

flexuose, the mature spikelet scales and achenes appressed and conforming closely 

to it, not spreading; achene narrowly ellipsoid to ellipsoid-obovoid; scales widely 

ovate-elliptic, 1.4-2 mm wide; frequent, throughout Puerto Rico and the Virgin 

Islands ................................................................. 40. C. unifolius 

24. Culms smooth at apex; inflorescence obtriangular-hemispherical; rachilla weakly 

flexuose, the mature spikelet scales and achenes spreading away from it; achene 

linear-oblong; scales ovate elliptic, 0.8-1.2 mm wide; rare, only known from Puerto 

Rico on Mona Island and in Guánica ................................ 16. C. floridanus

256 


23. Spikelet scales convexly curvate-keeled or straight-keeled................................... 25 

25. Terminal spikelet scale dorsally gibbous or pouched, sterile....... 38. C. swartzii 

25. Terminal spikelet scale not dorsally gibbous or pouched, fertile. ................... 26 

26. Involucral bracts strongly deflexed before disarticulation of spikelets from 

rachis, rarely the uppermost one slightly ascending; spikelets strongly 

deflexed, the terminal one often erect. ..................................................... 27 

27. Spikelet scales 1.2-1.6 × 0.8-1 mm, finely 3- to 4-nerved; achenes oblongellipsoid, 

1-1.2 × 0.5-0.6 mm, straight on adaxial side.......... 26. C. nanus 

27. Spikelet scales 1.5-2 × 1.2-1.6 mm, coarsely 4- to 5-nerved; achenes 

ovoid-ellipsoid, 1.1-1.4 × 0.5-0.7 mm, curvate on adaxial side 

...................................................................................... 36. C. subtenuis 

26. Involucral bracts ascending to divergent, rarely slightly deflexed; spikelets 

ascending, radiately or hemispherically spreading, or rarely the lowermost 

several weakly deflexed........................................................................... 28 

28. Apex of spikelet scales excurved-cuspidate, the sides often whitish 

stramineous with reddish or blackish lineations; margins and abaxial 

midvein of leaf blades often with both antrorsely and retrorsely set barbs 

............................................................................... 42. C. urbanii 

28. Apex of spikelet scale straight, the sides yellowish or reddish brown or 

dark brown to purple-black; margins and abaxial midvein of leaf blades 

with antrorsely set barbs only................................................. 29 

29. Leaf blades (1-) 1.3-3 (-3.5) mm wide; inflorescence rounded or 

hemispherical; spikelets radiately spreading; scales straight-keeled 

to convexly curvate-keeled; achenes ellipsoid-obovoid to oblongobovoid, 

1-1.3 mm long .......................... 17. C. fuligineus 

29. Leaf blades 0.2-1.2 mm wide; inflorescence obtriangularhemispherical; 

spikelets ascending; scales often concavely curvatekeeled; 

achenes linear-oblong, 1.3-1.6 mm long ...... 15. C. floridanus 

17. Inflorescence with elongated rays (at least some spikes borne on elongated primary rays); spikes 

arranged in open to contracted, simple or compound, umbel-like corymbs with ascending rays... 30 

30. Terminal spikelet scale dorsally gibbous or pouched, sterile ........................... 38. C. swartzii 

30. Terminal spikelet scale not dorsally gibbous or pouched, fertile......................................... 31 

31. Culms, leaves, rays, and involucral bracts finely papillose....................... 24. C. ligularis 

31. Culms, leaves, rays, and involucral bracts not finely papillose.................................... 32 

32. Spikelet scales (at least some) with a dark purplish red splotch along lower margin on 

each side ................................................................................... 34. C. sphacelatus 

32. Spikelet scales lacking a dark purplish red splotch along lower margin on each side. 

................................................................................................................................. 33 

33. Apex of spikelet scale with an elongate, strongly recurved awn...................... 

................................................................................................ 35. C. squarrosus 

33. Apex of spikelet scale straight or recurved-cuspidate.................................... 34 

34. Spikelets arranged in narrowly to widely linear or cylindrical spikes; spikes 

digitately arranged at ray tips or in a simple digitate cluster at the summit of 

the culm .................................................................................. 35 

35. Inflorescence a simple, digitate cluster of spikes at the summit of the 

culm, essentially sessile or sometimes 1-2 short rays present............... 

..................................................................................... 1. C. aggregatus 

35. Inflorescence an open, compound corymb with ascending rays....... 36 

36. Spikelets oriented at right angles to the axis of the spike............. 37


37. Spikelets with 1-3 (-5) florets, oblong-ellipsoid, slightly compressed to 

subquadrate, or broadly elliptic; scales 2.2-3.4 × 1.5-2.1 mm, coarsely (3-) 4-5 

nerved on each side, light brownish to brown or reddish brown, dark brown 

lineolate, at least on margins ......................................................... 25. C. mutisii 

37. Spikelets with 12-30 florets, linear, compressed-rhombic; scales 1.4-2 × 1-1.3 

mm, laterally weakly 1- to 2-nerved along carina, light to reddish brown on sides, 

with 2 dark brown stripes paralleling either side of carina ........... 8. C. digitatus 

36. Spikelets strongly ascending to appressed to the axis of the spike....................... 38 

38. Caespitose annual or short-lived perennial, (5-) 12-60 (-75) cm tall; sheaths pale 

green to pale brown, the lowermost reddish tinged, reddish lineolate; leaf blades 

(1.5-) 2-6 mm wide, both the blades and involucral bracts not cross-veined 

abaxially or indistinctly so ...................................................... 22. C. iria 

38. Coarse, rhizomatous perennial, 70-150 cm tall; sheaths purple-black proximately, 

fading to brown streaked with black distally; leaf blades 4-15 (-18) mm wide, 

both the blades and involucral bracts cross-veined abaxially 

................................................................................................. 20. C. imbricatus 

34. Spikelets arranged in globose, subglobose, or ovoid spikes; spikes solitary, spicately, or digitately 

arranged at ray tips........................................................................................................................ 39 

39. Spikelets digitately arranged, hence spikes without a conspicuous rachis.......................... 40 

40. Plant with sticky leaves and inflorescence when fresh; culms remotely scabrous; spikelets 

(2-) 3-5 (-7) mm wide; scales 2.2-3.2 × 2-2.8 mm, with a slightly excurved-cuspidate or 

straight apex; achenes 1.3-1.9 × 0.7-1 mm ............................ 11. C. elegans 

40. Plants not sticky; culms smooth; spikelets ranging from 0.8-1.6 (-2) mm wide; scales 

ranging from 0.6-1.9 × 0.6-1 mm, with a straight, mucronate apex; achenes ranging from 

0.5-0.8 × 0.3-0.5 mm. .......................................................................................... 41 

41. Caespitose annual; spikes in dense, globose, often irregularly lobate clusters at ray 

tips, each with 20-140 spikelets; scales obovate to orbiculate, 0.6-0.8 mm long 

.............................................................................................................. 7. C. difformis 

41. Rhizomatous perennial with short, horizontally creeping rhizome; spikes in 

hemispherical or rounded, open clusters at ray tips, each with (1-) 3-13 spikelets; 

scales oblong-ovate, 1.4-1.9 mm long ................................................. 19. C. haspan 

39. Spikelets spicately arranged on a short to elongate spike rachis ........................................ 42 

42. Spikelets 0.5-0.6 mm wide; florets widely spreading to 1.8 mm wide with maturing achenes; 

spikelet scales remotely 2-ranked ................................................... 9. C. distans 

42. Spikelets 0.7-3 mm wide; florets not widely spreading; spikelet scales closely 2-ranked. 

.........................................................................................................................................43 

43. Apex of spikelet scales cuspidate, slightly excurved; achene apiculum bulbousthickened 

............................................................................................. 5. C. confertus 

43. Apex of spikelet scales mucronate or short-awned, essentially straight; achene apiculum 

not bulbous-thickened ........................................................................... 44 

44. Caespitose annuals (lacking rhizomes); hyaline rachilla wings deciduous, not 

persistent ........................................................................................................ 45 

45. Spikelets 2.5-3 mm wide; spikelet scales lacking a dark purplish red splotch 

along lower margin on each side .......................... 4. C. compressus 

45. Spikelets 1.2-1.8 (-2) mm wide; spikelet scales often with a dark purplish 

red splotch along lower margin on each side or sometimes absent............... 

........................................................................................ 34. C. sphacelatus 

44. Rhizomatous or caespitose perennials; hyaline rachilla wings persistent ...... 46 

46. Spikelets 0.7-1.3 mm wide ..................................................................... 47 

47. Inflorescence rays well-developed, often longer than the length of spikes 

borne at their tips; spikes subglobose .................... 6. C. croceus

258 


47. Inflorescence rays often shorter than the length of spikes borne at their tips or wanting; spikes 

loosely ovoid to oblong-ovoid ................................................................... 39. C. tenuis 

46. Spikelets (1.2-) 1.4-2.8 mm wide ................................................................................................ 48 

48. Rhizomes stoloniferous, bearing tubers .............................................................................. 49 

49. Involucral bracts elongate, the lowermost 2 generally exceeding the inflorescence, the 

lowest one 7-60 cm long; spikelet scales light brown or yellowish brown with 2-3 welldeveloped 

lateral nerves evident the length of the scale; eligulate ........ 12. C. esculentus 

49. Involucral bracts generally shorter than to equaling the length of the inflorescence, or only 

the lowest one shortly exceeding the inflorescence, 1-9 (-14) cm long; spikelet scales dark 

reddish brown to purple-brown or blackish with only 1-2 indistinct lateral nerves evident 

only at proximal base of scale, the sides smooth for the most part; faintly short-ligulate on 

lower sheaths ................................................................. 33. C. rotundus 

48. Rhizomes short, non-stoloniferous ..................................................................................... 50 

50. Spikelet scales uniformly light brown or light yellowish brown on sides with a conspicuous, 

elongate thickened mucro ............................................. 32. C. pulguerensis 

50. Spikelet scales often stained dark brown, reddish brown, or reddish on sides, with an 

inconspicuous, minute or very short mucro ................................................................... 51 

51. Scales brownish, 2-2.8 mm wide, the nerves coarse, evenly spaced, extending down 

to the scarious margins; achene obovoid to ellipsoid-obovoid, plane on the adaxial 

side, 0.8-1.1 mm wide ......................................................................... 3. C. brunneus 

51. Scales reddish, 1.4-1.8 (-2) mm wide, the nerves, crowded towards the carina, not 

extending down to scarious margins; achene ellipsoid-obovoid to oblong-obovoid, 

slightly curvate on the adaxial side, 0.6-0.9 mm wide .................... 30. C. planifolius 

1. Cyperus aggregatus (Willd.) Endl., Cat. Horti 

Vindob. 1: 93. 1842; Mariscus aggregatus 

Willd., Enum. Pl. 1: 70. 1809; Cypreus flavus 

var. aggregatus (Willd.) Kük. in Engler, 

Pflanzenr. IV. 20 (Heft 101): 532. 1936. Type: 

based on a cultivated plant (holotype: B- 

Willd. 1426). 

Kyllinga cayennensis Lam., Tabl. Encycl. 1: 49. 

1791; Cyperus cayennensis (Lam.) Britton, 

Bull. Dept. Agric. Jamaica 5: 8. 1907, non 

Willdenow ex Link, 1820; Mariscus 

cayennensis (Lam.) Urb., Symb. Antill. 2: 

165. 1900. Type: French Guiana. Von Rohr 

s.n. (holotype: P-Lam.). 

Mariscus flavus Vahl, Enum. Pl. 2: 374. 1805; 

Cyperus flavus (Vahl) Nees, Linnaea 19: 698. 

1847, non J. Presl & C. Presl, 1828. 

Lectotype: St. Croix, U.S. Virgin Islands. Von 

Rohr 70 (C-Vahl), designated by G. C. Tucker 

& McVaugh in McVaugh, Fl. Novo-Galiciana 

13: 279. 1993. 


rhizome short, decumbent, knotty. Culms loosely 

tufted, rigid, triquetrous distally, trigonous 

proximally, finely ribbed, often antrorsely 

spinulose-scabrous on angles distally, smooth 

proximally, (0.8-) 1-2.7 (-3.3) mm wide, sheathing 

bases 2-7 mm wide. Leaves 5-8, basal and lower 

cauline; sheaths purple-tinged or light red-brown 

proximally, green distally; ligule absent; blades 

linear, flattened-plicate or V-shaped proximally, 

9-60 cm × 2-7 mm, antrorsely spinulose-scabrous 

on the margins and abaxial midvein, attenuate to 

triquetrous apex. Inflorescence a simple umbellike 

corymb with few rays, often contracted to a 

radiate cluster of 2-8 (-10) sessile to subsessile 

spikes, 2-7 (-10) × 1.5-5 cm; involucral bracts 4- 

8, leaf-like, spreading, the lowermost to 30 cm 

long; rays essentially absent or 3-10 short ones, 

several sometimes to 5 cm long; spikes 6-20 (-30) 

× 5-8 (-10) mm, cylindrical to oblong-cylindrical 

with 20-80 densely disposed, spicately arranged 

spikelets, which are each subtended by a linearsetaceous 

bract at base that equals or exceeds the 

spikelet; spikelets spreading, elliptic to oblongelliptic, 

subquadrate, 2-3.5 × 0.9-1.3 mm, acute at 

apex, cuneate to short-cuneate at base, with 1-2 

(-4) florets; rachilla broadly hyaline-winged; 

scales 4-5, the lowermost two reduced and sterile; 

fertile scales ovate to ovate-elliptic or broadly 

ovate-elliptic, obtusely curvate-keeled, 2.3-3.2 × 

1.8-2.6 mm, submembranous, with 3-4 coarse 

nerves on each side above the broad margins which


often clasp the mature achenes, narrowly scarious, 

stramineous to yellowish brown, often dark red 

lineolate on sides, semi-glossy, carina 3-nerved, 

green, shortly prolonged beyond the obtuse apex 

as a mucro. Stamens 3, the anthers 0.4-0.7 mm 

long, bluntly apiculate; styles 3-branched. Achene 

trigonous with concave sides, obovate to ellipticobovate 

or broadly ellipsoid, 1.5-1.9 (-2.1) × 0.8- 

1.1 mm, obtuse to rounded at apex, apiculate, 

short-cuneate at base, estipitate, essentially smooth 

to finely puncticulate, dark brown to reddish brown 

at maturity. 


(Louisiana, west to New Mexico), West Indies, 

Mexico, Central America, tropical and subtropical 

South America; adventive in Australia. 


Islands: In sandy soils of a variety of habitats 

including road banks, rocky woods and hillsides, 

marshy areas, stream margins, savannas, cultivated 

fields, lawns, pastures, and waste areas. Adjuntas, 

Bayamón, Cabo Rojo, Coamo, Dorado, Guayama, 

Humacao, Jayuya, Juncos, Loíza, Manatí, 

Maricao, Mayagüez, Moca, Peñuelas, Río Piedras, 

Salinas, Sabana Grande, San Germán, San Juan, 

San Sabastián, Utuado, and Vega Baja; Buck 

Island, St. Croix, and St. Thomas. 


Adjuntas, Alto de la Bandera, N.L. Britton & Hess 

2137 (US). Cabo Rojo: Sierra Bermeja, Proctor 

& McKenzie 44021 (US). Dorado: Bo. Higuillar, 

just E of Aeropuerto de Dorado, Proctor 42454 

(US). Jayuya, Sargent 3093 (US). Juncos: in Mt. 

Florida, Sintenis 1904 (US). Loíza, Stevenson 5480 

(US). Manatí: Bo. Tierras Nuevas Saliente, Proctor 

& Thomas 43802 (US). Moca: Sargent 402 (US). 

Salinas: Sabana Llaña, Stevenson 1256 (US). San 

Juan: Along the beach at Santurce, Heller 4683 

(US). Utuado: vicinity of Utuado, E.G. Britton 

5213 (US). Vega Baja: Tortuguero, Rd. 687, km 

2, González-Más 1047 (US). ST. CROIX: Bassin, 

A.E. Ricksecker 483 (US). 

2. Cyperus articulatus L., Sp. Pl. 44. 1753. 

Lectotype: Jamaica. Sloane, Voy. Jamaica 1: 

t 81, f. 1. 1707, designated by G. C. Tucker, 

Syst. Bot. Monogr. 2: 42. 1983. 

Rhizomatous perennial, (50-) 70-260 (-300) 

cm tall; rhizome elongate, horizontally creeping, 

hardened, scaly, 2-8 mm thick, often tuberous- 

thickened below base of culm. Culms forming in 

a row along the rhizome, approximate or widely 

spaced, terete to obtusely trigonous distally, 

tapering to apex, with transverse septa, essentially 

smooth, 2-14 mm wide, sheathing bases 4-18 mm 

wide. Leaves reduced to bladeless sheaths or 

occasionally 1-2 (-4) blades present, basal and 

lower cauline; sheaths prominently cross-veined, 

tinged with purple or purple-brown proximally, 

becoming fibrillose, the bladeless ones oblique at 

orifice, dorsally acute to acuminate; ligule absent; 

blades when present, linear-lanceolate, flattenedplicate 

or V-shaped, 9-20 (-40) cm × 3-6 (-9) mm, 

prominently cross-veined, antrorsely scabrous on 

margins, -short-attenuate to triquetrous apex. 

Inflorescence an open, compound corymb, 4-15 

(-18) × 3-11 (-16) cm; involucral bracts 2-3, leaflike, 

erect, 1-2 (-10) cm long, often inconspicuous, 

shorter than the inflorescence; primary rays 5-7, 

to 10 cm long, wire-like, erect to curving; 

secondary rays sometimes present, to 20 mm long; 

spikes 4-30, broadly ovoid to obloid, 1-4.5 × 1- 

3.5 (-4.5) cm, with (1-) 4-10 spikelets; spikelets 

linear 8-32 (-55) × 1.1-2 mm, compressed, tapering 

to acuminate apex, cuneate at base, with 10-43 

(-60) florets; rachilla narrowly to broadly hyalinewinged 

at maturity; scales oblong-elliptic to ovate, 

acutely curvate-keeled, 2.6-3.7 × 1.1-1.8 mm, 

submembranous, with 1-2 nerves on each side 

above the scarious margins, light brown to brown 

on sides, carina 3-nerved, green, narrowed to a 

short subulate tip just below the acute to obtuse 

apex. Stamens 3, the anthers 1.3-2.4 mm long, 

bluntly apiculate; styles 3-branched. Achene 3sided 

with concave faces, slightly dorsiventrally 

compressed, obovoid-ellipsoid, 1.2-1.6 × 0.4-0.6 

mm, rounded to markedly apiculate apex, shortcuneate 

at base, substipitate, puncticulate, brown 

to dark brown at maturity. 

General distribution: Pantropical and warmtemperate 

regions; southeastern United States 

(South Carolina to Texas), West Indies, Mexico, 

Central America, South America; and Old World 

tropics. 


Islands: In wet areas of low swampy grounds, 

marshes, lake borders, pastures, and roadside 

ditches. Cabo Rojo, Caguas, Carolina, Fajardo, 

Guánica, Gurabo, Humacao, Juana Díaz, Juncos, 

Lajas, Loíza, Naguabo, Santa Isabel, Vega Baja, 

and Yabucoa; St. Croix, St. Thomas, and Tortola.

260 


Common name: Puerto Rico: Junco cimarrón. 


Cabo Rojo: Bo. Llanos Costa, Proctor & McKenzie 

43770 (US). Caguas: Sintenis 2549 (US). Carolina: 

Canovanillas, Hwy 165, km 14.9, González-Más 

129 (US). Guánica: West end of Lake Guánica, 

Sargent 578 (US). Gurabo: Sintenis 2548 (US). 

Humacao: Playa de Humacao, Eggers 670 (US). 

Juncos: Sintenis 2503 (US). Lajas: Laguna 

Cartagena, Liogier et al. 30568 (US). Loíza: Vacía 

Talega, Woodbury et al. s.n. (US); Loíza road, 

Stevenson 5467 (US). Naguabo: N.L. Britton & 

Hess 2319 (US); Sierra de Naguabo, Loma Icaco, 

Shafer 3416 (US). Vega Baja: Pto. Nuevo, Rd. 686, 

km 3.4, González-Más 1109 (US). Yabucoa: 

Sintenis 5119 (US). 

3. Cyperus brunneus Sw., Fl. Ind. Occid. 1: 116. 

1797; Mariscus brunneus (Sw.) C. B. Clarke 

in Urban, Symb. Antill. 2: 51. 1900; Cyperus 

planifolius var. brunneus (Sw.) Kük., Repert. 

Spec. Nov. Regni Veg. 23: 189. 1926. Type: 

West Indies. Swartz s.n. (holotype: S-Sw. no. 

R-1383; isotype: M). 

Cyperus brizaeus Vahl, Enum. Pl. 2: 316. 1805; 

Mariscus brizaeus (Vahl) C. B. Clarke in 

Urban, Symb. Antill. 2: 52. 1900. Lectotype: 

French Guiana; Cayenne. Richard s.n. (C- 

Vahl), designated by G. C. Tucker, Syst. Bot. 

Monogr. 43: 145. 1994. 

Cyperus evaginatus Boeck., Linnaea 36: 351. 

1870. Type: United States; Florida. Collector 

unknown (holotype: B, destroyed). 

Cyperus ottonis Boeck., Linnaea 36: 350. 1870. 

Type: Cuba. Otto s.n. (holotype: B, 

destroyed). 

Cyperus discolor Boeck., Beitr. Cyper. 1: 7. 1888. 

Type: St. Thomas, U.S. Virgin Islands. Kuntze 

s.n. (holotype: B, destroyed). 

Cyperus krugii Boeck., Beitr. Cyper. 1: 8. 1888. 

Type: Puerto Rico; Bayamón. Sintenis s.n. 

[collection number not designated in 

protologue] (holotype: B, destroyed). 

Rhizomatous perennial, (10-) 15-80 cm tall; 

rhizome short, thickened. Culms trigonous to 

obtusely so proximally, 1.2-4 mm wide, coarsely 

ribbed, smooth, sheathing bases 6-15 mm wide. 

Leaves 5-13, crowded at base, 1-2 lower cauline; 

sheaths short, red-brown to purple-black 

proximally; ligule absent; blades narrowly linear, 

somewhat stiff, flattened-plicate, 10-90 cm × 2-9 

mm, the lowermost often equaling or overtopping 

culms, pale green to green, faintly reddish brown 

lineolate on adaxial surface, whitish green 

abaxially, indistinctly cross-veined proximally, 

antrorsely scabrous on margins and midvein 

beneath, long-acuminate to triquetrous apex. 

Inflorescence a simple to partially compound 

umbel-like corymb, open, or more often congested 

in one head-like lobed cluster, 2-8 (-13) cm in 

diam.; involucral bracts 3-5, leaf-like, elongate, 

the lowest to 60 cm long, the upper linear, subulate; 

rays (when evident) 2-7, 1-7 cm long; spikes 

broadly ovoid, 2-4 cm diam., with 12-38 (-50) 

spikelets; spikelets lanceolate or elliptic-lanceolate 

to oblong-lanceolate, (6-) 8-20 × 2-2.8 mm, 

divergent, subcompressed, abruptly acute or 

obtuse at apex, cuneate at base, with 6-26 florets; 

rachilla thickened, broadly hyaline-winged; scales 

broadly ovate to oval, obtusely keeled in cross 

section, 2.4-3.2 × 1.5-2.2 mm, chartaceous to 

thinly coriaceus, coarsely 4-6 nerved on each side 

above the narrow scarious margins, light brown 

and stained with red-brown or dark yellowish 

brown, carina 3-nerved, narrow, inconspicuous, 

pale-green, prolonged at the acute to obtuse apex 

as a short mucro. Stamens 3, the anthers 1-1.3 mm 

long, with a minute, rounded, black appendage at 

apex; styles 3-branched. Achene trigonous with 

adaxial face plane to slightly concave, abaxial 

faces slightly convex, subdorsiventrally 

compressed, obovoid to ellipsoid-obovoid, 1.2-1.8 

× 0.8-1.1 mm, obtuse at apex, apiculate, cuneate 

at base, estipitate, puncticulate, blackish at 

maturity. 


(southern Florida), coastal areas and cays of 

eastern Mexico and Central America, and the West 

Indies. 


Islands: In sand or sandy soils of coastal habitats 

including borders of salt ponds, dunes, ecotones 

between beach and scrub, scrub, coastal thickets, 

limestone slopes and flats, strand vegetation, 

roadsides, and disturbed areas. Adjuntas, 

Aguadilla, Arecibo, Cabo Rojo, Carolina, Cayo 

Ahogado, Cayo Diablo, Cayo Icacos (Ratones), 

Cayo Lobos, Cayo Palominos, Cayo Ramos, 

Desecheo, Fajardo, Guánica, Guayama, 

Guayanilla, Hatillo, Humacao, Isabela, Isleta 

Marina, Lajas, Loíza, Manatí, Mona Island, Ponce,


Quebradillas, Rincón, Río Grande, Salinas, San 

Juan, Vega Baja, Vieques, and Yauco; Anegada, 

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

Island. 


Adjuntas: Sintenis 4664 (US). Arecibo: Sargent 

767 (US); Bo. Islote, Punta Caracoles, Proctor & 

Rivera 46277 (US). Cabo Rojo: Rt. 102, Joyudas, 

González-Más 628 (US). Carolina: Boca de 

Cangrejos, González-Más 1029 (US); Punta 

Cangrejos, Stevenson & Johnston 1707 (US).Cayo 

Diablo, La Cordillera Cays, Fosberg 52282 (US). 

Fajardo: Fajardo Lighthouse, Heller & Heller 1168 

(US). Guánica: below Guánica State Forest, Strong 

et al. 441 (US). Humacao: Playa Humacao, 

González-Más 93 (US). Isleta Marina (Obispo), 

Woodbury et al. s.n. (US). Lajas: La Parguera, 

Sargent 43 (US). Manatí, Sintenis 6690 (US). 

Mona Island: Sardinera, Otero & Chardón 982 

(US). Ponce: Rt. 2, km 16.6, Los Cuchares, 

González-Más 798 (US). Quebradillas: Bo. San 

José, Axelrod 10811 (UPRRP, US). Rincón, ad 

Puntas, Sintenis 5694 (US). Salinas: Bo. Aguirre, 

Jobos Bay, Proctor & Delgado 46362 (US). San 

Juan: Park of Santurce, Hioram 811 (US); San 

Juan, Underwood & Griggs 915 (US). Vega Baja: 

Bo. Cibuco, Proctor et al. 45555 (US). Vieques 

Island: Santa Maria to Caballo, Shafer 2685 (US). 

ANEGADA: West End, Britton & Fishlock 982 (US). 

ST. CROIX: Sandy Point, Acevedo-Rdgz. et al. 5309 

(NY, US); Pinetree Bay, Ricksecker 334 (US). ST. 

THOMAS: Kaornsight, Eggers s.n. (US). 

4. Cyperus compressus L., Sp. Pl. 46. 1753. 


t. 76, f. 1. 1707, designated by Du Puy et al. 

in A.S. George et al. (eds.), Fl. Australia 50 

(Oceanic Islands 2): 446. 1993. 

Cyperus compressus var. capillaceus C. B. Clarke 

in Urban, Symb. Antill. 2: 32. 1900. Type: 

Dominica. Imray 430 (holotype: probably at 

K). 

Densely tufted annual, (3-) 5-40 (-60) cm tall; 

roots many, fine, capillary. Culms 3-angled, finely 

ribbed, smooth, 0.8-1.7 (-2.0) mm wide. Leaves 

2-7 primarily basal, often 1-2 lower cauline; 

sheaths red-tinged to purple-brown; ligule absent; 

blades folded to subflattened, frequently curved, 

3-30 cm × 1.5-3 mm, smooth to remotely 

antrorsely scabrous on margins, acuminate to 

triquetrous apex. Inflorescence an umbel-like 

corymb, 5-15 (-24) cm diam., or a sessile or 

subsessile, solitary spike; involucral bracts 3-5, 

2.5-18 cm × 1.2-3.0 mm, leaf-like; rays 0-5, 

unequal, 3-angled to subcompressed, spreading, 

to 9 cm long, 0.5-0.9 mm wide, ribbed, smooth; 

spikes hemispherical to rhomboid-obovoid, 1.1- 

3.5 × 1.2-5.0 cm, with 3-18 spikelets; spikelets 

approximate, oblong-elliptic to lance-oblong, 

spreading, 8-25 × 2.5-3.0 mm, subcompressed, 

acute at apex, short-cuneate at base, with (9-) 12- 

38 (-44) florets; rachilla flat, hyaline-winged, the 

wings caducous; scales deciduous, ovate to 

broadly ovate, acutely keeled dorsally, 2.6-3.5 × 

2.0-2.5 (-3.0) mm, herbaceous, 4- to 5-nerved on 

each side above the whitish scarious margins, the 

sides green or light brown, sometimes tinged with 

yellow, carina broad, green, 3-nerved, prolonged 

beyond the acute to acuminate apex as a stiff, welldeveloped, 

sharp-tipped mucro, 0.5-0.7 mm long. 

Stamens 3, the anthers 0.6-0.9 mm long, with a 

prolonged lanceolate apex; style 3-branched. 

Achene trigonous with concave sides, 1.1-1.5 × 

1-1.1 mm, obovoid to broadly obovoid, obtuse to 

3-lobed at apex, minutely apiculate, short-cuneate 

at base, brown, shiny, minutely reticulate, 

appearing smooth, obscurely ribbed on the angles. 

General distribution: Cosmopolitan; although 

more common in warmer regions. 


Islands: Sandy soils of wet areas, meadows, 

depressions, roadsides, road banks, river banks, 

clearings, grassy flood plains, coastal habitats, 

cane fields, lawns, and waste areas. Arroyo, 

Bayamón, Cabo Rojo, Caguas, Carolina, Cataño, 

Ceiba, Culebra, Fajardo, Guayanilla, Humacao, 

Juana Díaz, Juncos, Lajas, Las Piedras, Loíza, 

Manatí, Maunabo, Mayagüez, Naguabo, Patillas, 

Río Grande, San Juan, Vega Alta, Vega Baja, 

Vieques, and Yabucoa; Anegada, St. John, St. 


Note: G. C. Tucker (Syst. Bot. Monogr. 43: 

103. 1994) cites Clayton 598 (BM) as the type of 

Cyperus compressus L. However, that specimen 

is a different species, Cyperus bipartitus Torr. 

Fortunately, Du Puy et al. had lectotypified the 

name in its current sense a year earlier. 


Bayamón: Rd. 2, km 14.6, González-Más 1125 

(US). Carolina: Boca de Cangrejos, González-Más 

1021 (US). Cataño: Palo Seco, González-Más 2117

262 


(US). Ceiba: Bo. Machos, Axelrod & Rogowitz 

10822 (UPRRP). Culebra Island: N.L. Britton & 

Wheeler 46 (US). Guayanilla: Rd. 336, González- 

Más 818 (US). Humacao: Cotto Wakí, km 27.1, 

González-Más 74 (US). Juana Díaz: Pastillo, 

González-Más 751 (US). Juncos: locis cultis in 

monte Florida, Sintenis 1939 (US). Manatí: Bo. 

Tierras Neuvas Saliente, Axelrod et al. 11400 

(UPRRP). Maunabo: La Pica, km 105.1, González- 

Más 1339 (US). Mayagüez: Athletic field, College 

of Agriculture, González-Más 515 (US). Naguabo: 

Rd. 3, km 4.1, González-Más 1365 (US).Vieques 

Island: Playa de Esperanza, Shafer 2748 (US). ST. 

JOHN: Lameshur, Woodbury 775a/7208 (VINPS). 

VIRGIN GORDA: Near Valley, N.L. Britton & 

Fishlock 1067 (US). 

5. Cyperus confertus Sw., Prodr. 20. 1788; 

Mariscus confertus (Sw.) C. B. Clarke in 


Jamaica. Swartz s.n. (lectotype: S-Sw. R- 

1384; isolectotypes: B-Willd. 1349, S), here 

designated. 

Rhizomatous perennial, (2-) 6-50 (-75) cm 

tall; rhizome short; culms tufted or solitary, 

forming small cormlike enlargements at base with 

age, trigonous, smooth, (0.5-) 1-2 (-2.5) mm wide, 

sheathing bases 1-5 mm wide. Leaves 2-6, all basal 

or 1-3 lower cauline; sheaths often cross-veined 

proximally, pale green, tinged with red or brownish 

purple; ligule absent; blades linear, 3-35 cm × 

(0.5-) 1-5 (-7.5) mm, subflattened, soft, thinly 

herbaceous, smooth or with antrorsely scabrous 

margins and abaxial midvein distally, longacuminate 

to triquetrous apex. Inflorescence a 

simple umbel-like corymb with ascending rays or 

congested in a head in poorly developed 

individuals, (0.9-) 1.5-11 (-16) cm in diam.; 

involucral bracts leaflike, 2-6, elongate, 2-3 times 

as long as the inflorescence; rays 2-5, slender, 

patent, 1-9 (-13) cm long; spikes globose-ovoid, 

10-15 mm in diam., with 3-32 spikelets; spikelets 

subdigitately to spicately disposed on an 

abbreviated axis, dense, divergent proximally, 

jointed at base, oblong-lanceolate, 6-10 × 2-2.2 

mm, compressed, obtuse at apex, short-cuneate to 

sub-rounded at base, with 6- to 18-flowered; 

rachilla very narrowly hyaline-winged; scales 

ovate, dorsally acute, 2-2.5 × 1.8-2 mm, thinly 

herbaceous, 3- or 4-nerved on both sides, straw- 

colored or light yellowish brown, carina 3-nerved, 

wide, green, extending beyond the apex as a 0.3 

mm long distinctly to slightly excurved cusp. 

Stamens 3, the anthers 0.2-0.5 mm long, bluntly 

apiculate; style 3-branched. Achene trigonous with 

concave faces, obovoid to ellipsoid-obovoid, 1- 

1.4 × 0.7-0.8 mm, rounded above to a mucronulate, 

somewhat bulbous-thickened apex, tapered at 

base, puncticulate, brown to blackish. 

General distribution: West Indies, South 

America, and Galapagos Islands. 


Islands: Sandy soils of coastal areas, along forest 

trails, grassy areas, scrub forests, and thickets. 

Cataño, Coamo, Guayama, Loíza, Manatí, and 

Salinas; St. Croix, St. Thomas, Tortola, and Virgin 

Gorda. 


Manatí: Bo. Tierras Nuevas Saliente, Axelrod et 

al. 11491 (UPRRP). ST. THOMAS: Eggers s.n. (US). 

6. Cyperus croceus Vahl, Enum. Pl. 2: 357. 1805. 

Type: United States; South Carolina (cited by 

Vahl as “Puerto Rico” in protologue, see note 

below) Bosc s.n. (holotype: C-Vahl). 

Cyperus globulosus sensu many authors, non 

Aublet, 1775. 

Mariscus echinatus sensu C. B. Clarke in Urban, 

Symb. Antill. 2: 43. 1900, non (Linnaeus) 

Elliott, 1816. 

Mariscus globulosus sensu Urban, Symb. Antill. 

4: 113. 1903, non (Aublet) Urban, 1900. 

Rhizomatous perennial, 9-75 cm tall; rhizome 

short and thickened or shortly horizontally 

creeping. Culms caespitose, slightly bulbousbased, 

trigonous to obtusely so proximally, 

coarsely ribbed, smooth, 1-2.2 (-3) mm wide, 

sheathing bases 4-9 mm wide. Leaves 5-13, 

primarily basal, 1-3 lower cauline; sheaths short, 

pale brown, often reddish tinged and fibrillose 

proximally; ligule absent; blades linear, flattened 

to widely V-shaped or subplicate, 10-44 (-50) cm 

× (1.5-) 2.2-5 (-7) mm, antrorsely scabrous on the 

margins and abaxial midvein, attenuate to 

triquetrous apex. Inflorescence a simple umbellike 

corymb of ovoid to spherical heads of spikelets 

at the tips of 6-13 ascending to divergent rays, (2- 

) 3-12 (-14) × 4-16 cm; involucral bracts 4-11, 

leaf-like, spreading, the lowermost elongated, 

greatly exceeding the inflorescence, to 30 cm long;


spikes globose, 7-15 (-20) × 7-18 mm, with 14-60 

spikelets; spikelets lanceolate to oblonglanceolate, 

subcompressed, quadrate, 5-10 × 0.7- 

1.3 mm, acuminate at apex, cuneate at base, with 

4-6 florets; rachilla broadly hyaline-winged; scales 

ovate, acute to obtuse dorsally, 2.3-3.2 × 1.8-2.2 

mm, herbaceous, with 3-4 coarse nerves on each 

side above the scarious margins, greenish brown 

to yellowish brown, carina 3-nerved, green, 

prolonged beyond the obtuse to acute apex as a 

short, antrorsely scabrous mucro. Stamens 3, the 

anthers 0.4-0.6 mm long, bluntly apiculate; style 

3-branched. Achene trigonous, curved on side 

facing rachilla, with plane or concave faces, 

oblong-ellipsoid, 1.5-2 × 0.4-0.8 mm, obtuse to 

subtruncate at apex, apiculate, short-cuneate at 

base, substipitate, puncticulate, reddish brown to 

blackish. 


States, West Indies, Central America, and South 

America. 

Distribution in Puerto Rico: Sandy savannas, 

roadside ditches, lawns, pastures, and disturbed 

areas; primarily from Laguna Tortuguero to 

Dorado. Cabo Rojo, Carolina, Dorado, Manatí, 

San Juan, and Vega Baja. 

Note: Carter & Kral, (Taxon 39: 326. 1990) 

consider the type locality to probably be 

Charleston, South Carolina based on a collection 

of Bosc, not “Bosch” as erroneously cited by Vahl 

on the type specimen. 


Carolina: Punta Cangrejos, Stevenson & Johnston 

1707 (US); Stevenson 5477 (US). Manatí: Bo. 

Tierras Nuevas Saliente, Proctor & Thomas 43803 

(US). Vega Baja: Tortuguero, González-Más 2149 

(US). Dorado: to Cerro Gordo, Rd. 693, km 13.0, 


7. Cyperus difformis L., Cent. Pl. 2: 6. 1756. 

Lectotype: India. Collector unknown (LINN- 

70.10), designated by G. C. Tucker, Syst. Bot. 

Monogr. 43 : 50. 1994. 

Caespitose annual, (4-) 7-40 (-50) cm tall. 

Culms loosely tufted, soft, easily compressed, 

triquetrous, coarsely ribbed on margins, finely 

veined on sides, dark green, smooth, (0.7-) 1.2-3 

mm wide (when flattened). Leaves 2-7, primarily 

basal and 1-2 lower cauline; sheaths short, 1-2 mm 

long, dark green; ligule present, well-developed, 

membranous; blades linear, flattened, often folded 

proximally, (2-) 7-22 cm × 1.8-4 mm, dark green, 

essentially smooth on margins and abaxial midvein 

except for antrorsely scabrous tip, attenuate to 

apex. Inflorescence a simple or sometimes 

compound umbel-like corymb of 3-20 orbicular 

to glomerulate dense heads of spikelets on 

relatively short rays, or contracted and heads 

subsessile in an irregularly lobate mass; involucral 

bracts 2-4, leaf-like, to 22 cm long, the lowermost 

often erect or ascending, the others horizontal to 

divergent; primary rays 2-6, several sometimes to 

30 cm long, secondary rays (when present) 

inconspicuous; spikes globose, often irregularly 

lobate, 7-17 mm in diam., with 20-140 spikelets; 

spikelets oblong-ovate to lanceolate, (2-) 3-5 (-6) 

× 0.8-1.2 mm, subcompressed, obtuse at apex, 

acute at base, with (6-) 12-20 (-30) florets; rachilla 

unwinged; scales obovate to orbiculate, laterally 

compressed, 0.6-0.8 × 0.6-0.9 mm, curvate-keeled, 

dorsally acute, submembranous, lateral nerves 

indistinct, stramineous to brown-purple, carina 3nerved, 

greenish, yellowish, or whitish 

stramineous, subtranslucent, prolonged beyond the 

obtuse apex as a short mucro. Stamens 1 or 2, the 

anthers 0.2-0.4 mm long, short-apiculate; styles 

3-branched. Achene trigonous with plane or 

slightly concave faces, ellipsoid-obovoid, 0.6-0.8 

× 0.3-0.4 mm, obtuse at apex, apiculate, cuneate 

at base, finely reticulate-papillose, light brown or 

yellowish. 

General distribution: Native in the Old World 

from southern Europe, Africa, Southeast Asia, 

Malesia, Pacific Islands, and Australia. 

Naturalized in the southern United States, 

Hawaiian Islands, West Indies, Mexico, Central 

America, and Northern South America. 

Distribution in Puerto Rico: Wet sandy or clay 

soils of low marshy areas, pond and lake margins, 

pastures, rice fields, canals in agricultural areas, 

roadside ditches, and disturbed areas. Aguada, 

Arecibo, Bayamón, Caguas, Carolina, Cataño, 

Ceiba, Dorado, Fajardo, Humacao, Manatí, 

Maunabo, Mayagüez, Río Grande, San Juan, and 

Vega Baja. 



10695 (UPRRP, US). Bayamón: Bo. Minillas, 

along E side of Río Hondo, Proctor 43555 (US). 

Caguas: Bo. Bairóa, off Rt. 30 near old Central 

Santa Juana, Axelrod & L. Pérez 9479 (US).

264 


Cataño: Bo. Palmas, Rt 5 at El Puente, Axelrod & 

Leal 4267 (US). Mayagüez: College of 

Agriculture, González-Más 2135 (US). Río 

Grande: Bo. Zarzal, beyond end of Rd. 968, 

Proctor 42671 (US). San Juan: Bo. Hato Rey 

Norte, Proctor 43035 (US). Vega Baja: Tortuguero 

Lake, Woodbury s.n. (US). 

8. Cyperus digitatus Roxb., Fl. Ind. 1: 209. 1820. 

Type: India. Roxburgh s.n. (holotype: K). 

Coarse rhizomatous perennial, 70-180 cm tall; 

rhizomes short, thickened and corm-like. Culms 

solitary or two approximate, erect, stiff and 

hardened, trigonous or triquetrous proximally, 

triquetrous distally, often with a medial 

longitudinal furrow proximally, coarsely and 

finely ribbed, 4-12 mm wide, smooth or rarely 

sparsely scabrellate on angles distally, sheathing 


cauline; sheaths elongate, herbaceous, brown to 

brown-black proximally, pale brown distally, open, 

with scarious margins, the inner band essentially 

absent; ligule absent; blades linear-lanceolate, 

folded or V-shaped proximally, plicate or 

subflattened distally, spongy-thickened 

proximally, 60-130 cm × 7-20 mm, antrorsely 

scabrous on margins and abaxial midvein, 

attenuate to triquetrous apex. Inflorescence a large, 

compound or decompound, hemispherical, umbellike 

corymb with ascending rays, 14-33 (-38) × 

15-30 (-34) cm, with numerous spikes of densely 

aggregated spikelets; involucral bracts 8-12 (-15), 

(5-) 9-90 cm × 3-20 mm, leaf-like, plicate, 

spreading-ascending to horizontal, often purplish 

brown at base; primary rays 7-10, to 25 cm long, 

secondary rays to 5 cm long; spikes cylindrical, 

(2.5-) 3.5-7 (-7.5) × 1-2 (-2.5) cm, with 30-140 

spikelets; spikelets linear, compressed-rhombic, 

5-11 × 1-1.2 mm, acute at apex, cuneate at base, 

with 12-30 florets, spreading at right angles to the 

rachis, appearing serrated along edge due to the 

spreading scales; rachilla broadly hyaline-winged, 

the wings light brown, caducous; scales ovate or 

ovate-elliptic, acutely keeled dorsally, 1.4-2 × 1- 

1.3 mm, submembranous, laterally weakly 1-2nerved 

along carina, light to reddish brown on 

sides, with 2 dark brown stripes paralleling either 

side of carina, carina 3-nerved, green, apiculate at 

the acute to obtuse apex. Stamens 3, the anthers 

0.4-0.6 mm long, bluntly apiculate; style 3- 

branched. Achene subtrigonous with the side 

facing the rachilla concave and the other faces 

plane or slightly convex, ellipsoid, oblongellipsoid, 

or ellipsoid-obovoid, 0.8-1 × 0.4-0.5 

mm, dorsiventrally compressed, obtuse at apex, 

short-apiculate, cuneate at base, estipitate, glossy 

and appearing essentially smooth, indistinctly 

finely puncticulate, light brown to light grayish 

brown at maturity. 

General distribution: Pantropical; southern 

United States (Florida, Louisiana, and Texas), West 

Indies, Mexico, Central America, Africa, East 

Asia, Malesia, and Australia. 

Distribution in Puerto Rico: In wet grounds 

of marshy areas, pond and lagoon margins, and 

pastures. Cabo Rojo, Lajas, and Toa Baja. 


Toa Baja: Bo. Sabana Seca, area S of Rt. 867, near 

Naval Station, Axelrod & L. Pérez 9825 (US). 

9. Cyperus distans L.f., Suppl. Pl. 103. 1782 [as 

a new name]; Cyperus elatus Rottb., Descr. 

Icon. Rar. Pl. 37. 1773, non Linnaeus, 1756. 

Type: India; Kerala, Malabar. Collector 

unknown. (holotype: C-Rottb.; isotype: C- 

Vahl). 

Perennial, (20-) 30-115 (-138) cm tall; 

rhizome very short, corm-like. Culms solitary or 

sometimes 2-3, trigonous, finely to coarsely ribbed 

and channeled, smooth, 2-6 mm wide proximally, 

sheathing bases (4-) 5-15 (-20) mm wide. Leaves 

5-7, basal and lower cauline; sheaths cross-veined, 

pale brown to reddish brown; ligule absent; blades 

folded or V-shaped proximally, flattened or plicate 

distally, 20-60 cm × (3-) 5-12 mm, subabruptly 

acuminate at apex, the margins, mid rib below, 

and the two lateral veins above antrorsely 

scabrous. Inflorescence a compound, 

hemispherical, umbel-like corymb, (6-) 11-26 (- 

34) cm diam., with numerous spikes at ray tips; 

involucral bracts leaf-like, (5-) 6-7 (-10), the lower 

to 70 cm long, 3.5-13.5 mm wide, the uppermost 

linear to subulate, shorter; primary rays elongate, 

6-9, unequal, to 17 cm × 0.5-1.2 (-2) mm, with 

tubular prophylls prolonged dorsally into 2, long, 

linear-subulate teeth; secondary rays very short or 

the spikes sessile; spikes oblong-ovate to broadly 

ovate, 2.5-3.5 (-4) × 1.5-3 (-4) cm, with 5-30 laxly 

disposed spikelets; spikelets linear, cylindrical, 

slightly ascending to divergent, 7-30 (-70) × 0.4-


0.6 mm (to 1.8 mm wide after spreading of mature 

florets), acuminate at apex, cuneate at base, with 

(5-) 7-35 (-75) florets; rachilla flattened, zigzagged, 

hyaline-winged (wings caducous), 

persistent; scales elliptic to oblong-elliptic, boatshaped 

to obtusely keeled, 1.6-2 × 0.8-1.1 mm, 

remotely 2-ranked, spreading at maturity, 

membranous with broad, scarious margins, weakly 

1- to 2-nerved on each side along carina, reddish 

brown on sides, carina 3-nerved, greenish, not 

prolonged beyond the scarious, obtuse, emarginate 

apex, at most forming a minute apiculum. Stamens 

3, the anthers 0.5-0.7 mm long, apiculate; style 3branched 

from just above the base. Achene 

trigonous with plane or slightly convex faces, 

ellipsoid-obovoid or oblong to oblong-ellipsoid, 

1.2-1.7 × 0.4-0.5 mm, slightly curved on side 

facing rachilla, acutely narrowed to apiculate apex, 

cuneate at base, estipitate, minutely reticulate, dark 


General distribution: Cosmopolitan; more 

common in warmer regions. 


Islands: In wet soils of meadows, riverbanks, 

marshy areas, grassy areas, scrub forests, rocky 

thickets and clearings in forests, hillsides, 

agricultural fields, pastures, roadside ditches, and 

disturbed open areas. Adjuntas, Aguas Buenas, 

Aibonito, Añasco, Barranquitas, Bayamón, 

Caguas, Canóvanas, Carolina, Cayey, Ceiba, 

Cidra, Coamo, Comerío, Culebra, Dorado, 

Fajardo, Guayama, Guayanabo, Humacao, Las 

Piedras, Loíza, Maunabo, Maricao, Mayagüez, 

Naguabo, Patillas, Peñuelas, Ponce, Río Grande, 

San Juan, Utuado, Vega Alta, Vieques, Villalba, 

and Yabucoa; St. Croix, St. John, St. Thomas, and 

Tortola. 


Adjuntas: near Pico Guilarte, Liogier 10004 (US). 


(US). Canóvanas, Stevenson 3004 (US). Cayey: 

Sintenis 2376 (US). Coamo: Bo. San Ildefonso, 

Baños de Coamo, Proctor 42430 (US); Coamo 

Springs, Goll 669 (US). Comerío: Río Hondo, 

González-Más 382 (US). Dorado: Rd. 693, km 1.9, 

González-Más 1123 (US). Fajardo: Bo. Río Arriba, 

Axelrod & Thomas 10724 (UPRRP). Humacao, 

González-Más 1352 (US). Las Piedras: Rd. 5, km 

26.4, González-Más 3 (US). Maricao: Maricao 

Forest near Hatchery, González-Más 1643 (US). 

Mayagüez: Buena Vista, González-Más 850 (US). 

Naguabo: González-Más 1359 (US). Ponce: Cotto 

Laurel, Rd. 14, km 122.2, Bucaná, González-Más 

779 (US). San Juan: Río Piedras, Sabana Llana, 

Highland Park grounds, González-Más 1217 (NY), 

González-Más 1214 (US). Yabucoa: Rd. 3, km 

96.2, González-Más 1345 (US). ST. CROIX: 

Fountain Valley, Fosberg 59329 (US). ST. JOHN: 

Bordeaux Mt., Woodbury 59/6545 (VINPS); Cruz 

Bay Quarter; Gift Hill, Acevedo-Rdgz. 5282 (NY, 

US). ST. THOMAS: St. Peter, E.G. Britton & Marble 

1223 (US); Signalhill, Eggers s.n. (NA). TORTOLA: 

Fishlock, W.C. 60 (US). Road Town to High Bush, 

N.L. Britton & Shafer 785 (US). 

10. Cyperus eggersii Boeck., Beitr. Cyper. 1: 53. 

1888; Torulinium eggersii (Boeck.) C. B. 

Clarke in Urban, Symb. Antill. 2: 56. 1900. 

Lectotype: Dominican Republic. Eggers 2627 

(B; isolectotype: C), designated by G. C. 

Tucker, Syst. Bot. Monogr. 43: 181. 1994. 

Rhizomatous perennial, often loosely 

caespitose, 14-65 cm tall; rhizome short, 

thickened; roots medium to coarse. Culms erect, 

triquetrous, sometimes trigonous proximally, stiff 

and hardened, finely ribbed, smooth, pale green, 

1.5-3.5 mm wide, sheathing bases 4-18 mm wide. 

Leaves 3-7, ascending, basal and lower cauline, 

those at base often with short blades; sheaths 

elongate, herbaceous, dark brown with pale veins; 

ligule absent; blades linear, broadly V-shaped 

proximally just above sheath apex, plicate or 

sometimes subflattened distally, 14-45 (-60) cm × 

3.5-8 (-11) mm, herbaceous, abaxial mid and 

lateral veins antrorsely scabrous, margins 

antrorsely scabrous, attenuate to triquetrous tip. 

Inflorescence a simple to compound, umbel-like 

corymb with ascending rays, or reduced to a single, 

essentially sessile, lobate head of spikes, (1.5-) 2.5- 

14 × (2-) 3.5-18 cm; primary involucral bracts 4- 

8, leaf-like, greatly exceeding the inflorescence; 

secondary involucral bracts linear-setaceous; rays 

2-8 or absent; spikes ovoid, in lobate heads of 

(2-) 3 (-4) at ray tips, with 20-60 congested 

spikelets; spikelets linear, subcylindrical to slightly 

compressed, 5-15 × 0.8-1.2 (-1.5) mm, with 5-22 

florets, the scales spreading with maturing 

achenes; rachilla wings spongy-thickened, 

enveloping achene at maturity; scales elliptic to 

elliptic-obovate, broadly rounded in cross section, 

1.7-2.2 (-2.4) × 1-1.5 mm, submembranous, light 

brown, smooth, laterally 2-3 nerved on each side 

along carina, carina finely 5-nerved, green,

266 


prolonged beyond the obtuse apex as short mucro. 


apiculate; style 3-branched. Achene oblongobovate, 

compressed-trigonous, with plane to 

slightly convex sides, falcate, 1.3-1.5 × 0.4-0.6 

mm, obtuse at apex, apiculate, cuneate at base, 

stipitate, puncticulate, light brown to brown. 

General distribution: Southwestern United 

States, West Indies, Mexico, Central America, and 

Andean South America. 

Distribution in Puerto Rico: In wet sandy 

soils, often swampy or marshy areas bordering 

lakes and ponds. Isabela, Manatí, Mayagüez, 

Quebradillas, San Sebastián, Vega Alta, and Vega 

Baja. 

Note: This plant was included within the 

circumscription of Cyperus odoratus L. by G. C. 

Tucker (1994). However, C. eggersii consistently 

differs from C. odoratus by the dark brown 

sheaths, lobate heads of spikes with crowded 

spikelets, and closely overlapping spikelet scales 

(vs. green to purple-tinged sheaths, laxly disposed 

spikes, and spikelet scales that overlap, at the most, 

only the base of the next scale above). 


Isabela: Bo. Coto, W. bank, estuary of Río 

Guajataca near its mouth, Proctor et al. 42358 

(US). San Sebastián: Bo. Aibonito, Area 

Recreativa de Guajataca, Proctor & Vives 49274 

(US). Vega Alta: Bo. Sabana, Proctor 45794 (NY, 

US). Vega Baja: Along the shoreline of Laguna 

Rica near Camp Tortuguero National Guard, off 

Rt. 687, Strong et al. 430 (US). 

11. Cyperus elegans L., Sp. Pl. 45. 1753. 


t. 75, f. 1. 1707, designated by G. C. Tucker & 

McVaugh in McVaugh, Fl. Novo-Galiciana 

13: 292. 1993. 

Cyperus viscosus Sw., Prodr. 20. 1788; Scirpus 

viscosus (Sw.) Lam., Tabl. Encycl. 1: 142. 

1791. Type: Jamaica. Swartz s.n. (holotype: 

S-Sw. R-1387). 

Cyperus confertus sensu Grisebach, Fl. Brit. W. I. 

563. 1864, non Swartz, 1788. 

Fig. 48. I-K 


rhizome short, ascending to vertical, 2.6 mm thick. 

Culms obtusely trigonous to subcylindrical, 

remotely antrorsely scabrous, 1.5-3.2 (-3.8) mm 

wide, sheathing bases 5-11 mm wide. Leaves 2-3, 

primarily basal; sheaths elongate, cross-veined, 

stramineous or tinged with brown; ligule absent; 

blades linear, stiffly ascending, thickly herbaceous, 

inrolled along margins, laterally compressed, 

channeled towards apex, 6-75 cm × (1.6-) 2-4 

(-5) mm (unfolded), often sticky, smooth, soft and 

spongy proximally, cross-veined distally, coarsely 

and remotely scabrous on the margins, often 

bearing pairs or small groups of antrorse and 

retrorse barbs, long-attenuate to a bristly, 

somewhat triquetrous apex. Inflorescence a 

compound, umbel-like corymb with ascending 

rays, (5-) 7-20 (-26) cm diam.; involucral bracts 

5-6, leaf-like, cross-veined, the lowermost one to 

75 cm long; primary rays 5-9, unequal, ascending 

at anthesis, to 22 cm long; secondary rays 1-5, to 

5 cm long, ascending at anthesis, becoming 

reflexed at maturity; spikes in dense clusters or 

glomerules at ray tips, 7-30 mm diam., with 5-40 

palmately disposed spikelets; spikelets ovate, 

oblong-ovate, or ovate-elliptic, subflattened, 5-22 

× (2-) 3-5 (-7) mm, acute at apex, widely cuneate 

at base, with 6-34 florets; rachilla wingless; scales 

broadly ovate, obtusely curvate-keeled, 2.2-3.2 × 

2-2.8 mm, with 2-3 coarse nerves on each side 

above the scarious margins, pale green, tinged with 

reddish brown on sides, sticky near base at 

maturity, carina 5-nerved, pale greenish, coarsely 

antrorsely scabrous on carina distally, the apex 

slightly excurved-cuspidate. Stamens 3, the 

anthers 0.6-1.2 mm long, apiculate; styles 3branched 

from just above the base. Achene 

trigonous with plane, concave or slightly convex 

faces, obovoid, 1.3-1.9 × 0.7-1 mm, obtuse to 

rounded at apex, with thickened margins, shortbeaked, 

narrowly cuneate at base, short-stipitate 

puncticulate, maturing blackish. 


Mexico, Nicaragua, South America, and 

throughout the West Indies. 


Islands: Wet, coastal habitats, often in brackish 

water of swampy or marshy areas, borders of 

mangroves, pastures, roadside ditches, shallow 

depressions on limestone, and disturbed areas. 

Arecibo, Cabo Rojo, Cataño, Guánica, Guayama, 

Guayanilla, Juana Díaz, Mona Island, Peñuelas, 

Ponce, Río Grande, Salinas, San Juan, Santa 

Isabel, and Vega Baja; Anegada, St. Croix, St. 

John, and St. Thomas. 


Arecibo: Sargent 678 (US). Cabo Rojo: Bo.


Boquerón, just NW of village of Boquerón, 

Proctor & McKenzie 43765 (US). Cataño: Bo. 

Palmas, Rt. 5 at El Puente, Axelrod & Leal 4261 

(US). Guayama: Puerto Jobos, Rt. 707, González- 

Más 648 (US). Juana Díaz: Rt. 14, González-Más 

775 (US). Mona Island: E of Cabo Noroeste, 

Breckon et al. 6124 (US). Peñuelas: Rd. 2, km 10.9, 

González-Más 810 (US). Ponce: Rt. 1, Mucadita, 

González-Más 764 (US). Salinas: entrance to Rd. 

705, km 1, González-Más 723 (US). San Juan: Bo. 

Hato Rey Norte, along Kennedy Ave. (Highway 

2), Proctor 43036 (US). Santa Isabel: Rt. 1, km 

110.5, González-Más 757 (US).Vega Baja: Bo. 

Cabo Caribe, between Los Naranjos and Caño Las 

Pozas, Proctor et al. 45594 (US). ST. CROIX: Bassin 

Lagoon, A.E. Ricksecker 59 (US). ST. JOHN: 

Lameshur, N.L. Britton 605 (US); Coral Bay 

Quarter; trail to Fortsberg, Acevedo-Rdgz. 4084 

(JBSD, MO, NY, UPR, US, VINPS). ST. THOMAS: 

Magens Bay, E.G. Britton & Marble 1299 (US); 

Long Cay, Eggers s.n. (NA). 

12. Cyperus esculentus L., Sp. Pl. 45. 1753. 

Lectotype: “Habitat Monspelii, inque Italia, 

Oriente.” Bauhin, Theatr. Botanici, 222, 1658, 

designated by Simpson in C. E. Jarvis et al., 

Regnum Veg. 127: 41. 1993. 

Cyperus lutescens Torr. & Hook., Ann. Lyceum 

Nat. Hist. New York 3: 433. 1836. Cyperus 

esculentus var. macrostachyus Boeck., 

Linnaea 36: 291. 1870. Type: United States; 

Texas, Drummond 452 (holotype: NY; 

isotype: GH). 

Cyperus cubensis Steud., Syn. Pl. Glumac. 2: 42. 

1854. Type: Cuba. Collector unknown 

(holotype: probably at P). 

Stoloniferous perennial, (20-) 25-70 (-90) cm 

tall; stolons flexible, slightly spongy, elongate, 1- 

2 mm thick, sometimes bearing small tubers. 

Culms solitary, trigonous proximally, triquetrous 

distally, firm, finely ribbed, smooth, 1.4-3.8 mm 

wide, sheathing bases 4-14 mm wide. Leaves 6- 

13 (-23), primarily basal, 1-3 lower cauline; 

sheaths short, indistinctly cross veined, pale 

brown; ligule absent; blades V-shaped proximally 

to plicate distally, (3-) 10-70 (-85) cm long, the 

lowermost often equaling length of culm, (-2) 3-7 

(8.5) mm wide, finely antrorsely scabrous on 

margins and abaxial midvein, at least distally, 

attenuate to triquetrous apex. Inflorescence an 

open, compound, hemispherical, umbel-like 

corymb, 3.5-14 (-18) × 3-12 (-15) cm, with 

ascending rays; involucral bracts 3-5 (-7), leaf like, 

ascending to horizontal, the lowermost 2 generally 

exceeding the inflorescence, with the lowest one 

being 7-60 cm long; primary rays 3-11, to 20 cm 

long, ascending, secondary rays shorter, often not 

evident; spikes 1 (-3) at ray tips, broadly ovoid to 

ellipsoid, 18-35 mm in diam., with (3-) 8-40 (-60) 

spikelets; spikelets linear, subquadrate, (5-) 9-25 

(-30) × (1.2-) 1.5-1.8 (-2.3) mm, acuminate at apex, 

cuneate at base, with (7-) 9-32 florets; rachilla 

hyaline-winged; scales ovate to elliptic, acutely 

keeled, 2.7-3.8 (4.2-) × 1.4-2.2 mm, submembranous, 

with 2-3 well-developed lateral 

nerves evident the length of the scale, light 

yellowish brown to brown or golden-brown, carina 

3-nerved, inconspicuous, greenish, prolonged 

beyond the obtuse to acute apex as a short, slightly 


long, apiculate; styles 3-branched. Achene 

trigonous with the adaxial face slightly concave 

and the abaxial faces plane or slightly convex, 

ellipsoid to oblong-ellipsoid or ellipsoid-obovoid, 

(1-) 1.3-1.6 × 0.3-0.8 mm, obtuse at apex, shortcuneate 

at base, short-apiculate, estipitate, 

puncticulate-reticulate, glossy, grayish brown to 

brown. 

General distribution: A cosmopolitan weed; 

in temperate and tropical regions; often cultivated. 


Islands: In waste areas, gardens, pastures, lawns, 

and other disturbed habitats. Carolina, Dorado, 

Fajardo, Guayama, Mayagüez, Río Grande, 

Quebradillas, San Juan, Toa Baja, and Vieques; 

Anegada and St. Thomas. 

Common name: Puerto Rico: Chufas. 

Note: This species is closely related to 

Cyperus rotundus L., but can be distinguished from 

it by its light brown or yellowish brown spikelet 

scales with 2-3 well-developed lateral nerves 

evident the length of the scale (vs. dark reddish 

brown to purple-brown or blackish scales that have 

1-2 indistinct lateral nerves evident only at 

proximal base of scale, the sides smooth for the 

most part); and longer inflorescence bracts, the 

lowermost 2 generally exceeding the 

inflorescence, with the lowest one being 7-60 cm 

long (vs. all bracts generally shorter than to 

equaling or only the lowest shortly exceeding the 

inflorescence, 1-9 (-14) cm long).

268 



Guayama: On road from Guayama to Aguirre, 

Underwood & Griggs 379 (US). Toa Baja: Palo 

Seco, Hioram 809 (US). Vieques Island: Near 

terminal building, Proctor 46586 (US). ANEGADA: 

The Settlement, Proctor & Haneke 45978 (US). 

13. Cyperus filiformis Sw., Prodr. 20. 1788. 

Torulinium filiforme (Sw.) C. B. Clarke in 


Jamaica. Swartz s.n. (holotype: C, no. R-1385; 

isotype: B-Willd. 1292). 


rhizome short, knotty, branching horizontally. 

Culms loosely tufted, firm but flexuous, often 

arching, very slender, triquetrous with obtuse 

angles, angles sometimes antrorsely scabrous just 

below inflorescence, finely ribbed, smooth, 0.3-1 

mm wide. Leaves 1-3, basal and lower cauline, 

the lowermost with short blades or bladeless; 

sheaths short, brown or purplish brown, inner band 

membranous or thinly herbaceous, with truncate 

to slightly convex orifice; ligule absent; blades 

narrowly linear, plicate, 3-40 cm × (0.5-) 1-2.3 

mm, semi-glossy and cellular-reticulate adaxially, 

antrorsely scabrous on the margins and abaxial 

midvein towards apex, smooth below, acuminate 

to triquetrous apex. Inflorescence a terminal 

corymb, 1.2-5 × 0.7-3 cm, or reduced to 2-6 

subfastigate spikelets; involucral bracts 1-3, leaflike, 

the lowest well-developed, erect and 

appearing as a continuation of the culm, exceeding 

the inflorescence, upper bracts setaceous; rays 1- 

3, often inconspicuous, sometimes elongating up 

to 4 cm; spikes 1-3, loose, obconic, 12-30 × 7-20 

mm, with 1-6 spikelets; spikelets linear-subulate, 

subterete, (8-) 10-30 × 0.8-1.2 mm, acuminate at 

apex, cuneate at base, with 4-13 florets; rachilla 

flexuose, broadly winged, with coriaceous wings 

that clasp the achene at maturity; scales ovateelliptic, 

dorsally obtuse to rounded, 2.5-3.5 × 1.7- 

2 mm, thickly herbaceous to subcoriaceous, with 

5-7 nerves on each side above the scarious 

margins, stramineous to yellowish or reddish 

brown, dark reddish lineolate, carina 3-nerved, 

green, prolonged at subobtuse apex as a short 

mucro Stamens 3, the anthers 0.4-0.6 mm long, 

truncate-apiculate or sometimes 2-lobed at tip; 

styles 3-branched. Achene trigonous with plane 

to slightly concave faces, oblong-obovate, curvate 

on adaxial side, 1.3-1.9 × 0.5-0.8 mm, abruptly 

narrowed to an obtuse apex, apiculate, subrounded 

at base, short-stipitate, puncticulate, 

brown to dark brown or grayish brown at maturity. 

General distribution: Endemic to the West 

Indies. 

Distribution in the Virgin Islands: Moist 

sandy, gravelly, or rocky hillsides and road banks. 

St. Croix and St. Thomas. 


Eggers 77 (US); Eggers s.n. (NA, US); Vélez 

30175 (US). 

14. Cyperus flavescens L., Sp. Pl. 46. 1753. 

Pycreus flavescens (L.) P. Beauv. ex Rchb., 

Fl. Germ. Excurs. 72. 1830. Lectotype: Herb. 

Burser 1: 81 (UPS), designated by Kukkonen 

in Cafferty & C. E. Jarvis (ed.), Taxon 53: 

178. 2004. 

Slender annual, (6-) 10-30 (-40) cm tall. 

Culms densely tufted, slender, soft, flexuous, 

triquetrous, finely ribbed, smooth, 0.4-1.3 mm 

wide. Leaves 2-4 per culm, basal and lower 

cauline; sheaths purple-tinged to purple-brown 

proximally, pale green distally; ligule absent; 

blades narrowly linear, V-shaped or folded 

proximally, flattened to subplicate distally, 3-20 

cm × 0.5-2 (-2.2) mm, antrorsely scabrous on the 


attenuate to triquetrous apex. Inflorescence a 

simple, umbel-like corymb with few rays, often 

reduced to a single spike, 1-7 cm in diam.; 

involucral bracts 2-3, leaf-like, spreading, the 

lowermost to 15 cm long; rays not evident or 1-3, 

short, to 4 cm long; spikes widely ovoid to 

subglobose, 1-4 cm diam., with 4-25 (-30) 

spikelets; spikelets strongly laterally flattened, 

oblong to oblong-lanceolate, 4-12 × 2-2.5 mm, 

excluding the denuded rachilla, subacute at apex, 

obtuse at base, with (8-) 12-24 (-40) florets; 

rachilla 4-angled, unwinged; scales ovate to ovateelliptic, 

narrowly acute dorsally, curvate-keeled, 

1.5-2 × 1-1.6 mm, submembranous, with 1 nerve 

on each side bordering carina, the sides essentially 

nerveless above the narrowly scarious margins, 

light yellowish brown, carina 3-nerved, greenish, 

ending in a short cusp at the broadly acute apex. 

Stamens 2 or sometimes 3, the anthers 0.4-0.5 mm 


thickly biconvex, orbicular-obovate, 0.8-1 × 0.5-


0.7 mm, sub-rounded to rounded at apex, often 

asymmetrically so, apiculate, abruptly cuneate at 

base, stipitate, transversely rugulose-reticulate, 

often inconspicuously so, with fine transverse 

often whitish undulations that sharply contrast with 

the brown to black color of the achene. 

General distribution: Cosmopolitan; in 

temperate and tropical regions. 

Distribution in Puerto Rico: In wet habitats 

including seepages, marshy areas, swales, and 

ditches. Bayamón, Cidra, Dorado, and Río Grande. 


Bayamón: Hioram 359 (US). Dorado: N.L. Britton 

& E.G. Britton 7534 (US). Río Grande: Sierra de 

Luquillo, Proctor 42281 (US). 

15. Cyperus flexuosus Vahl, Enum. Pl. 2: 359. 

1805; Torulinium flexuosum (Vahl) T. 

Koyama, Phytologia 29: 74. 1974; Diclidium 

vahlii Schrad. ex Nees in Martius, Fl. Bras. 

2: 53. 1842, nom. illeg.; Cyperus vahlii Steud., 

Syn. Pl. Glumac. 2: 48. 1854, nom. illeg.; 

Torulinium vahlii C. B. Clarke in Urban, 

Symb. Antill. 2: 56. 1900, nom. illeg. Type: 

St. Croix, U.S. Virgin Islands. von Rohr 173 

(holotype: C-Vahl; photo at F and US). 

Cyperus ehrenbergii Kunth, Enum. Pl. 2: 89. 1837. 

Lectotype: St. Thomas, U.S. Virgin Islands. 

Ehrenberg 77 (HAL), here designated. 

Cyperus insignis Kunth, Enum. Pl. 2: 92. 1837. 

Cyperus flexuosus var. insignis (Kunth) Kük. 

in Engler, Pflanzenr. IV. 20 (Heft 101): 622. 

1936. Type: Guadeloupe. Bertero s.n. 

(holotype: probably at TO or P; isotype: B, 

destroyed). 

Rhizomatous perennial, 20-90 cm tall; 

rhizome short, subbulbous, woody, 7-15 mm thick. 

Culms 1 to several, trigonous proximally to 

obtusely triquetrous distally, smooth, 2.5-4.5 mm 

wide near base, sheathing bases 0.6-1 cm wide. 

Leaves 4-8, all basal; sheaths short, cross-veined, 

light brown, purplish proximally; ligule absent; 

blades linear, flattened, 14-45 cm × 4.5-11 mm, 

remotely antrorsely scabrous on margins, 

smoothish on midvein beneath, long-acuminate to 

triquetrous apex. Inflorescence a simple or 

compound umbel-like corymb with ascending 

rays, (2-) 5-25 cm diam.; involucral bracts 7-8, 

elongate, leaf-like, 3-51 cm long; rays 8-10, to 14 

cm long; spikes 1-7 at ray tips, widely ovate to 

rounded, 1.6-3.5 × 1.3-3 cm, with 30-80 spikelets; 

spikelets linear, subcylindrical, 0.5-2 cm × 0.8- 

1.1 mm, spicate or sometimes radiate, somewhat 

congested, with the internodes of the rachis 

between spikelets less than 1 mm, the lower ones 

reflexed at maturity, with 3-12 florets, 

disarticulating at base and rachilla nodes; rachilla 

with broad scarious wings, these at maturity 

becoming thick and corky, yellowish to brown, 

enveloping the achene, the rachilla node and 

flower falling together; scales ovate to ellipticovate, 

somewhat dorsally compressed, dorsally 

rounded to broadly so, inrolled along margins, 2.3- 

3.5 × 2.0-2.5 mm, obscurely 1-2 nerved on each 

side bordering carina above the narrowly scarious 

margins, thickly membranous to subcoriaceous, 

chestnut-colored, shiny, reddish brown to dark 

brown on sides, carina finely 5- to 7-nerved, green, 

extending as a short, prickly-tipped mucro below 

the obtuse apex. Stamens 3, the anthers 0.5-1 mm 


trigonous with plane to slightly convex faces, 

oblong-obovoid, dorsiventrally compressed, 1.2- 

1.9 × 0.5-0.7 mm, slightly falcate at maturity, 

narrow acutely to short-beaked apex, cuneate at 

base, estipitate, minutely puncticulate, light brown 

to brown at maturity. 

General distribution: West Indies, Mexico, 

and Brazil. 

Distribution in the Virgin Islands: Wet or 

moist soils of roadside ditches. St. John and St. 

Thomas. 

Note: Reported for Puerto Rico (Culebra 

Island) by Axelrod (Caribbean J. Sci. 29: 126: 

1993), based on Axelrod et al. 1695 (UPRRP; 

xerox at US), but that specimen is Cyperus 

planifolius Rich. 


Lameshur, N.L. Britton & Shafer 623 (US); Reef 

Bay Quarter; along new road from Bordeaux Road 

to Coral Bay, Acevedo-Rdgz. & Siaca 3799 (MO, 

NY, US). ST. THOMAS: St. Peter, E.G. Britton & 

Marble 1224 (US). 

16. Cyperus floridanus Britton in Small, Fl. s.e. 

U.S. 170. 1903; Mariscus floridensis C. B. 

Clarke, Bull. Misc. Inform. Kew Addit. Ser. 

8: 20. 1908, nom. illeg. Type: United States; 

Florida, Key West. Blodgett s.n. (holotype: 

NY; fragment at US!). 

Cyperus filiformis var. densiceps Kük., Repert. 

Spec. Nov. Regni Veg. 23: 190. 1926.

270 


Lectotype: Cuba; Oriente. Ekman 7803 (NY!; 

isolectotype fragment: US), here designated. 

Cyperus kingsii C.D. Adams ex Proctor, Sloanea 

1: 1. 1977. Type: Cayman Islands; Grand 

Cayman. W. Kings GC410 (holotype: BM). 


short, knotty. Culms erect or sometimes reclining, 

obtusely trigonous, filiform, wiry, 0.3-0.8 mm 

wide, finely ribbed and often coarsely channeled 

on sides, smooth, sheathing bases 0.8-1.8 mm 

wide. Leaves primarily basal, 1-2 lower cauline, 

often equaling to overtopping culms; sheaths short, 

light brown to brown, the veins often reddish; 

ligule absent; blades U-shaped, V-shaped, or often 

involute and appearing filiform, 0.5-30 cm × 0.2- 

1.2 mm, finely veined abaxially, subglossy and 

cellular-reticulate adaxially, smooth to remotely 

antrorsely scabrous on the margins, attenuate to 

triquetrous apex. Inflorescence a narrowly 

hemispherical, terminal cluster of 1-5, shortpeduncled 

spikes, appearing pseudolateral, the 

lowest bract erect, appearing as a continuation of 

the culm, 5-15 mm in diam.; involucral bracts 2- 

4, leaf-like, the lowermost one to 20 cm long; rays 

short, essentially absent; spikes with 4-16 densely 

disposed spikelets; spikelets linear-subulate, 3-14 

(-17) × 0.5-1 (-1.5) mm, the scales widely 

spreading with maturing achenes, acute at apex, 

narrowly cuneate at base, with 5-16 florets; rachilla 

with hyaline wings; scales ovate-elliptic, dorsally 

obtuse, slightly falcate-spreading, concavely 

curvate-keeled or keel straight, 1.4-2.2 × 0.8-1.2 

mm, herbaceous to thickly so, with 4-5 nerves on 

each side above the narrowly scarious margins, 

maroon, carina 3-nerved, green, prolonged beyond 

the obtuse apex as a short mucro. Stamens 3, the 


3-branched. Achene trigonous, slightly curvate, 

with adaxial face concave and adaxial faces plane, 

linear-oblong, 1.3-1.6 × 0.4-0.6 mm, shortapiculate, 

obtuse to sub-rounded at base, estipitate, 

puncticulate, brown to dark brown or brown-black. 


(Florida) and the West Indies. 

Distribution in Puerto Rico: Soil-filled 

depressions on limestone. Guánica and Mona 

Island. 


Guánica: San Jacinto Beach, González-Más 966 

(MAPR). Mona Island: between Uvero and Bajura 

de los Cerezos, Proctor & Haneke 43147 (SJ). 

17. Cyperus fuligineus Chapm., Fl. South. U. S. 

511. 1860; Mariscus fuligineus (Chapm.) C. 

B. Clarke in Urban, Symb. Antill. 5: 290. 

1907. Type: United States; Florida, Key West. 

Chapman s.n. (holotype: NY!; fragment at 

US!). 


short, knotty. Culms slender but somewhat stiff, 

firm, thickened at base, obtusely trigonous, finely 

ribbed, channeled medially on sides, smooth, 0.6- 

1.5 mm wide, sheathing base of culm 1-3 mm 

wide. Leaves 2-6, primarily basal, 1-2 lower 

cauline; sheaths short, to 5 cm long, the lower 

bladeless or nearly so, dark red, reddish purple or 

purple-brown, the orifice truncate, often with a 

cleft medially; ligule absent; blades linear, Vshaped 

to subplicate, sometimes folded, stiffish 

and incrassate proximally, slightly revolutemargined, 

0.5-21 cm × (1-) 1.3-3 (-3.5) mm, finely 

veined abaxially, faintly veined or smooth and 

semi-glossy adaxially, antrorsely scabrous on the 


attenuate to triquetrous apex. Inflorescence a 

spherical or hemispherical, head-like or lobate 

corymb of 8-50 (-70) radiate spikelets, 0.8-2 cm 

in diam.; involucral bracts 2-3, leaf-like, ascending 

to divergent, often reflexed at maturity, the 

lowermost one elongated, greatly exceeding the 

inflorescence, to 12 cm long; spikes indistinct; 

spikelets elliptic-lanceolate to oblong-lanceolate, 

subcompressed, 4-10 × 1-1.5 (-2) mm, acute to 

acuminate at apex, short-cuneate at base, with 6- 

16 florets; rachilla narrowly hyaline-winged; 

scales ovate to broadly ovate, dorsally obtuse, or 

acute when immature, 1.6-2.1 × 1.2-1.6 mm, 

chartaceous, semi-glossy, lineolate adaxially, with 

4-5 coarse nerves on each side above the narrowly 

scarious margins, light yellowish brown and tinged 

with red-brown, uniformly dark reddish brown, 

or purple black distally, carina relatively broad, 

obscurely 3-nerved, greenish, prolonged beyond 

the obtuse to sub-rounded apex as a short, scabrous 


bluntly apiculate; style 3-branched. Achene 

triquetrous or with the 2 abaxial faces plane, 

ellipsoid-obovoid to oblong-obovoid, 1-1.3 × 0.5- 

0.8 mm, apiculate, short-cuneate at base, estipitate, 

puncticulate, maturing black. 


(Florida) and the West Indies.



Islands: Growing in depressions of dry limestone 

rock in coastal areas. Cayo Icacos; Anegada. 

Selected specimens examined: ANEGADA: West 

End, N.L. Britton & Fishlock 951 (US). 

18. Cyperus giganteus Vahl, Enum. Pl. 2: 364. 

1805. Type: Puerto Rico. West s.n. (holotype: 

C-Vahl, photo at F, US; isotypes: B, C, 

fragment at US!). 

Papyrus comosus Kunth in Humb., Bonpl., & 

Kunth, Nov. Gen. Sp. 1 [quarto ed.]: 218. 

1816; Cyperus comosus (Kunth) Poir. in 

Lamarck, Encycl. Suppl. 5. 185. 1817, non 

Sibthorp & Smith, 1806; Cyperus giganteus 

var. comosus (Kunth) Kük., Repert. Spec. 

Nov. Regni Veg. 32: 74. 1933. Type: Ecuador; 

Guayaquil. Humboldt & Bonpland 3803 

(holotype: P-HBK). 

Coarse rhizomatous perennial, 2-4 m tall; 

rhizomes indurate, scaly, 1-6 cm thick, to 30 cm 

long; roots coarse. Culms approximate, erect, firm, 

with a whitish pith, somewhat spongy and 

subterete proximally, obtusely trigonous distally, 

very finely striate, smooth, (0.6-) 0.8-3 cm wide, 

sheathing bases 1-5 cm wide. Leaves 4-6, basal 

and lower cauline, reduced to bladeless sheaths, 

dorsally acute to acuminate; sheaths eligulate, 

elongate, herbaceous, glossy adaxially, open, with 

scarious red-speckled margins, the inner band 

essentially absent, brown. Inflorescence a large, 

decompound, hemispherical, umbel-like corymb 

with ascending rays, (17-) 20-54 × (13-) 18-32 

cm, with numerous spikes of densely aggregated 

spikelets; primary involucral bracts (6-) 8-13 

(-15), 4-45 cm × 4-22 mm, linear-lanceolate, 

flattened to plicate, spreading-ascending to 

horizontal, glabrous except for antrorsely 

appressed-strigose edges of blade both abaxially 

and adaxially, the margins and abaxial midvein 

antrorsely scabrous, at least distally, gradually and 

narrowly acuminate at apex; primary rays 7-25, 

to 32 cm long, secondary rays 5-7, to 18 cm long; 

spikes cylindrical, 1.5-4 × 0.7-1.7 cm, with 24- 

130 spikelets; spikelets linear-lanceolate, 

subcompressed, 3-9 (-11) × 0.8-1.2 mm, acute to 

acuminate at apex, cuneate at base, with 6-19 

florets; rachilla wings light brown, caducous; 

scales ovate-elliptic, dorsally obtuse, 1.3-2.2 × 1- 

1.5 mm, submembranous, laterally nerveless, light 

to reddish brown on sides, margins scarious, carina 

5- to 7-nerved, light green to dark green, prolonged 

beyond the acute to obtuse apex as a slightly 


long, with a blackish, papillate, subulate-tipped 

apex; style 3-branched. Achene subtrigonous with 

the side facing the rachilla concave and the other 

faces convex, oblong, dorsiventrally compressed, 

0.9-1.2 × 0.4-0.5 mm, obtuse at apex, scarcely to 

short-apiculate, sub-rounded at base, estipitate, 

puncticulate, stramineous to pale brown or light 

reddish brown at maturity. 


(Louisiana and Texas), eastern Mexico, Central 


Distribution in Puerto Rico: In water or wet 

grounds along streams and creeks, riverine 

intertidal zones, swales, swampy or marshy areas, 

drainage canals, pond margins, roadside ditches, 

and waste areas. Aguas Buenas, Añasco, Bayamón, 

Cataño, Cidra, Dorado, Loíza, Mayagüez, Río 

Grande, San Juan, Toa Baja, Vega Alta, and 

Yabucoa. 

Common name: Puerto Rico: Junco de 

ciénaga. 


Añasco: Sintenis 5621 (US). Bayamón: Johnston 

913 (US). Cataño: Bo. Palmas, Axelrod & Díaz 

12735 (UPRRP). Cidra: Pueblo Viejo, Stevenson 

554 (US). Dorado: Rd. 165, González-Más 259 

(US). Loíza: At km 2.8 to Loíza on Rt. 951, 

Howard et al. 15943 (US). Mayagüez, Sintenis 

1217 (US). Río Grande: Bo. Herreras, Rd. 187, 

Proctor & Rivera 48357 (US). Yabucoa: Sintenis 

4941b (US). 

19. Cyperus haspan L., Sp. Pl. 45. 1753. 

Lectotype: India. Herbarium Hermann 2: 43, 

No. 37 (BM), designated by McGivney, 

Catholic Univ. Amer., Biol. Series 26(1-17) : 

45. 1938. 

Cyperus juncoides Lam., Tabl. Encycl. 1: 147. 

1791. Cyperus haspan subsp. juncoides 

(Lam.) Kük., Repert. Spec. Nov. Regni Veg. 

23. 184. 1926. Type: North America. Collector 

unknown (holotype: P-Lam.). 

Rhizomatous perennial, 20-132 (-180) cm tall; 

rhizome short, horizontally creeping, 2-6 mm 

thick. Culms forming in a row along the rhizome 

at short intervals, triquetrous to nearly 3-winged,

272 


soft and friable, easily compressed, closely and 

finely ribbed, smooth, (1-) 1.8-5 (-5.5) mm wide, 

sheathing bases 3-8 mm wide. Leaves reduced to 

bladeless sheaths or occasionally 1-3 blades 

present, basal and lower cauline; sheaths thinly 

herbaceous, subtranslucent, reddish or purplish, 

finely and closely veined with dark red veins, the 

bladeless ones oblique at orifice, the blade-bearing 

ones with a convex to truncate orifice; ligule 

absent; blades when present, linear, flattened, Vshaped, 

or often folded (at least proximally) and 

appearing unifacial, 1-20 (-30) cm × 2-8 (-10) mm, 

essentially smooth on margins proximally, 

antrorsely scabrous distally, attenuate to 

triquetrous apex. Inflorescence a compound, open 

or contracted, umbel-like corymb with ascending 

rays, (3-) 4-15 cm in diam.; involucral bracts 2 

(-3), leaf-like, ascending to horizontal, to 18 cm 

long, rarely the lowermost overtopping the 

inflorescence; primary rays 5-16, ascending, short 

to elongate and up to 20 cm long, secondary rays 

sometimes present, to 15 mm long; spikes 

hemispherical or rounded, with subdigitately 

clustered spikelets, 8-20 mm in diam., with (1-) 

3-13 spikelets; spikelets linear-lanceolate to linear, 

3-18 × 1-1.6 (-2) mm, compressed, acuminate at 

apex, short-cuneate at base, with 7-23 (-31) florets; 

rachilla unwinged; scales oblong-ovate, dorsally 

acute, 1.4-1.9 × 0.8-1 mm, submembranous, with 

1 nerve on each side above the narrowly scarious 

margins, reddish to greenish brown or stramineous 

on sides, carina 1-nerved, greenish, prolonged as 

a short prickly mucro at the subacute apex. 

Stamens 3, the anthers 0.3-0.6 mm long, often with 

a tuft of minute crystalline trichomes at apex; style 

3-branched. Achene trigonous to very obtusely so, 

with convex faces, subglobose to obovoid or 

broadly ellipsoid, 0.5-0.8 × 0.3-0.5 mm, obtuse to 

acute at apex, apiculate to entire, cuneate at base, 

with small basal torus, granular to papillose, 

lustrous-white or pale brown, sometimes reddish 

brown. 


Distribution in Puerto Rico: In wet or 

saturated sandy soils along roadsides, ditches, 

pastures, grasslands, pond borders, marshy areas, 

and waste places. Canóvanas, Carolina, Cayey, 

Ceiba, Cidra, Dorado, Guayama, Humacao, 

Luquillo, Manatí, Mayagüez, Naguabo, Río 

Grande, San Juan, San Sebastián, Utuado, Vega 

Alta, and Vega Baja. 


Canóvanas: Sierra de Luquillo, Proctor & Taylor 

46298 (US). Carolina: Lago San José, Hioram 229 

(US). Cayey: Sierra de Cayey, Carite Forest, 

Proctor & Milety 43962 (US); Guavate State 

Forest, Liogier 10359 (US). Humacao: Pozal, 

Playa de Humacao, Eggers 707 (US). Mayagüez: 

Las Mesas, Holm 222 (US). Naguabo: Sierra de 

Naguabo, Río Icaco and adjacent hills, Shafer 3469 

(US); Río Grande: Sierra de Luquillo, Carribean 

National Forest, Proctor & Thomas 43198 (US). 

San Juan: Martín Peña, Stevenson 539 (US). Río 

Grande: El Yunque, Sargent 304 (US). San 

Sebastián, Sargent 245 (US). Utuado, Sargent 

3277 (US). 

20. Cyperus imbricatus Retz., Observ. Bot. 5: 

12. 1788. Type: India. Koenig s.n. (holotype: 

LD; isotype: C). 

Cyperus radiatus Vahl, Enum. Pl. 2: 369. 1805. 

Lectotype: Africa; Guinea. 1784, Isert s.n. (C- 

Vahl), designated by G. C. Tucker & McVaugh 

in McVaugh, Fl. Novo-Galiciana 13: 304. 

1993. 

Cyperus radiatus var. elongatus Boeck., Linnaea 

36: 317. 1870. Cyperus imbricatus var. 

elongatus (Boeck.) Kük. in Engler, Pflanzenr. 

IV. 20 (Heft 101): 71. 1936. Type: Philippine 

Islands. Cuming 437 (holotype: B, destroyed). 

Coarse, rhizomatous perennial, 70-150 cm 

tall; rhizomes short, 1-3 cm long, 5-10 mm thick, 

hardened; roots coarse. Culms erect, trigonous, 

often subtriquetrous distally, firm, coarsely ribbed, 

smooth, (3-) 3.5-10 mm wide, sheathing bases 1- 

3 cm wide. Leaves 3-7, basal and cauline; sheaths 

eligulate, spongy-thickened and purple-black 

proximately, fading to brown streaked with black 

distally; ligule absent; blades linear, folded to Vshaped 

proximally, plicate distally, 35-90 cm × 4- 

15 (-18) mm, with numerous cross veinlets, 

antrorsely scabrous on the margins, abaxial 

midvein, and adaxial lateral veins, long-attenuate 

to triquetrous apex. Inflorescence a compound 

umbel-like corymb with ascending rays, 12-30 × 

14-30 (-40) cm; involucral bracts 5-10, leaf-like, 

spreading, ascending to horizontal, the lowermost 

to 90 cm long; rays 6-12, to 25 cm long; spikes 

linear-cylindric, 1-6 (-8) cm × 3-10 (-15) mm, in 

subradiate groups of (1-) 3-20 at ray tips, with 

(20-) 30-130 (-160) densely disposed spikelets;


spikelets ascending initially, divergent with age, 

ovate-lanceolate to oblong-lanceolate, 

compressed, often slightly twisted, 3-6 × 1-1.4 

mm, acute to obtuse at apex, obtuse at base, with 

8-22 florets; rachilla unwinged; scales ovate, 1- 

1.3 (-1.5) × 0.7-1 mm, dorsally obtuse, 

submembranous, with 2-3 indistinct nerves on 

each side above the scarious margins, stramineous 

to yellowish or reddish brown, carina 3-nerved, 

greenish, prolonged beyond the obtuse to subrounded 

apex, as a short, straight to slightly 

excurved mucro. Stamens 3, the anthers 0.2-0.5 

mm long, apiculate; styles 3-branched. Achene 

trigonous, dorsiventrally compressed, with the 

adaxial face plane and the abaxial faces broadly 

rounded, ellipsoid to ellipsoid-obovoid, 0.5-0.6 × 

0.3-0.4 mm, obtuse to sub-rounded at apex, 

apiculate, sub-rounded at base, short-stipitate, very 

finely puncticulate to essentially smooth and 

glossy at maturity, dull whitish to stramineous. 



Distribution in Puerto Rico: In wet or 

saturated soils of low, swampy habitats, lake 

margins, edges of rivers and canals, old fields, 

roadsides, lawns, and disturbed areas, often 

emergent. Aguada, Aibonito, Añasco, Bayamón, 

Carolina, Cataño, Guayama, Humacao, Mayagüez, 

Naguabo, Quebradillas, San Juan, San Sebastián, 

Trujillo Alto, Vega Alta, and Vega Baja. 


Aguada: Sargent 567 (US). Añasco: Heller 4536 

(US). Bayamón: Bo. Minillas, along E side of Río 

Hondo, Proctor 43553 (US). Carolina, Stevenson 

& Johnston 1390 (US). Humacao: Playa de 

Humacao, Eggers 682 (US). Mayagüez, Sintenis 

126 (US). Vega Alta: Bo. Sabana, ca. 1.8 km due 

NE of jct. Rds. 690 and 691, Proctor 45087 (US). 

21. Cyperus involucratus Rottb., Descr. Pl. Rar. 

22. 1772; Cyperus flabelliformis Rottb., 

Descr. Icon. Pl. Rar. 42. 1773, nom. illeg.; 

Cyperus alternifolius subsp. flabelliformis 

(Rottb.) Kük. in Engler, Pflanzenr. IV. 20 

(Heft 101): 193. 1936. Lectotype: Arabia. 

Forsskål s.n. (holotype: C-Rottb.), designated 

by Baijnath, Kew Bull. 30: 522. 1975. 

Cyperus alternifolius of many authors, non 

Linnaeus, 1771. 


tall; rhizomes horizontally creeping, stout, 10-20 

mm thick, woody. Culms solitary, crowded to 

narrowly spaced and forming in a row along the 

rhizome, erect, trigonous to obtusely trigonous or 

subterete proximally, finely ribbed, smooth, 

sometimes scabrid at apex, 3-9 mm wide, 

sheathing bases 7-23 mm wide. Leaves basal and 

lower cauline, reduced to bladeless sheaths; upper 

sheaths elongate, pale green, obliquely truncate 

at orifice, dorsally acute at apex, the lower 

cataphylloid, yellow-brown or brown. 

Inflorescence a compound umbel-like corymb with 

ascending rays, 5-16 × 5-25 (-30) cm; involucral 

bracts 12-25, foliaceous, linear-lanceolate, 

flattened, 9-30 (-35) cm × (2-) 4-18 mm, subequal 

in length, horizontally spreading, appearing 

spirally disposed, antrorsely scabrous on the 

margins and abaxial midvein, abruptly acute to 

acuminate at apex; primary rays 9-16, to 20 cm 

long, secondary rays 7-12, to 30 mm long; spikes 

in subglobose or lobate clusters, 7-20 (-30) mm in 

diam., disposed in digitate clusters of 7-35 

spikelets at ray tips; spikelets ovate to lanceolate 

or oblong-lanceolate, strongly compressed, 4-10 

× 1.8-2.2 mm, acute at apex, short-cuneate at base, 

with 8-34 florets; rachilla unwinged; scales ovatedeltate, 

1.6-2.4 × 1.2-1.7 mm, obtusely curvatekeeled, 

bicarinate basally, membranous and 

translucent, lateral nerves indistinct above the 

scarious margins, whitish stramineous or light 

brown to brown on sides proximally, carina 3nerved, 

light brown, prolonged beyond the acute 

apex as a short mucro,. Stamens 3, the anthers 0.7- 

1 mm long, with prolonged subulate, papillose 

apex; style 3-branched. Achene plano-convex to 

subtrigonous, with the adaxial face plane or 

slightly convex and the abaxial faces convex, 

broadly ellipsoid, 0.6-0.8 × 0.4-0.6 mm, shortapiculate 

at obtuse apex, sessile or short-stipitate 

at abruptly cuneate base, puncticulate, brown. 

General distribution: Native to Arabia and 

tropical East and South Africa; escaping 

cultivation and naturalized in the New World from 

the southern United States (Florida, Louisiana, 

Texas, Arizona, and southern California) Hawaii, 

West Indies, Mexico, Central America, South 

America, and in the Old World tropics. 


Islands: Wet areas of creek and stream borders, 

riverbanks, swampy areas, pond margins, small 

pools, secondary forest, roadside ditches, fields, 

pastures, and disturbed areas. Cultivated and often 

used as an ornamental, naturalized. Adjuntas,

274 


Aibonito, Arecibo, Cayey, Coamo, Guánica, 

Guayama, Guayanilla, Naguabo, Isabela, Jayuya, 

Juana Díaz, Maricao, Mayagüez, Naguabo, 

Peñuelas, Ponce, San Germán, Santa Isabel, San 

Juan, Río Grande, Trujillo Alto, Utuado, Vega 

Baja, and Yauco; St. Croix, St. John, and St. 

Thomas. 

Common name: Puerto Rico: Paragüita. 


Adjuntas: Rd. 10, km 23.5, along creek, González- 

Más 875 (US). Arecibo: Río Abajo State Forest, 

Acevedo-Rdgz. 10728 (MAPR, NY, UPRRP, US). 

Guayanilla: Rd. 336, González-Más 813 (US). 

Jayuya: Toro Negro Forest Reserve, Acevedo- 

Rdgz. & Angell 9443 (FTG, NY, UPR, UPRRP, 

US). Mayagüez: Guanajibo, González-Más 822 

(US). Naguabo: Bo. Río, along estuary of the Río 

Blanco, Proctor & Rodríguez 42847 (US). San 

Juan: Río Piedras, Stevenson 5115 (US). Utuado: 

Vicinity of Utuado, N.L. Britton & Cowell 902 

(US). Yauco: banks of Río Loro, Sargent 669 (US). 

ST. THOMAS: Raphune, Acevedo-Rdgz. 11347 (NY, 

UPRRP, US). 

22. Cyperus iria L., Sp. Pl. 45. 1753. Lectotype: 

India. Osbeck s.n. (LINN-70.16), designated 

by G. C. Tucker, Syst. Bot. Monogr. 43: 91. 

1994. 

Caespitose annual or short-lived perennial, 

(5-) 12-60 (-75) cm tall. Culms erect, loosely 

caespitose, triquetrous, stiff and hardened, ribbed, 

smooth, (1-) 1.2-3 mm wide, sheathing bases (2-) 

3-6 (-8) mm wide. Leaves (1-) 3-5, basal and lower 

cauline; sheaths elongate, pale green to pale 

brown, the lowermost reddish tinged, reddish 

lineolate; ligule absent; blades linear, V-shaped 

proximally, subflattened to plicate distally, (3-) 8- 

30 (-40) cm × (1.5-) 2-6 mm, antrorsely scabrous 

on the margins and abaxial midvein, at least 

distally, attenuate to triquetrous apex. 

Inflorescence a simple to compound umbel-like 

corymb with ascending rays, 2.5-15 (-19) × 2.5- 

18 cm; involucral bracts (3-) 4-8 (-9), leaf-like, 

ascending to spreading, to 2-40 (-50) cm long; 

primary rays 4-8, to 12 cm long, secondary lateral 

rays absent or very short; spikes oblong to 

narrowly ovoid, 2-6 (-8) at ray tips, 1-5 cm × 2.2- 

11 (-15) mm, with 5-26 (-30) loosely disposed 

spikelets; spikelets linear-lanceolate, compressed, 

4-12 (-20) × 1.5-2 mm, obtuse at apex, short- 

cuneate at base, with 6-22 (-26) florets; rachilla 

unwinged; scales broadly elliptic to ellipticobovate 

or subrounded, 1.3-1.8 × 1.2-1.8 mm, 

dorsally acute, curvate-keeled, submembranous, 

with 1 lateral nerve on each side above the scarious 

margins, brown to golden brown, finely cellularreticulate, 

carina 3-nerved, green, slightly shorter 

than to equaling the scarious, obtuse, emarginate 

apex as a short mucro. Stamens (1-) 2, the anthers 

0.2-0.4 mm long, bluntly apiculate; style 3branched. 

Achene trigonous, with concave faces, 

narrowly obovoid, 1.2-1.4 × 0.5-0.8 mm, obtuse 

to sub-rounded at apex, apiculate, short-acuminate 

at base with a striate, crystalline basal torus, 

puncticulate, yellowish to dark brown. 

General distribution: Native to the Old World, 

introduced and now naturalized in the southeastern 

United States, West Indies, Mexico, Central 

America, and South America. 

Distribution in Puerto Rico: In wet soils 

bordering rivers and canals, swales, meadows, 

pastures, agricultural fields, roadside ditches, and 

waste places. Añasco, Arecibo, Bayamón, Caguas, 

Carolina, Cataño, Ceiba, Dorado, Gurabo, 

Humacao, Juncos, Lajas, Loíza, Maunabo, 

Mayagüez, Naguabo, Río Grande, San Juan, San 

Lorenzo, Toa Alta, Vega Baja, and Vieques. 


Bayamón: Bo. Minillas, along E side of Río 

Hondo, Proctor 43558 (US). Caguas: Bo. Bairoa, 

along Río Bairoa just NE of Highway 30 bridge, 

Proctor 46118 (US). Carolina: 65 Inf. Road Río 

G. de Loíza, González-Más 1231 (US). Gurabo: 

At Sub-Station Gurabo, Woodbury et al. s.n. (US). 

Humacao: Mariana, González-Más 1351 (US). 

Lajas: Cartagena Lake, González-Más 949 (US). 

Mayagüez: Miradero’s Road, km 1.15, González- 

Más 881 (US). Naguabo: Bo. Río Blanco, Axelrod 

11018 (UPRRP). San Juan: Bo. Hato Rey Norte, 

along Kennedy Ave. (Highway 2), Proctor 43034 

(US). 

23. Cyperus laevigatus L., Mant. Pl. 2: 179. 1771; 

Pycreus laevigatus (L.) Nees, Linnaea 10: 

130. 1836; Juncellus laevigatus (L.) C. B. 

Clarke in Hooker f., Fl. Brit. India 6: 596. 

1893. Lectotype: South Africa; Cape of Good 

Hope. Koenig s.n. (LINN-70.13), designated 

by G. C. Tucker & McVaugh in McVaugh 

(ed.), Fl. Novo-Galiciana 13 : 308. 1993.



rhizome horizontally creeping, 1-4 mm thick. 

Culms solitary or clustered along the rhizome at 

short or sometimes long intervals, obtusely 

trigonous to subterete, subspongy, coarsely ribbed 

and channeled, smooth, 0.5-2.7 (-3) mm wide, 

sheathing bases 1-6 mm wide. Leaves 2-3, basal 

and lower cauline, reduced to bladeless sheaths 

or the uppermost with short blades; sheaths reddish 

brown or dark brown, with cross veinlets, the 

bladeless ones obliquely truncate at orifice, bladebearing 

sheaths with a membranous U-shaped 

orifice; ligule absent; blades when present 

subulate, involute or convolute, often canaliculate 

distally, 1-10 cm × 0.3-1 (-1.6) mm, smooth on 

margins, midvein indistinct, acuminate or shortacuminate 

to an often bent callous tip, glaucousgreen, 

smooth, with cross veinlets. Inflorescence 

a pseudolateral capitate head of 1-22, loosely 

subdigitate spikelets at the summit of the culm, 4- 

16 (-22) mm in diam.; involucral bracts (1-) 2 

(-3), leaf-like, the lowermost one elongate, erect, 

appearing as a continuation of the culm, to 7 cm 

long, the upper ones short, ascending to horizontal 

or reflexed at maturity; rays not evident; spikelets 

ovate to oblong-ovate or oblong-lanceolate, 4-11 

(-15) × (1.6-) 1.8-2.8 mm, compressed-turgid, 

subacute at apex, sub-rounded at base, with 10-40 

(-56) florets; rachilla broad, appearing scaly on 

margins from the persistent decurrent bases of the 

shed scales; scales broadly ovate or subrounded, 

1.5-2 × 1.5-2.2 mm, thinly herbaceous, with 3-4 

indistinct nerves on each side above the broad 

scarious margins, whitish or stramineous, 

distinctly reddish lineolate adaxially, indistinctly 

so abaxially, carina 3-nerved, light greenish, 

prolonged as a short mucro at the obtuse to subrounded 

apex, invaginated and 2-keeled abaxially 

at base. Stamens (2-) 3, the anthers 0.6-1.2 mm 

long, with a lanceolate-subulate connective 

bearing crystalline papillae; style 2-branched, 

reddish lineolate. Achene lenticular, dorsiventrally 

compressed, with a plane to concave adaxial face 

and convex abaxial face, oblong-elliptic to ovateelliptic, 

1.1-1.5 × 0.7-1 mm, obtuse at apex, 

apiculate, short-cuneate at base, estipitate, the 

surface finely reticulate, glossy, grayish brown or 

dark brown to blackish. 

General distribution: Pantropical; in the New 

World occurring in the southwestern United States 

(Texas, Arizona, California, and Nevada), West 

Indies, Mexico, and South America. 


Islands: In coastal areas on wet soils of lake and 

pond borders, marshes, ditches, and pastures. 

Aibonito, Coamo, Guánica, Guayama, Guayanilla, 

Mayagüez, Ponce, Salinas, San Juan, and Santa 

Isabel; St. Croix, St. Thomas, and Virgin Gorda. 


Coamo: Sintenis 2984 (US). Guánica: Sintenis 

3847 (US). Guayama: Pto. Jobos, Rd. 707, 

González-Más 703 (US). Guayanilla: N.L. Britton 

& Shafer 1792 (US). Ponce: Rd. 2, km 16.6, Las 

Cucharas, González-Más 802 (US). Salinas: 

Aguirre, N.L. Britton et al. 6031 (US). Santa 

Isabel: Playita Cortada Sign, Rd. 1, González-Más 

746 (US). ST. CROIX: Bassin Lagoon, A.E. 

Ricksecker 54 (US). ST. THOMAS: at Bolongo, 

Eggers s.n. (US). 

24. Cyperus ligularis L., Syst. Nat. ed. 10, 2: 867. 

1759; Mariscus ligularis (L.) Urb., Symb. 

Antill. 2: 165. 1900. Lectotype: West Indies; 

Jamaica. Browne s.n. (LINN-70.37), 

designated by G. C. Tucker, Syst. Bot. 

Monogr. 2: 49. 1983. 

Mariscus rufus Kunth in Humb., Bonpl., & Kunth, 

Nov. Gen. Sp. 1 [quarto ed.]: 216. 1816. Type: 

Mexico; Michoacán. Humboldt & Bonpland 

s.n. (holotype: P-HBK). 

Cyperus sintenisii Boeck., Beitr. Cyper. 1: 12. 

1888. Type: Puerto Rico, Sintenis 4952 


Cyperus trigonus Boeck., Beitr. Cyper. 1: 11. 1888. 

Type: Puerto Rico, Sintenis s.n. [collection 

number not designated in protologue] 


Fig. 48. A-D 

Rhizomatous perennial, sometimes 

caespitose, (40-) 50-120 (-190) tall; rhizome short, 

stout, 1-2 cm thick; roots coarse. Culms solitary 

or sometimes two together, obtusely trigonous to 

sub-rounded proximally, often trigonous at apex, 

light green to glaucous, minutely papillose, 3-6 

mm wide, sheathing base of culm stout, 1-3 cm 

wide. Leaves 5-10, crowded basally, sometimes 

1-3 lower cauline; sheaths elongate, closely 

overlapping at base, conspicuously cross-veined, 

reddish to purplish brown proximally; ligule 

absent; blades flattened to subplicate, rarely the 

margins somewhat inrolled, 30-120 cm × 5-15 

mm, conspicuously cross-veined, margins and 

abaxial midvein harshly antrorsely scabrous, long-

276 


acuminate to triquetrous apex. Inflorescence an 

open to subcontracted, simple to compound, 

umbel-like corymb with ascending rays, 5-25 cm 

diam.; involucral bracts 6-9, leaf-like, ascending, 

the lowermost ones elongate, to 90 cm long, the 

uppermost ones short, linear to subulate; primary 

rays 5-12, unequal, to 20 cm long; secondary rays 

short, inconspicuous; spikes oblong-cylindrical to 

subglobose, 5-30 × 8-15 mm, 3-7 congested at ray 

tips or solitary, the terminal one erect and elongate, 

the lateral ones (when present) short, divergent, 

with 15-300 or more densely disposed spikelets; 

spikelets oblong-elliptic, subcylindrical to 

subcompressed, 3-8 × 1-2 mm, disarticulating at 

base, falling entire, with 3-8 florets; rachilla 

narrowly hyaline-winged; scales ovate, dorsally 

obtuse to rounded, boat-shaped, 2-3 × 1.5-2 mm, 

3- to 5-nerved on each side above the broadly 

scarious margins, reddish to reddish brown, carina 

inconspicuous, finely 3-nerved, green, slightly 

prolonged beyond the acute apex as a short mucro. 

Stamens 3, the anthers 0.5-1 mm long, apiculate; 

style 3-branched from just above the base. Achene 

trigonous with concave faces, obovoid to broadly 

ellipsoid, slightly curvate on adaxial side, 1.2-1.8 

× 0.6-0.8 mm, acute at apex, apiculate, cuneate at 

base, estipitate, puncticulate, red to reddish brown 

or blackish at maturity. 


States (Florida), West Indies, Mexico, Central 

America, South America, tropical West Africa, 

Chagos Archipelago (Indian Ocean), Seychelles, 

and western Pacific islands and atolls. 


Islands: Sandy soils of coastal habitats in 

savannas, marshy or swampy areas, borders of 

mangrove swamps, margins of ponds and lakes, 

thickets, scrub forest, beaches, fields, pastures, 

roadsides, and waste areas. Aguada, Aibonito, 

Arecibo, Arroyo, Bayamón, Cabo Rojo, Camuy, 

Carolina, Cataño, Cayey, Cayo Icacos, Cayo 

Ramos, Ceiba, Coamo, Culebra, Dorado, Fajardo, 

Guayama, Guayanilla, Humacao, Isabela, Isleta 

Marina, Loíza, Luquillo, Manatí, Maunabo, 

Mayagüez, Mona Island, Naguabo, Patillas, 

Peñuelas, Ponce, Río Grande, Salinas, San Juan, 

Santa Isabel, Toa Baja, Utuado, Vega Alta, Vega 

Baja, Vieques, and Yabucoa; St. Croix, St. John, 

St. Thomas, and Tortola. 

Common name: Puerto Rico: Junco de agua. 


Arecibo: Sargent 682 (US). Bayamón: 2 km W of 

Bayamón, McKee 10596 (US). Cabo Rojo: 

Boqueron, González-Más 930 (US). Cataño: 

Heller & Heller 35 (US). Cayey: Reserva Forestal 

de Guavate-Carite, Stimson 1726 (US). Coamo: 

near hot-spring, Underwood & Griggs 532 (US). 

Culebra Island: N.L. Britton & Wheeler 183 (US). 

Guayama, Kuntze 573 (US). Guayanilla: Rd. 336, 

González-Más 820 (US). Humacao: Santa Teresa, 

Liogier et al. 31335 (US); lecheria, Goll 24 (US). 

Mayagüez: Sintenis 1218 (US). Mona Island: 

Sardinera, Britton et al. 1833 (NY). Ponce: Bo. 

Cañas, along dirt road to Punta Cucharas, Axelrod 

& Chávez 7330 (US). San Juan: Martín Peña, 

Stevenson 266 (US). Santa Isabel: Rd. 1, km 110.5, 

González-Más 753 (US). Vega Baja: Pto. Nuevo, 

Rd. 686, km 4, González-Más 1106 (US). Vieques 

Island: Resolución to Punta Arenas, Shafer 2900 

(US). ST. CROIX: Salt River, Ricksecker 107 (US). 

ST. JOHN: Coral Bay Quarter; along dirt road to 

Bordeaux Mountain, Acevedo-Rdgz. 5213 (NY, 

UPRRP, US). ST. THOMAS: Pease 22914 (US); 

Jumbee Gut, Eggers s.n. (NA). 

25.Cyperus mutisii (Kunth) Andersson, 

Galapagos Veg . 53. 1854; Mariscus mutisii 

Kunth in Humb., Bonpl., & Kunth, Nov. Gen. 

Sp. 1 [quarto ed.]: 216, pl. 66. 1816. Type: 

Colombia; Near Bogotá. Mutis s.n. (holotype: 

P; possible isotype: Mutis 338, US!). 

Cyperus ochreatus Boeck., Linnaea 36: 386. 1870. 

Type: Dominican Republic. Meyerhoff 218 


Cyperus semitribrachiatus Boeck., Linnaea 36: 

386. 1870. Cyperus mutisii var. 

semitribrachiatus (Böck) Kük. in Engler, 


Dominican Republic. Meyerhoff 217 


Cyperus compresso-triqueter Boeck., Beitr. Cyper. 

1: 10. 1888. Type: Puerto Rico, vicinity of 

Coamo. Sintenis s.n. (holotype: B, destroyed; 

isotype: B). 

Cyperus martinicensis Boeck., Beitr. Cyper. 2: 38. 

1890. Cyperus mutisii var. martinicensis 

(Boeck.) Kük. in Engler, Pflanzenr. IV. 20 

(Heft 101): 483. 1936. Type: Martinique. Duss 


Mariscus incompletus sensu Urban, Symb. Antill. 

4: 113. 1903, non (Jacquin) Urban, 1900. 


cm tall; rhizome short, knotty, thickened, 0.8-15


mm thick. Culms triquetrous, sometimes trigonous 

proximally, hardened, finely ribbed, smooth or 

scabridulous on angles distally, (1.4-) 1.8-5 mm 

wide, sheathing bases 3-10 mm wide. Leaves 2-6, 

basal and cauline; sheaths elongate, often crossveined 

distally, brown, purplish brown proximally; 

ligule absent; blades linear, plicate, 6-50 (-70) cm 

× 4-10 (-13) mm, scabridulous adaxially (at least 

distally), smooth abaxially, antrorsely scabrous on 

the margins and abaxial midvein, long-acuminate 

to narrowly plicate apex. Inflorescence a simple, 

open or contracted, umbel-like corymb with 

ascending rays, 6-20 (-32) × 5-18 (-22) cm; 

involucral bracts 5-9, leaf-like, ascending, the 

lowermost 20-50 cm long; rays 3-7 or absent, 2- 

26 cm long; spikes, cylindrical, solitary or in 

digitate clusters of 3-6 ray tips, the lateral ones 

shorter, 1-6 cm × 5-11 mm, with (-8) 17-140 

(-160) densely disposed spikelets which are each 

subtended by a linear bract at base that is often 

shorter than the spikelet; spikelets oblongellipsoid, 

2.5-7 × (0.9-) 1.2-2 mm, slightly 

compressed to subquadrate, or broadly elliptic in 

cross section, obtuse at apex, short-cuneate at base, 

with 1-3 (-5) florets; rachilla broadly hyalinewinged; 

scales ovate to ovate-elliptic, 2.2-3.4 × 

1.5-2.1 mm, thinly herbaceous, coarsely (3-) 4-5 

nerved on each side above the scarious margins, 

light brownish to brown or reddish brown, dark 

brown lineolate, at least on margins, carina 3nerved, 

greenish, equaling or prolonged beyond 

the obtuse apex as a short mucro. Stamens 3, the 

anthers 0.5-1 mm long, short or scarcely apiculate; 

style 3-branched. Achene trigonous, with the 

adaxial face concave and the abaxial faces plane 

to slightly concave, adaxially curvate, slightly 

dorsiventrally compressed, ellipsoid to broadly 

ellipsoid or oblong-ellipsoid, 1.5-1.8 × 0.7-0.9 

mm, obtuse at apex, apiculate, short-cuneate to 

obtuse at base, short-stipitate, puncticulate, brown 

to reddish brown. 


(Arizona), West Indies, Mexico, Central America, 


Distribution in Puerto Rico: Grassy slopes, 

hillsides, and roadbanks. Jayuya, Peñuelas, Ponce, 

Sabana Grande, Villalba, and Yauco. 


Jayuya: Rd. 140 to Doña Juana, km 9, González- 

Más 2185 (US). Ponce: On the Adjuntas road 11 

mi. from Ponce, Heller 6358 (US). Yauco: Bo. 

Sierra Alta, upper slopes and summit area of Pico 

Rodadero, Proctor & Rivera 48004 (US). 

26. Cyperus nanus Willd., Sp. Pl. 1: 272. 1797. 

Type: “Habitat in Guinea” (from Martinique 

fide Vahl, Enum. Pl. 2: 372. 1805) Isert s.n. 

(holotype B-Willd. 1419). 

Schoenus capillaris Sw., Prodr. 20. 1788, [non 

Cyperus capillaris Koenig ex Roxb., 1820]; 

Mariscus capillaris (Sw.) Vahl, Enum. Pl. 2: 

372; 1805. Kyllinga capillaris (Sw.) Griseb., 

Syst. Veg. Karaiben. 120. 1857; Cyperus 

tenuis var. capillaris (Sw.) Kük., Repert. Spec. 

Nov. Regni Veg. 23: 188. 1926. Type: 

Hispaniola. Swartz s.n. (holotype: S-Sw. R- 

5607). 

Fig. 48. E-H 

Slender, caespitose perennial, 2-35 cm tall; 

rhizome short, knotty; roots filiform. Culm 

ascending, often arching, trigonous proximally, 

triquetrous distally, wiry, 0.3-0.4 mm wide, 

sparsely antrorsely scabrous on angles near apex. 

Leaves 2-5, basal and lower cauline, the lowermost 

reduced to bladeless sheaths; sheaths short, 

uppermost elongating to 3 cm, submembranous, 

reddish brown to dark brown proximally; ligule 

absent; blades narrowly linear, folded to V-shaped 

proximally, subflattened or plicate distally, 1-30 

cm × 0.3-1.6 mm, antrorsely scabrous on margins 

and abaxial midvein, attenuate to triquetrous apex. 

Inflorescence a single, globose to widely-ellipsoid 

spike at the summit of the culm, 4-8 × 3-7 mm, 

with (3-) 5-30 spikelets, rarely with an additional 

lateral spike; involucral bracts 2-3, leaf-like, 

reflexed or the lowermost one often ascending at 

anthesis, 1-8 cm long; spikelets oblong-ovate to 

oblong-lanceolate, subcompressed, 2-4 × 0.8-1.2 

mm, bearing 3-5 florets, the terminal flower 

staminate or wanting, reflexed at maturity, 

sometimes the terminal spikelet remaining erect, 

disarticulating at base, falling entire, the scales 

spreading with maturing achenes; rachilla 

narrowly hyaline-winged; scales ovate to broadly 

ovate-elliptic, 1.2-1.6 × 0.8-1 mm, dorsally acute, 

coarsely 3- to 4-nerved above the scarious margins, 

green to light reddish brown, carina 3-nerved, dark 

green to brown, prolonged beyond the acute to 

obtuse apex as a short mucro. Stamens 2 or 

sometimes 1, the anthers oblong-elliptic, 0.3-0.5 

mm long, rounded-apiculate; style 3-branched

278 


Fig. 48. A-D. Cyperus ligularis. A. Habit, inflorescence, and detail of abaxial surface of leaf blade. B. Portion of inflorescence 

spike rachis with spikelets. C. Spikelet. D. Flower and subtending spikelet scale. E-H. Cyperus nanus. E. Habit. F. Inflorescence. 

G. Spikelet. H. Spikelet scale, flower and achene. I-K. Cyperus elegans. I. Inflorescence. J. Spikelet. K. Flower and spikelet 

scale. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78.


from just above the base. Achene trigonous with 

plane to slightly concave faces, the adaxial face 

straight, oblong-ellipsoid, 1-1.2 × 0.5-0.6 mm, 

obtuse at apex, apiculate, short-cuneate at base, 

estipitate, puncticulate, light brown to purplish 


General distribution: Throughout the West 

Indies and Mexico. 


Islands: Sandy, shaded, often moist areas on grassy 

slopes, secondary thickets, hillsides, rocky woods, 

ridges, scrub thickets, trailsides, and disturbed 

areas. Cabo Rojo, Mayagüez, Mosquito Island, 

Palomino Island, and Vieques; Anegada, Big 

Tobago, Guana Island, St. Croix, St. John, Tortola, 

Virgin Gorda, and Whistling Key. 


Cabo Rojo: Sierra Bermeja, Bo. Llanos Costa, 

Proctor & McKenzie 44022 (US). Fajardo: 

Palominos Island, Woodbury s.n. (US). Mayagüez: 

near San Antonio Hospital, González-Más 2181 

(MAPR, US). ST. JOHN: Cruz Bay, Raunkiaer s.n. 

(US); Along trail to Reef Bay by petrogliph, 

Acevedo-Rdgz. et al. 2929 (NY, UPR, UPRRP, 

US). TORTOLA: Little Camanoe Island, Proctor 

46512 (US). 

27. Cyperus ochraceus Vahl, Enum. Pl. 2: 325. 

1805. Type: St. Croix; U.S. Virgin Islands. 

West 15 (holotype: C-Vahl; photo at US; 

isotype: C-Schum.). 

Cyperus ochraceus var. excelsior Kük. in Engler, 

Pflanzenr. IV. 20 (Heft 101): 182. 1936. 

Lectotype: Bolivia, Villa Montes, Prov. Tarija, 

Pflanz 651 (B), designated by Denton, Contr. 

Univ. Michigan Herb. 11: 234. 1978. 

Cyperus ochraceus var. minor Kük. in Engler, 


Puerto Rico; near Guánica. Sintenis 3839 

(holotype: B). 

Caespitose perennial, (5-) 10-75 (-102) cm 

tall; rhizome short, horizontal to ascending, 3-10 

mm thick. Culms obtusely trigonous or 

subtriquetrous distally, slender, hardened and stiff, 

smooth, (1-) 2-4 (-5) mm wide, narrowing 

subabruptly at apex; sheathing bases 3-12 mm 

wide. Leaves 3-7, primarily basal, 1-2 lower 

cauline; sheaths short, thickly herbaceous and 

stiffened, pale greenish or stramineous, often 

tinged with brown; ligule absent; blades U- or V- 

shaped or subfolded proximally, subplicate 

distally, (5-) 10-50 (-72) cm × (1.5-) 2-8 (-9) mm, 

substiffened, semi-glossy and finely cellularreticulate 

adaxially, finely vined abaxially, smooth 

or remotely antrorsely scabrous on the margins 

and abaxial midvein, the apex long-acuminate to 

triquetrous apex. Inflorescence a simple to 

compound, open or contracted, umbel-like corymb 

with ascending rays, (1-) 2-15 × (1.3-) 2.2-16 cm; 

involucral bracts (4-) 5-8, leaf-like, ascending to 

horizontal, the lowermost to 30 (-50) cm long; 

primary rays (3-) 4-13, ascending, elongating up 

to 12 cm, several short secondary rays often 

present; spikes glomerulate, spherical to 

hemispherical, (0.6-) 1-3 cm diam., with (4-) 6- 

40 (-50) densely disposed, digitately arranged 

spikelets; spikelets oblong to linear-oblong, (4-) 

6-14 × 2-3 mm, compressed, obtuse at apex, 

broadly obtuse at base, with 10-36 florets; rachilla 

unwinged; scales broadly ovate, (1.2-) 1.5-2 × 1.2- 

2 mm, boat-shaped, dorsally 2-keeled proximally, 

thickly herbaceous, glossy, spongy-thickened 

proximally, faintly rugulose-reticulate, yellowish 

brown or stramineous, reddish lineolate adaxially, 

with 1 indistinct nerve on each side above the 

scarious margins, carina 3-nerved, the lateral 

nerves forming two pronounced keels proximally 

producing a declivity between the two, at the base 

of which lies the midnerve, all three nerves 

converging into a single one at the acute apex, there 

forming a short mucronate-cuspidate, slightly 

recurved apex, gray-green. Stamen 1, the anther 

0.7-1.2 mm long, apiculate; style 3-branched. 

Achene obtusely trigonous, with slightly convex 

faces, ellipsoid to broadly ellipsoid, 1-1.5 × 0.4- 

0.7 mm, short-acuminate at apex, gradually 

narrowing to a short, subulate beak, acute to 

subcuneate at base, stipitate, finely cellular- or 

rugulose-reticulate, reddish brown, maturing dark 

purplish brown. 

General distribution: Southern United 

States, West Indies, Mexico, Central America, 



Islands: Wet places in swampy or marshy areas, 

swales, stream or river edges, pond or lake 

margins, agricultural fields, pastures, roadside 

ditches, and waste areas. Añasco, Arecibo, Arroyo, 

Bayamón, Cabo Rojo, Camuy, Carolina, Cataño, 

Cayey, Coamo, Dorado, Florida, Guánica, 

Guayama, Guayanilla, Isabela, Ponce,

280 


Quebradillas, Santa Isabel, Toa Alta, Toa Baja, and 

Vega Baja; St. Croix (type locality). 


Arecibo: Liogier & Martorell 34872 (US). 

Bayamón: Heller & Heller 409 (US). Cabo Rojo: 

Bo. Llanos Costa, Cabo Rojo National Wildlife 

Refuge, Proctor & McKenzie 43990 (US). Cataño: 

Rd. 165, km 5, González-Más 1151 (US). Coamo: 

Coamo Springs, E.G. Britton & Marble 2317 (US). 

Guánica: Guánica Lake, Sargent 40 (US). 

Guayama: Pto. Jobos, Rd. 709, González-Más 657 

(US). Guayanilla: N.L. Britton 1790 (US). Ponce: 

Two miles W of Ponce, Heller 6145 (US). Santa 

Isabel: Rd. 1, km 110.5, González-Más 756 (US). 

Vega Baja: Bo. Cibuco, just SW of Playa Cerro 

Gordo, Proctor et al. 45564 (US). ST. CROIX: 

Orange Grove, A.E. Ricksecker 308 (US). 

28. Cyperus odoratus L., Sp. Pl. 46. 1753; 

Torulinium odoratum (L.) Hooper, Kew Bull. 

26: 579. 1972. Lectotype: Jamaica. Sloane, 

Voy. Jamaica 1: t. 74, f. 1, 1707, designated 

by Dandy in Exell (ed.), Cat. Vasc. Pl. S. 

Tomé : 360. 1944. 

Cyperus ferax Rich., Actes Soc. Hist. Nat. Paris 

1: 106. 1792; Torulinium ferax (Rich.) Urb., 

Symb. Antill. 2: 165. 1900. Type: French 

Guiana; Cayenne. Leblond s.n. (holotype: P; 

isotypes: G, P). 

Cyperus michauxianus Schult., Mant. 2: 123. 

1824; Torulinium michauxianum (Schult.) C. 

B. Clarke in Urban, Symb. Antill. 2: 56. 1900. 

Type: United States. “In Carolina”, collector 

unknown. (holotype: probably at M; isotype: 

possibly at BM). 

Cyperus fastuosus Desv. ex Ham., Prodr. Pl. Ind. 

Occid. 12. 1825. Type: Puerto Rico. Collector 

unknown (holotype: probably at P-Desv.). 

Cyperus poeoides Desv. ex Ham., Prodr. Pl. Ind. 

Occid. 12. 1825. Type: Puerto Rico. Collector 

unknown (holotype: probably at P-Desv.). 

Torulinium confertum Desv. ex Ham., Prodr. Pl. 

Ind. Occid. 15. 1825. Type: French Guiana; 

Cayenne. Collector unknown (holotype: 

probably at P-Desv.). 

Diclidium aciculare Schrad. ex Nees in Martius, 

Fl. Bras. 2(1): 55. 1842; Cyperus acicularis 

(Schrad. ex Nees) Steud., Syn. Pl. Glumac. 

2: 45. 1854, non (Linnaeus) Withering, 1796; 

Cyperus ferax var. acicularis (Schrad. ex 

Nees) Kük. in Engler, Pflanzenr. IV. 20 (Heft 

101): 619. 1936. Type: Brazil; Bahia. Martius 

s.n. (holotype: M). 

Cyperus cubanus Liebm., Mexic. Halvgr. 34. 

1850. Type: Cuba; near Havana. Liebmann 

14355 (holotype: C). 

Cyperus sanctae-crucis Liebm., Mexic. Halvgr. 

35. 1850. Type: St. Croix, U.S. Virgin Islands. 

Oersted 14374 (holotype: C). 

Cyperus parvispiculatus Boeck., Beitr. Cyper. 1: 

6. 1888. Type: St. Thomas, U.S. Virgin 

Islands. Eggers s.n. (holotype: B, destroyed). 

Cyperus flexuosus sensu Grisebach, Fl. Brit. W. I. 

566. 1864, pro parte, non Vahl, 1805. 

Coarse annual, 25-130 cm tall. Culms solitary 

or sometimes tufted in reduced forms, trigonous 

proximally, obtusely triquetrous distally, smooth 

on angles, 1.8-7 (-8.5) mm wide, sheathing bases 

stout, 0.5-2.7 cm wide. Leaves 4-9, basal and lower 

cauline; sheaths loose, the basal ones inflated, 

obscurely cross-veined, green to purple-tinged or 

sometimes brown-black proximally, the veins 

often pale; ligule absent; blades flattish to plicate, 

up to 1 m long, 5-17 mm wide, antrorsely scabrous 

on margins and abaxial midvein, acuminate to 

blunt or rounded tip. Inflorescence a simple or 

compound umbel-like corymb with ascending 

rays, 6-32 (-40) cm in diam; involucral bracts 6- 

15, leaf-like, the lowermost 1-6 dm long; primary 

rays 5-12, to 22 cm long; secondary rays up to 6 

cm long; spikes 1-7 at ray tips, oblong-cylindrical, 

2-7 × 1.4-5 cm, with 20-60 spikelets; spikelets 

linear-lanceolate, subterete, 6-32 × 0.8-1.2 mm, 

ascending near apex of spike, at right angles to 

the rachis below, with the internodes of spike 

rachis between spikelets 1-5 mm wide, the 

spikelets never radiate, rarely reflexed, with 4-24 

florets, disarticulating at the base and at rachilla 

nodes when mature; rachilla with broad, scarious 

wings, these becoming thick and corky at maturity, 

yellowish to brown, enveloping the achene, the 

rachilla node and floret falling together; scales 

ovate to elliptic, somewhat dorsally compressed, 

dorsally rounded or broadly so, 2.2-3.5 (-4) × 

(1.3-)1.5-2 mm, 1-2 nerved on each side bordering 

carina above the narrowly scarious margins, light 

yellowish brown, carina 5- to 7-nerved, green, 

extending as a short prickly-tipped mucro below 

the obtuse apex. Stamens 3, the anthers 0.5-0.9 

mm long, bluntly apiculate; style 3-branched. 

Achene trigonous with convex to nearly plane


faces, oblong-obovate to narrowly so, 

dorsiventrally compressed or only slightly so, 

(1.6-)1.8-2 × 0.5-0.7 mm, slightly falcate at 

maturity, acutely narrowed to short beak, cuneate 

at base, estipitate, minutely puncticulate, lightbrown 

to grayish brown. 




Islands: Wet or damp sandy or gravelly soils in 

marshy and swampy areas, agricultural fields, 

pastures, roadside ditches, coastal swales; edges 

of pools and mangrove swamps; pond margins, 

alluvium along rivers and estuaries, forest trails, 

and waste areas. Adjuntas, Aguas Buenas, 

Aibonito, Añasco, Arecibo, Arroyo, Bayamón, 

Cabo Rojo, Caguas, Camuy, Carolina, Cataño, 

Cayey, Cayo Ramos, Culebra, Dorado, Fajardo, 

Florida, Guánica, Guayama, Guayanilla, 

Humacao, Isabela, Juana Díaz, Juncos, Lajas, 

Lares, Las Piedras, Loíza, Luquillo, Manatí, 

Maricao, Maunabo, Mayagüez, Naguabo, Patillas, 

Peñuelas, Ponce, Río Grande, Río Piedras, Sabana 

Grande, San Germán, San Juan, San Lorenzo, San 

Sebastián, Santa Isabel, Toa Baja, Utuado, Vega 

Alta, Vega Baja, Vieques, Yabucoa, and Yauco; 

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

Gorda. 


Adjuntas: Alto de la Bandera, near Adjuntas, N.L. 

Britton & Shafer 2140 (US). Añasco, Stevens 3214 

(NY). Arecibo; Río Abajo State Forest, Acevedo- 

Rdgz. 10683 (UPRRP, US). Arroyo: Playa 

Guilarte, González-Más 1318 (NY). Bayamón: 

Heller & Heller 410 (NY, US). Cabo Rojo: 

Boqueron, Rd. 4, km 2.5, González-Más 940 (NY, 

US). Caguas: Jct. Rts. 52 & 156, Axelrod et al. 

981 (NY). Carolina: Along Rt. 3 at the Río Grande 

de Loíza, Taylor & Druitt 8008 (NY). Cataño: Palo 

Seco, González-Más 2108 (NY, US). Cayo Ramos: 

South side, Liogier et al. 34643 (NY). Culebra, 

Britton & Wheeler 58 (NY). Dorado: Rd. 165, km 

3.9, González-Más 250 (NY, US). Fajardo: El 

Convento, Liogier 32289 (NY). Guánica: near 

Ochoa Factory, González-Más 956 (NY, US). 


(NY, US). Guayanilla: Rd. 336, Bay, González- 

Más 811 (NY, US). Humacao: University of Puerto 

Rico campus, Acevedo-Rdgz. et al. 5425 (US). 

Juncos: Sintenis 2502 (US). Lajas: Laguna 

Cartagena, Liogier et al. 30553 (NY). Luquillo: 

El Verde Field Station, Taylor 10157 (MO, NY). 

Mayagüez: Sintenis 166 (US). Naguabo: Sierra de 

Luquillo, Caribbean National Forest, Proctor & 

Thomas 43205 (US). Patillas: Road 3, km 129.4, 

González-Más 1326 (NY). Ponce: Cotto Laurel, 

Rd. 14, km 122.2, González-Más 782 (NY, US). 

Sabana Grande: Sargent 209 (US). San Germán: 

Sargent 93 (US). San Juan: Río Piedras, Stevenson 

51 (US). Santa Isabel: Rd 1, km 110.5, González- 

Más 759 (NY). Utuado: Road from Utuado to 

Lares, Underwood & Griggs 69 (NY, US). Vega 

Alta: Rd. 167, km 3, González-Más 1124 (NY, US). 

Vega Baja: Bo. Río Nuevo, Rd 686, km 4, 

González-Más 1107 (NY, US). Vieques Island: 

Isabel Segunda to Sierra Encantada, Shafer 2522 

(NY, US). ST. CROIX: Constitution Hill, A.E. 


29. Cyperus papyrus L., Sp. Pl. 47. 1753. 

Lectotype: “θ Papyrus 15”, Herb. Linn. 

(UPS), designated by D. A. Simpson in 

Cafferty & C. E. Jarvis (ed.), Taxon 53: 179. 

2004. 

Fig. 65. F 

Robust perennial, 100-500 cm tall; rhizome 

short, decumbent, coarse, woody, 1-3 cm thick. 

Culms loosely tufted along rhizome, erect, firm, 

obtusely triquetrous, smooth, 5-16 mm wide 

midculm, to 20 mm wide at base. Leaves reduced 

to bladeless sheaths or sheaths of sterile shoots 

sometimes bearing short blades; proximal sheaths 

coriaceous, distal ones herbaceous distally, finely 

and closely veined, scabridulous to smooth on 

veins abaxially, obliquely truncate at orifice, 

brown; ligule absent. Inflorescence an 

umbelliform, compound corymb with numerous, 

ascending to spreading rays, 15-50 × 10-40 cm; 

involucral bracts short, 7-14, like the sheaths, 

horizontally spreading, much shorter than corymb 

rays, 2-15 cm long; rays numerous, subequal in 

length, 8-50 cm long; secondary rays bearing 

spikes 3-5, to 8 cm long; spikes cylindrical, 10-35 

× 7-18 cm, subloosely to subdensely bearing 5-40 

spicately arranged spikelets; spikelets linear, 

subcompressed, wavy-margined, 4-10 (-12) × 0.7- 

1 mm, acuminate, cuneate at base, with 5-20 

florets; rachilla hyaline-winged; scales elliptic to 

ovate-elliptic, boat-shaped, dorsally obtuse, 1.7- 

2.3 × 0.7-1.4 mm, submembranous, brownish and 

closely 2- to 3-nerved, often obscurely so along 

either side of carina, stramineous to golden brown 

on sides with scarious margins, carina 3-nerved,

282 


green or light brownish, shortly prolonged beyond 

the obtuse apex as a mucro. Stamens 3, the anthers 

0.7-1.7 mm long, with a rounded or linear, papillate 

apiculum; style 3-branched. Achene trigonous, 

plane on abaxial faces, concave and slightly 

curvate on adaxial face, oblong or subobovoidoblong, 

0.8-1 × 0.4-0.5 mm, obtuse at apex, not 

apiculate, obtuse at base, estipitate, shiny, smooth, 

greenish, maturing grayish brown. 

General distribution: Native to eastern 

tropical Africa and Madagascar. Widely cultivated 

and planted as an ornamental elsewhere. 

Introduced and naturalized or escaped from 

cultivation in the New World, occurring in the 

United States (Florida, Louisiana, and California) 

and the West Indies. 


wet areas of standing water along river banks, 

creeks, swales, and roadside ditches; Ciales, Lares, 

and Río Grande. 


Ciales: Bo. Cordillera, finca of José Colón, ca. 0.6 

km N of Rd. 146 at km 21.6., Proctor & Quevedo 

48890 (US). Lares: Bo. Callejones, Rt. 453, just 

before Rt. 454, Axelrod 12640 (UPRRP). 

30. Cyperus planifolius Rich., Actes Soc. Hist. 

Nat. Paris 1: 106. 1792; Mariscus planifolius 

(Rich.) Urb., Symb. Antill. 2: 165. 1900. Type: 

St. Croix, U.S. Virgin Islands. Richard s.n. 


Cyperus purpurascens Vahl, Enum. Pl. 2: 359. 

1805; Mariscus purpurascens (Vahl) C. B. 

Clarke, Bull. Misc. Inform. Kew 1893: 283. 

1893. Type: St. Croix; U.S. Virgin Islands. 

West s.n. (holotype: C-Vahl). 

Cyperus brunneus sensu Grisebach, Fl. Brit. W. I. 

565. 1864, in part, non Swartz, 1797. 

Robust, rhizomatous perennial, (20-) 40-100 

cm tall; rhizome short and thick. Culms solitary, 

3-angled, (1.5-) 2-4 (-5) mm wide at base, smooth, 

the sheathing bases 1-2.5 cm wide. Leaves 8-20, 

primarily basal, crowded with short nodes, 

subdistichous; sheaths short, distinctly fineveined, 

reddish to purple-brown, the inner band 

finely veined; ligule absent; blades stiff, flattish, 

50-90 cm × (4-) 6-12 (-15) mm, glaucous, 

antrorsely scabrous on margins and midvein 

beneath, cross-veined, sometimes obscurely so, 

frequently pale beneath, the surface reddish brown 

lineolate, long-acuminate to triquetrous apex. 

Inflorescence an open, compound umbel-like 

corymb or rarely congested in 1 or 2 head-like 

clusters, (4-) 8-14 (-20) cm diam.; involucral bracts 

6-9, leaf-like, the lowest 18-50 cm × 4-10 mm, 

the upper linear, subulate; rays (6-) 8-9 (-12), 

unequal, obscurely 3-angled to slightly 

compressed, to 13 cm long, 0.6-1.2 mm wide, 

bearing clusters of spikes at apex; spikes broadly 

ovate to broadly oblong-ovate, 1.3-2.5 × 1.5-2.8 

cm, with 6-30 spikelets; spikelets narrowly linear, 

6-15 (-20) × 1.4-1.8 (-2) mm, spreading, falling 

entire, the lower ones divergent to reflexed at 

maturity, with 7-29 florets; rachilla broadly 

hyaline-winged; scales ovate-lanceolate, acutely 

keeled, 2.4-3.2 (-3.5) × 1.4-2.0 mm, closely 4- to 

5-nerved on each side along carina, membranous, 

pale brown to reddish or reddish brown, with 

scarious margins, the narrow, 3-nerved carina 

ending in a short mucro below the acute apex. 

Stamens 3, the anthers 1-1.2 mm long, with a 

minute, triangular, black appendage at apex; styles 

3-branched to below the middle. Achene trigonous 

with plane to slightly convex sides, curvate on 

adaxial side facing rachilla, ellipsoid-obovoid to 

oblong-obovoid, 1.3-1.7 × 0.6-0.9 mm, distinctly 

subulate-apiculate at obtuse apex, cuneate at base, 

minutely puncticulate, blackish at maturity. 


States (Florida, Georgia, and Alabama), West 

Indies, eastern coastal Mexico and Central 

America, and northeastern Brazil. 


Islands: Coastal habitats, in sand or sandy soil of 

beaches, dry scrub forest, rocky thickets, coastal 

rocks, sea-cliffs, edge of mangrove swamps, and 

coastal swales. Aguadilla, Cabo Rojo, Cayo Icacos, 

Culebra, Desecheo, Fajardo, Maunabo, Mona 

Island, Monito, Río Grande, San Juan, Vieques, 

and Yabucoa; Anegada, Buck Island, Capella 

Island, Curt Cay, Deadchest Island, French Cap 

Island, Flannigan Island, George Dog Island, Great 

Camonoe Island, Hilo Island, Leduck Island, Little 

Hans Lollick, Louange Cay, Norman Island, St. 

Croix, St. James, St. John, St. Thomas, Salt Island, 

Tortola, Virgin Gorda, Water Island, Watson Rock, 

and West Dog Island. 

Common names: Puerto Rico: Cortadora, 

Cortadera, Lambedora. 

Note: This species is rare on the mainland of 

Puerto Rico and is often confused with Cyperus


brunneus with which it is closely related. The 

inflorescence of C. planifolius is much broader 

and often more open [(4-) 8-14 (-20) cm diam.] 

than the often congested, head-like inflorescence 

of C. brunneus which is 2-8 (-13) cm in diam. The 

nerves on each side of the reddish scales of C. 

planifolius are crowded towards the dorsal carina 

and often less distinct than the brownish ones of 

C. brunneus which are more widely spaced and 

extend down to the scarious-margined edges of 

the scale. The achene of C. planifolius is slightly 

curvate on the adaxial side and is ellipsoid-obovoid 

to oblong-obovoid, 0.6-0.9 mm wide, while that 

of C. brunneus is plane and obovoid to ellipsoidobovoid, 

0.8-1.1 mm wide. 


Culebra Island: N.L. Britton & Wheeler 51 (US). 

Mayagüez: Monito Island Natural Reserve, 

Breckon et al. 5161 (US). Ponce: Bo. Cañas, Punta 

Cuchara-Laguna de las Salinas region, E side of 

El Tuque Recreation Area, Breckon & Lewis 4065 

(US). Vieques Island: Cerro Ventana, Shafer 2879 

(US). Yauco: Tamarindo Beach to Jacinto Point, 

along shoreline below Guánica State Forest, Strong 

et al. 441 (US). ST. JOHN: East End Quarter, east 

of Southside Pond, Acevedo-Rdgz. et al. 1831 (NY, 

US, VINPS). ST. THOMAS: Water Island, N.L. 

Britton et al. 134 (US); Eggers 224 (NA). VIRGIN 

GORDA: Near Valley, Britton & Fishlock 1089 

(US). 

31. Cyperus polystachyos Rottb., Descr. Pl. Rar. 

21. 1772; Pycreus polystachyos (Rottb.) P. 

Beauv., Fl. Oware 2: 48. 1807. Type: India. 

Koenig s.n. (holotype: C-Rottb.; isotype: C- 

Vahl)*. 

Cyperus hahnianus Boeck., Flora 61: 138. 1878; 

Pycreus polystachyos var. hahnianus (Boeck.) 

C. B. Clarke in Urban, Symb. Antill. 2: 18. 

1900. Type: Martinique. Hahn s.n. (holotype 


Pycreus odoratus sensu Urban, Symb. Antill. 4: 

110. 1903, non (Linnaeus) Urban, 1900. 

Cyperus odoratus sensu Britton & P. Wilson, Bot. 

Porto Rico 5: 81. 1923, non Linnaeus, 1753. 

Caespitose perennial or annual, (7-) 18-77 (- 

115) cm tall; rhizome (when present) short, 

thickened; roots aromatic. Culms erect, 

triquetrous, often obtusely so, subtrigonous 

proximally, firm to subhardened, finely ribbed, 

smooth, (0.8-) 1.2-2.3 (-2.8) mm wide, sheathing 


cauline; sheaths short proximally, reddish brown 

proximally, pale brown to greenish distally; ligule 

absent; blades V-shaped proximally, flattenedplicate 

distally, (3-) 9-45 (-67) cm × 1.2-4 mm, 

smooth, semi-glossy and finely cellular-reticulate 

adaxially, with remote, antrorse, swollen-based, 

whitish or crystalline prickles on the margins and 

abaxial midvein, long-acuminate to triquetrous 

apex. Inflorescence a simple to compound, umbellike 

corymb with ascending rays, or contracted into 

a single, head-like, lobed cluster, 1-6 × 1.4-11 cm; 

involucral bracts (3-) 4-7, leaf-like, ascending to 

horizontal, the lowermost one 3-23 (-30) cm long; 

primary rays 3-8, elongating up to 6 cm or 

essentially absent, secondary rays absent; spikes 

small, ovoid to turbinate, 9-16 × 4-8 mm, 

fasciculately or spicately disposed at ray tips, with 

2-6 somewhat loosely disposed spikelets; spikelets 

linear-lanceolate to linear, 6-13 × 1-1.6 mm, the 

scales spreading to 2 mm wide, compressed, shortacuminate 

at apex, short-cuneate at base, with 6- 

28 florets; rachilla hyaline-winged; scales ovateelliptic 

to oblong-elliptic, dorsally acute or 

subacute, (1.2-) 1.4-1.8 (-2) × 0.8-1.5 mm, thinly 

chartaceous, 1- to 2-nerved along carina above the 

broadly scarious margins, stramineous to 

yellowish brown or reddish brown, carina 3nerved, 

green, prolonged beyond the acute to 

obtuse apex as a short mucro. Stamens (1-) 2, the 

anthers 0.4-0.6 mm long, rounded-apiculate; style 

2-branched. Achene biconvex to broadly so with 

convex faces, oblong, 0.8-1 (-1.2) × 0.4-0.6 mm, 

truncate at apex, short-apiculate, asymetrically 

cuneate at base, substipitate, puncticulate to 

essentially smooth and glossy with an underlying 

reticulation, light brown to brown or grayish brown 

at maturity. 




Islands: Wet sandy or gravelly soils along 

riverbanks, borders of lagoons, pond margins, 

marshy areas, pastures, coastal thickets, sand bars, 

sandy barrens, roadside ditches, and disturbed 

areas. Arecibo, Bayamón, Cabo Rojo, Camuy, 

Carolina, Cataño, Ceiba, Ciales, Cidra, Coamo, 

Dorado, Fajardo, Guayama, Humacao, Isabela, 

Lajas, Loíza, Manatí, Mayagüez, Naguabo, 

Patillas, Río Grande, San Juan, San Lorenzo,

284 


Utuado, Vega Alta, Vega Baja, and Yabucoa, St. 

Thomas and Tortola. 

Notes: Plants in the flora area are referable to 

C. polystachyos var. polystachyos, with a more 

contracted inflorescence. *An illustration 

(Plukenet, Phytographia, Amaltheum botanicum, 

t. 416, f. 6. 1705) was designated as lectotype of 

Cyperus polystachyos by Corcoran, Catholic Univ. 

Amer., Biol. Ser. 37: 26. 1941. However, Tucker 

(1994) cites a Koenig collection at C as the 

holotype. Although we have not seen the material 

at C, the Rottbøll herbarium is there and he was 

director of the Copenhagen Botanical Garden from 

1770-1797. If what appears to be original material 

has in fact been rediscovered, then it supersedes 

Corcorans’ lectotypification (I.C.B.N. Art. 9.17). 


Arecibo: González-Más 1828 (US). Bayamón: 

Finca Santa Ana, Hioram 347 (US). Cataño: Rd. 

165, km 5, González-Más 1150 (US). Dorado: 

Cerro Gordo Beach, Liogier 10240 (US); San 

Antonio, González-Más 349 (US). Fajardo: 

Sintenis 1314 (US). Isabela: Bo. Coto, W bank, 

estuary of Río Guajataca near its mouth, Proctor 

et al. 42357 (US). Lajas: Laguna Cartagena, 

González-Más 945 (US). Manatí: Sintenis 6789 

(US). Mayagüez: Guanajibo, Rd. 102, km 5.3, 

González-Más 826 (US). Naguabo: Daguao River, 

Rd. 3, González-Más 233 (US). Río Grande: El 

Verde Forest entrance, González-Más 137 (US). 

San Juan: Stone Quarry, 2 mi. E of Santurce, Heller 

& Heller 1308 (US). Vega Baja: Bo. Cibuco, just 

SW of Playa Cerro Gordo, Proctor et al. 45587 

(US). 

32. Cyperus pulguerensis M.T. Strong sp. nov. 

Type: Puerto Rico. Manatí: Bo. Tierras 

Nuevas Saliente, Road 686, ca. 100 m E of 

intersection at El Pulguero, 24 Jun 1987, 

Proctor & Thomas 43804 (holotype: US; 

isotype: SJ). 

Fig. 49. A-K 

A Cyperus confertus Sw. spiculis squamis 

mucronatis rectis differt. 


knotty, bases cormose. Culm erect, triquetrous, 

ribbed and channeled, 1.3-3 mm wide, smooth. 

Leaves 4-7, primarily basal and lower cauline, 

spreading; ligule absent; sheaths short, the inner 

band membranous, minutely red-dotted, concave 

at orifice; blades linear, 6-30 cm × 2-3 mm, folded 

proximately, subflattened distally, finely veined 

and dull adaxially, finely cellular-reticulate and 

subglossy adaxially, attenuate to triquetrous tip. 

Inflorescence a simple umbel-like corymb with 5- 

6 ascending rays; involucral bracts 5-7, the 

lowermost elongate, leaf-like; rays 2-8 cm long; 

spikes broadly subglobose-oblong, dense, with 15- 

40 spikelets; spikelets spreading-divaricate or the 

lowermost reflexed, slender, 6-14 × 1.5-1.8 mm, 

5- to 14-flowered; rachilla hyaline winged; scales 

loosely imbricate, ovate-elliptic, 2.5-3 × 1.7-2 mm, 

uniformly light brown or light yellowish brown, 

carina light green, 3-nerved, prolonged beyond the 

obtuse apex as a thickened, elongate, straight, 

minutely prickle-tipped mucro, the sides coarsely 

3- to 4-nerved. Stamens 3, the anthers 0.4-0.6 mm 

long; style 3-branched. Achene narrowly ellipsoid, 

triquetrous, 1.3-1.6 × 0.6-0.7 mm, short-cuneate 

at base, acute to an apiculate apex, puncticulate, 

dark reddish at maturity. 

General distribution: Endemic to Puerto Rico. 

Distribution in Puerto Rico: On white sand 

at the western end of Laguna Tortuguero. Only 

known from the type collection. 

Note: Cyperus pulguerensis is similar in habit 

to C. confertus but differs in the spikelet scales 

which have the carina prolonged at apex as a 

straight thickened murcro vs. a distinctly to slightly 

excurved cusp in C. confertus. 

33. Cyperus rotundus L., Sp. Pl. 45. 1753. 

Lectotype: India. Herb. Hermann 1: 3, No. 

36 (BM-2), designated by G. C. Tucker, Syst. 

Bot. Monogr. 43: 100. 1994. 

Cyperus pseudo-variegatus Boeck., Beitr. Cyper. 

2: 37. 1880. Type: Martinique. Duss 449 


Stoloniferous perennial, (6-) 10-80 cm tall; 

stolons elongate, often bearing tubers. Culms 

solitary, tuberous at base, triquetrous, 

subcompressed, 0.7-3 mm wide, finely ribbed, 

smooth, sheathing base of culm 3-10 mm wide. 

Leaves 7-17 (-24), clustered at lower cauline and 

base; sheaths short, cross-veined, sometimes 

indistinctly so, pale brown; ligule present on lower 

sheaths, short and often faint, absent on upper 

sheaths; blades shorter than the culms, folded or 

V-shaped proximally, flattish or subplicate distally,


Fig. 49. A-K. Cyperus pulguerensis. A. Habit. B. Inflorescence. C. Inflorescence spike. D. Spikelet. E-F. Sterile bracts subtending 

base of spikelet. G. Lowermost spikelet scale (spread open). H. Upper spikelet scale (spread open, dorsal view). I. Upper spikelet 

scale (spread open, ventral view, showing winged rachilla and developing achene). J. Spikelet scale (side view) clasping developing 

achene. K. Achene. (A-K, from the type, Proctor & Thomas 43804).

286 


5-42 cm × 2-7 (-9.5) mm, antrorsely scabrous 

along margins and midvein beneath distally, 

acuminate to triquetrous apex. Inflorescence a 

simple to compound, narrowly hemispherical, 

umbel-like corymb, (2-) 3-14 (-17) × (1.5-) 2.5- 

10 (-21) cm; involucral bracts 2-5, leaf-like, the 

lowest 1-9 (-14) cm long; rays (1-) 2-7 or absent, 

unequal, 4-10 cm long; spikes 1 (-3) at ray tips, 

broadly ovoid, 1.5-4 × 1.5-7 cm, with 2-12 

spikelets; spikelets linear-lanceolate, 

subcompressed, 10-45 × 1.5-2 mm, acute at apex, 

cuneate at base, with 9-51 florets; rachilla 

flexuous, hyaline-winged; scales deciduous, ovate 

to broadly ovate or ovate-elliptic, 2.8-3.5 × 2-2.4 

mm, subacutely to obtusely keeled, 2- to 3-nerved 

on each side above the scarious margins, 

submembranous, dark reddish brown to purplish 

brown or blackish, with 1-2 indistinct lateral 

nerves evident only at proximal base of scale, the 

sides smooth for the most part, carina 3- or 5nerved, 

greenish, ending in a short, slightly 

excurved mucro at the cleft of the obtuse 

emarginate apex. Stamens 3, the anthers 1.5-2.2 

mm long, apiculate; styles elongate, 3-branched 

to below the middle, the unbranched portion 

subflattened. Achene trigonous with the adaxial 

face concave and the abaxial faces plane to slightly 

convex, ellipsoid to oblong-ellipsoid, 1.4-1.8 × 

0.7-0.9 mm, obtuse to sub-rounded at apex, shortapiculate, 

short-cuneate at base, estipitate, 

minutely puncticulate, brown to blackish. 




Islands: Sandy soils of coastal areas and cultivated 

grounds, in riverbanks, roadside ditches, sugar 

cane fields, pastures, lawns, disturbed areas, and 

waste places. Añasco, Bayamón, Cabo Rojo, 

Camuy, Carolina, Cataño, Ceiba, Coamo, Culebra, 

Desecheo, Dorado, Fajardo, Guánica, Guayama, 

Guayanilla, Humacao, Las Piedras, Loíza, 

Mayagüez, Mona Island, Naguabo, Peñuelas, 

Ponce, Salinas, San Germán, San Juan, Santa 

Isabel, Vieques, Yabucoa, and Yauco; St. Croix, 

St. John, St. Thomas, Tortola, and Virgin Gorda. 

Common names: Puerto Rico: Coquí, 

Coquillo, Yerba Coqui. 

Note: See discussion under C. esculentus for 

differences between it and C. rotundus. 



(US). Cabo Rojo: Boqueron, Rd. 4, km 2.4, 

González-Más 937 (US). Camuy: Underwood & 

Griggs 208 (US). Desecheo Island, south-facing 

slope just N of Puerto de los Botes, Breckon 4947 

(US). Dorado: Rd. 165, km 3.9, to Toa Baja, 

González-Más 258 (US). Guánica: Sargent 42 

(US). Guayama: Pto. Jobos, Rd. 707, González- 

Más 689 (US). Mayagüez: Sintenis 12 (US). Mona 

Island: Sardinera, Otero & Chardón 837 (US). 

Naguabo: Daguao River, Rd. 3, González-Más 211 

(US). Ponce: Rd. 14, banks of Inabón River, 

González-Más 776 (US). San Juan: Río Piedras, 

Stevenson 46 (US). Santa Isabel: Rd. 1, km 110.5, 

González-Más 758 (US). Vieques Island: 

Mosquito to Playa Grande, Shafer 2856 (US). 

Yabucoa: Sintenis 5089 (US). Yauco: Rd. 2, near 

Río Loco, González-Más 853 (US). ST. CROIX: 

Bassin Yard, A.E. Ricksecker 159 (US). ST. JOHN: 

Coral Bay, Raunkiaer s.n. (US). ST. THOMAS: 

Eggers s.n. (NA, US). TORTOLA: Fishlock 134 

(US). 

34. Cyperus sphacelatus Rottb., Descr. Pl. Rar. 

21. 1772 . Lectotype: Surinam. Rolander 32 

(C-Rottb., photo at F, US, IDC microfiche: 

type herbarium nr. 67 II, 4-5), designated by 

McLaughlin, Catholic Univ. Amer. Sci. Stud. 

5: 71. 1944. 

Cyperus balbisii Kunth, Enum. Pl. 2: 63. 1837. 

Type: Dominican Republic. Bertero s.n. 

(holotype: probably at TO). 

Caespitose annual, (10-) 13-67 (-90) cm tall. 

Culms slender, triquetrous, sometimes trigonous 

proximally, finely ribbed, smooth, 0.6-2.4 mm 

wide, sheathing bases (1.5-) 2-6 mm wide. Leaves 

2-6, basal and lower cauline; sheaths elongate, 

subloose, yellow-green or brownish; ligule absent; 

blades plicate, 3-42 cm × (1.3-) 1.5-4 (-5) mm, 

smooth, antrorsely scabrous on the margins and 

abaxial midvein, at least distally, narrowly 

acuminate to triquetrous apex. Inflorescence a 

simple or compound, open to contracted, umbellike 

corymb with ascending rays, (2-) 3-13 (-15) 

× (2-) 3-17 (-26) cm; involucral bracts (4-) 5-7, 

ascending to patent, leaf-like, the lowermost one 

to 30 cm long; primary rays (2-) 3-7 or essentially 

absent, unequal, elongating to 15 cm, the 

secondary lateral rays short or wanting; spikes 

solitary or 2-3 at ray tips (lateral spikes smaller 

than central one), broadly ovoid, 1-3.5 (-4.5) × 1-


5 (-6.5) cm, with 4-16 somewhat loosely disposed, 

spicately arranged spikelets; spikelets linearlanceolate, 

(5-) 6-23 (excluding denuded rachilla) 

× 1.2-1.8 (-2) mm, subflattened, acute to acuminate 

at apex, cuneate at base, with 6-22 (-28) florets, 

the scales spreading with developing achenes; 

rachilla unwinged; scales ovate-elliptic, dorsally 

acute to obtuse, 2.2-3 × 1.6-2.2 mm, thinly 

herbaceous, 2-3 nerved on each side above the 

broadly scarious margins, stramineous to light 

brown or yellowish brown, often reddish or brown 

lineolate, usually with a brown-black or purpleblack 

stain on the scarious margins, carina broad, 

finely 5- to 7-nerved, greenish, prolonged at the 

subobtuse to subacute emarginate apex as a short 


bluntly apiculate; style 3-branched. Achene 

trigonous with adaxial face concave and abaxial 

faces plane to slightly concave, narrowly ellipsoid 

or ellipsoid-obovoid, 1.2-1.5 × 0.6-0.9 mm, obtuse 

at apex, short-cuneate at base, short-stipitate with 

a small basal torus, obscurely puncticulate to 

essentially smooth, reddish brown to dark brown. 



Islands: In open, often wet areas on sandy and 

gravelly soils of washes along rivers and streams, 

swales, grassy slopes and hillsides, rocky ridges, 

marshy areas, clearings, pastures, ditches, 

roadsides, lawns, and waste areas. Adjuntas, Aguas 

Buenas, Aibonito, Añasco, Arecibo, Bayamón, 

Cabo Rojo, Caguas, Carolina, Cataño, Cayey, 

Cayo Ramos, Ceiba, Ciales, Corozal, Dorado, 

Fajardo, Guaynabo, Gurabo, Humacao, Isabela, 

Las Piedras, Loíza, Luquillo, Manatí, Maunabo, 

Maricao, Mayagüez, Naguabo, Orocovis, Patillas, 

Peñuelas, Ponce, Río Grande, Salinas, San Juan, 

San Lorenzo, San Sebastián, Trujillo Alto, Utuado, 

Vega Alta, Vega Baja, Vieques, and Yabucoa; St. 

Croix, St. Thomas, and Tortola. 



10639 (UPRRP, US). Bayamón: Underwood & 

Griggs 876 (US). Caguas: Bo. Cañabón, along W 

side of Río Cagüitas, just S of Rd. 156, Proctor 

46149 (US). Carolina: Boca de Cangrejos, 

González-Más 1026 (US). Corozal: At Sub-Station 

Corozal, Woodbury et al. s.n. (US). Dorado: to 

Cerro Gordo, Rd. 693, km 11.3, González-Más 284 

(US). Humacao: Mariana, González-Más 1349 

(US). Loíza: Bo. Medianía Alta, Miñi Miñi, 

Proctor & Colón 49565 (US). Manatí: Laguna 

Tortuguero, Acevedo-Rdgz. & Cedeño 9466 (FTG, 

NY, UPR, UPRRP). Maunabo: La Pica, Rd. 3, km 

105, González-Más 1338 (US). Maricao: Rd. 120, 

González-Más 416 (US). Mayagüez, Sintenis 90 

(US). Naguabo: Rd. 191, km 21.1, towards Sierra 

de Luquillo, González-Más 1399 (US). Patillas: 

Rd. 2, km 129.4, González-Más 1329 (US). Ponce: 

On the Adjuntas road 10 mi. from Ponce, Heller 

6239 (US). San Juan: Río Piedras, Sabana Llana, 

Highland Park, González-Más 1216 (US). Vega 

Baja: Tortuguero, Rd. 681, km 1, González-Más 

1049 (US). Vega Alta: Bo. Sabana, silica-sand area 

NE of Regadera, Proctor & Concepción 41847 

(US). Vieques Island: 1 km N of Laguna Playa, 

Fosberg 57571 (US). ST. THOMAS: Crown, E.G. 

Britton & Marble 1344 (US). TORTOLA: Road Town 

to High Bush, N.L. Britton & Shafer 758 (US). 

35. Cyperus squarrosus L., Cent. Pl. 2: 6. 1756; 

Mariscus squarrosus (L.) C. B. Clarke in 

Hooker f., Fl. Brit. India 6: 623. 1893. 

Lectotype: India. Koenig s.n. (LINN-70.8 B 

right hand specimen; isolectotype: S-LINN 

G-6816), designated by J. Kern, Blumea 10: 

642. 1960. 

Scirpus intricatus L., Mant. Pl. 182. 1771; Cyperus 

aristatus Rottb., Descr. Pl. Icon. Rar. 22. 1772, 

nom. illeg. Lectotype: South Africa. (LINN- 

71.50), designated by C. B. Clarke, J. Linn. 

Soc., Bot. 30: 313. 1894. 

Cyperus inflexus Muhl., Descr. Gram. 16. 1817. 

Type: United States; Pennsylvania. 

Muehlenberg Herb. no. 27 (holotype: PH; 

isotype: B-Willd. 1402). 

Cyperus cuspidatus sensu Britton & P. Wilson, 

1923, non Kunth, 1816. 

Caespitose annual, 1-17 (-20) cm tall; roots 

fine. Culms tufted, triquetrous, wing-angled, soft, 

finely ribbed, smooth, 0.4-1.3 (1.6) mm wide. 

Leaves 1-3, basal; sheaths short, loose and inflated, 

thin, pale green, light brown, or purple-black, 

reddish to brownish lineolate, semi-glossy, 

subcellular-translucent; ligule absent; blades 

linear, folded proximally, subplicate distally, (1-) 

2-10 (-15) cm × 0.8-2.7 (-3) mm (unfolded), 

adaxial surface distinctly cellular-reticulate, 

smooth on margins and abaxial midvein, longacuminate 


simple umbel-like corymb with few ascending

288 


rays, 0.5-4 (-6) × 0.5-3 (-4.5) mm; involucral bracts 

(1-) 2-4, leaf-like, the lowest one strongly 

ascending, to 11 cm long, the others divergent; 

rays 1-3 (-6) or absent, unequal, 4-40 mm long; 

spikes 1 (-2) at ray tips, ovoid or oblong, 5-20 

mm in diam., with (2-) 6-30 (-40) densely to 

subloosely disposed spikelets; spikelets ovatelanceolate 

to oblong-lanceolate, subcompressed, 

(2.5-) 3-11 × 0.8-2 (-2.2) mm (excluding awns), 

obtuse at apex, short-cuneate at base, with (6-) 8- 

26 (-34) florets; rachilla indistinctly winged; scales 

oblong-lanceolate, 1.3-1.8 × 0.6-1 (-1.2) mm 

(excluding awn), dorsally acute, coarsely 4-5 

nerved on each side above the entire margins, 

membranous, subtranslucent, reddish to blackish 

lineolate, stramineous, yellow brown, or brownish 

red, lustrous, carina 1-nerved, greenish, the 

subtending lateral nerves converging at the obtuse 

apex forming a 0.5-1.3 mm long excurved awn. 

Stamen 1, the anthers 0.2-0.4 mm long, bluntly 

apiculate; styles 3-branched. Achene trigonous 

with slightly convex to flat faces, lanceoloidobovoid 

or obovoid, 0.6-0.9 (-1.1) × (0.2-) 0.3- 

0.4 (-0.5) mm, broadly rounded to truncate or 3lobed 

at apex, apiculate, cuneate at base with a 

small torus, lustrous-papillose, light brown to 

blackish. 

General distribution: Cosmopolitan. 

Distribution in the Virgin Islands: Often in 

sandy, open habitats. Anegada and St. Thomas. To 

be expected in Puerto Rico. 

Selected specimens examined: ANEGADA: Near 

The Settlement, N.L. Britton & Fishlock 1006 

(US); Along track ca. 1.5 km N of The Settlement, 

Proctor & Haneke 45971 (US); Just inland from 

Loblolly Point, Proctor & Haneke 45968 (US). 

ST. THOMAS: Eggers s.n. (US). 

36. Cyperus subtenuis (Kük.) M.T. Strong, comb. 

& stat. nov.; Cyperus nanus var. subtenuis 

Kük. in Engler, Pflanzenr. IV. 20 (Heft 101): 

536. 1936. Type: Navassa Island. Ekman 

10806 (lectotype: US!; isolectotype: NY), 

here designated. 

Cyperus tenuis sensu Grisebach, Fl. Brit. West 

Indies Is. 563. 1864, non Swartz, 1788. 


short. Culms erect to ascending, slender, 

triquetrous, flattened-trigonous distally, firm, 

smooth to antrorsely scabrous on margins (at least 

distally), glabrous. Leaves 3-5 per culm, basal and 

lower cauline, ascending; sheaths short, 

herbaceous, distinctly veined, smooth to remotely 

scabrous, light brown to reddish brown proximally, 

glabrous, the inner band concave to truncate at 

orifice; ligule absent; blades linear, subflattened 

to folded, 4-25 cm × 0.7-2 (-2.5) mm, soft, 

herbaceous, green, abaxially distinctly veined and 

often scabrous, adaxially smooth, glabrous, finely 

cellular-reticulate, margins and abaxial midvein 

finely mixed antrorsely-retrorsely scabrous, 

attenuate to triquetrous tip. Inflorescence a single 

small spike at the summit of the culm; involucral 

bracts 2-4, divergent to reflexed, exceeding length 

of inflorescence; spikes obovoid to oblongobovoid, 

5-13 mm diam., the spikelets reflexed 

except for the terminal one which is often erect; 

spikelets 7-33 (-45), linear-lanceolate, 3.5-7 × 0.7- 

1.2 mm, rhombic, acuminate, cuneate at base, with 

4-8 florets, the scales slightly spreading; fertile 

scales ovate-elliptic, 1.5-2 × 1.2-1.6 mm, dorsally 

acute, thinly herbaceous, smooth, glabrous, 

yellow-brown, margins broadly scarious, carina 

3-nerved, greenish, prolonged beyond the obtuse 

to acute apex as a short, slightly excurved mucro, 

lateral nerves distinct, coarse, 4-5 on each side; 

rachilla broadly hyaline-winged. Stamens 3, the 

anthers 0.3-0.5 mm long, rounded-apiculate; style 

3-branched. Achene trigonous, ovoid-ellipsoid, 

1.1-1.4 × 0.5-0.7 mm, with plane to slightly 

concave sides, curvate adaxially, short-cuneate, 

short-apiculate, obtuse at base, puncticulate, 

brown to brown-black. 


Jamaica, Puerto Rico, and the Virgin Islands. 


Islands: Known only from the southwestern region 

of Puerto Rico along coastal areas on rocky slopes, 

ravines, and sea shores. Lajas and Yauco; St. Croix. 

Note: Cyperus subtenuis was described as a 

variety of Cyperus nanus Willd. by Kükenthal 

(1936) and is closely related to it. However, it 

consistently differs from the latter in its coarser 

habit, spikelet scales 1.5-2 × 1.2-1.6 mm and 4- to 

5-nerved (vs. 1.2-1.6 × 0.8-1 mm and 3- to 4nerved), 

and achenes which are ovoid-ellipsoid, 

1.1-1.4 × 0.5-0.7 mm with curvate adaxial side 

(vs. oblong-ellipsoid, 1-1.2 × 0.5-0.6 mm, and 

straight adaxial side) and is therefore treated here 

at the specific level. 


Lajas: Bo. Parguera, McKenzie s.n. (SJ); Proctor 

44138 (SJ). Yauco: Stevens & Hess 3309 (NY).


37. Cyperus surinamensis Rottb., Descr. Pl. Rar. 

20. 1772. Type: Surinam. Rolander s.n. 

(holotype: probably at C-Rottb.). 


tall; rhizome short, becoming thickened and 

knotty. Culms 2-6, caespitose or arising at short 

intervals along the rhizome, trigonous, scabrous 

distally with scattered retrorse and antrorse barbs, 

nearly smooth proximally, (1-)2-4 (-4.5) mm wide, 

sheathing base of culm (3-) 4-7 (-12) mm wide. 

Leaves 3-9, primarily basal, several sometimes 

lower cauline; sheaths short, cross-veined, pale 

green to brownish, the inner band finely-veined; 

ligule absent; blades flattened or folded, (6-) 12- 

45 (-65) cm × 2-5 (-6.5) mm, with antrorsely 

scabrous or scaberulous margins and abaxial 

midvein, acuminate to triquetrous apex. 

Inflorescence a compound, open to subcontracted, 

umbel-like corymb with ascending rays, (2-)5-18 

cm diam.; involucral bracts (3-) 4-10, leaf-like, 

the lowermost ones to 50 cm long, the uppermost 

short, linear-setaceous; rays (3-) 4-17 (-25), very 

unequal, the primary ones to 15 cm long, stiff, 

ascending, the secondary ones 3-6, 1-2 cm long, 

divergent to reflexed at maturity; spikes broadly 

ovate to hemispherical or subrounded, 5-18 (-20) 

mm diam., with 6-50 (-90) densely disposed 

digitately arranged spikelets; spikelets ovate, 

oblong-ovate or oblong-lanceolate, distinctly 

flattened, (3-) 4-8 (-12) × 1.5-2 mm (3-5 mm long 

if denuded rachilla excluded), acute at apex, 

widely obtuse to sub-rounded at base, with (9-) 

11-31 (-45) florets (5-22 florets if denuded rachilla 

excluded); rachilla unwinged; scales oblong-ovate 

to lanceolate, curvate-keeled, dorsally acute, 1.2- 

1.5 × 0.7-0.9 mm, membranous, minutely 

reticulate, pale yellow, pale brown, or reddish 

brown, carina 3-nerved, 2-keeled near base, 

greenish, prolonged beyond the acute apex as a 

short barb-tipped mucro. Stamen 1, the anther 0.7- 

0.8 mm long; style 3-branched to below the 

middle. Achene trigonous with plane to slightly 

convex faces, narrowly ovate-ellipsoid to narrowly 

oblong-ellipsoid, 0.8-0.9 × 0.3 mm, acute at apex, 

apiculate, obtuse at base, short-stipitate, shiny, 

papillate, indistinctly rugulose, brown to reddish 

brown. 


States, West Indies, Mexico, Central America, and 



Islands: Wet or moist areas; in sandy or gravelly 

soils of lake and pond margins, river and stream 

edges, marshy or swampy areas, meadows, sandy 

savannas; roadside ditches, coastal areas, pastures, 

agricultural fields, and waste areas. Primarily at 

lower elevations, along roadsides at higher 

elevations. Aguada, Aibonito, Añasco, Arecibo, 

Arroyo, Bayamón, Cabo Rojo, Caguas, Camuy, 

Carolina, Cataño, Ceiba, Dorado, Fajardo, 

Guayama, Humacao, Juana Díaz, Loíza: Luquillo, 

Manatí, Maunabo, Mayagüez, Morovis, Naguabo, 

Patillas, Río Grande, Río Piedras, San Juan, San 

Lorenzo, Toa Baja, Utuado, Vega Alta, Vega Baja, 

Vieques, and Yabucoa; St. John and St. Thomas. 


Añasco: Hwy. Mayagüez to Moca, km 145.2, 

González-Más 1782 (US). Arroyo: Playa de 

Guilarte, González-Más 1312 (US). Bayamón: 

N.L. Britton 2346 (US). Carolina: Sabana Abajo, 

near Carolina, N.L. Britton & Brown 5710 (US). 

Cataño: Rd. 165, km 5, González-Más 1155 (US). 

Dorado: Johnston 884 (US). Fajardo: Sintenis 

1256 (US). Guayama: Along Rd. 179, NE of 

Guayama, near km 14, Stimson 1739 (US). 

Humacao: Pozal, Playa de Humacao, Eggers 594 

(US). Juana Díaz: Rd. 14, Guayabal, González- 

Más 770 (US). Manatí: Bo. Tierras Nuevas 

Saliente, area 0.7-1.4 km E of Rd. 686 at Baldwin 

School (SE of El Pulguero), Proctor & Díaz 42159 

(US). Mayagüez: Guanajibo, Rd. 102, Puente Río 

Estero, González-Más 821 (US). Naguabo: Sierra 

de Luquillo, Caribbean National Forest, Proctor 

& Thomas 43203 (US). San Juan: Río Piedras Exp. 

Station, Johnston 776 (US). Vega Alta: Bo. Sabana, 

silica-sand area NE of Regadera, Proctor & 

Concepción 41845 (US); Laguna San José, 

Hioram 374 (US).Vega Baja: Laguna Rica, near 

Camp Tortuguero National Guard, off Rt. 687, 

Strong et al. 429 (GMUF). Vieques: Isabel 

Segunda to Sierra Encantada, Shafer 2524 (NY). 

Yabucoa: Rd. 3, km 96.2, González-Más 1343 

(US). ST. JOHN: Herman Farm, Woodbury 457/6911 

(VINPS). ST. THOMAS: Nazareth, Acevedo-Rdgz. 

11304 (FTG). 

38. Cyperus swartzii Boeck., Beitr. Cyper. 1: 10. 

1888; Kyllinga filiformis Sw., Prodr. 20. 1788, 

[non Cyperus filiformis Sw., 1788]; Mariscus 

filiformis Spreng., Syst. Veg. 1: 234. 1824; 

Mariscus swartzii A. Dietr., Sp. Pl. 2: 343.

290 


1833, nom. illeg. Type: Jamaica. Swartz s.n. 

(holotype: S-Sw., R-3118). 

Mariscus gracilis Vahl, Enum. Pl. 2: 373. 1805, 

[non Cyperus gracilis R. Br., 1810]. Type: 

South America. Richard s.n. (holotype: C- 

Vahl). 

Kyllinga granularis Desf. ex Boeck., Linnaea 35: 

432. 1868; Cyperus granularis (Desf. ex 

Boeck.) Britton, Bull. Torrey Bot. Club 43: 

44. 1916. Cyperus swartzii var. granularis 

(Desf. ex Boeck.) Kük., Repert. Spec. Nov. 

Regni Veg. 23: 186. 1926. Type: Hispaniola; 

collector unknown. (holotype: B-Willd. 1443, 

which stems from Desfontaines). 

Cyperus swartzii var. elongatus Kük., Repert. 

Spec. Nov. Regni Veg. 23: 187. 1926. 

Syntypes: Cuba. Ekman 10989 (B, destroyed); 

Puerto Rico. Sintenis 3808 (B, destroyed). 

Caespitose perennial (1-) 5-56 (-70) cm tall; 

rhizomes short, knotty, thickened, or often 

horizontal, 2-3 mm thick. Culms densely to loosely 

tufted, slender, filiform-triquetrous with obtuse to 

rounded angles, finely ribbed, medially channeled, 

smooth or essentially so, 0.5-1.3 (-1.5) mm wide, 

sheathing bases 1-2 mm wide. Leaves 3-6, basal, 

the lowermost bladeless; sheaths light brown or 

reddish brown, reddish or purplish dotted adaxially 

and on membranous inner band; ligule absent; 

blades narrowly linear, V-shaped proximally to 

plicate or subflattened distally, 8-30 cm × 1-3 mm 

(flattened), finely antrorsely scabrous on the 

margins and abaxial midvein at least distally, longattenuate 


contracted to open, umbel-like corymb with short 

ascending rays or often reduced to a single headlike 

cluster of 2-5 (-6) spikes, 4-40 mm in diam; 

involucral bracts 3-4 (-5), leaf-like, the lowermost 

ones erect or strongly ascending, 2-15 (-19) cm 

long, the others ascending to spreading; rays 1-3, 

short, to 35 mm long, or absent; secondary rays 

absent; spikes ovoid to oblong-ovoid, 3-9 × 3-4 

mm, with (6-) 16-60 densely disposed, spicately 

arranged spikelets; spikelets elliptic-obovoid, 

scarcely compressed, (1-) 1.4-2.3 × 0.6-1.3 mm, 

quadrate-rhombic, acute to short-acuminate at 

apex and base, with 1 floret; rachilla broadly 

hyaline-winged; fertile scale ovate, dorsally acute, 

1.4-2 × 1.4-1.8 mm, submembranous, 6-8 nerved 

above the scarious inrolled margins, light brown 

to tan or yellowish or reddish brown, reddish 

lineolate, carina 3-nerved, the medial one coarse, 

greenish brown, prolonged above the obtuse apex 

as a short slightly recurved mucro; terminal scale 

reduced, sterile, dorsally gibbous or pouched 

medially along carina. Stamens 3, the anthers 0.3- 

0.6 mm long, apiculate; style 3-branched. Achene 

trigonous with slightly convex faces, ellipticobovoid, 

1-1.3 × 0.6-0.9 mm, obtuse at apex, 

apiculate, cuneate at base, short-stipitate, 

puncticulate, dark brown or blackish. 


Antilles, and known from a single collection made 

in San Luis Potosí, Mexico. 


Islands: Known only from the southwestern region 

in Puerto Rico in open to shaded moist clearings, 

pastures, pond borders, savanna, and hillsides. 

Guánica and Peñuelas; St. Croix. 

Note: The following collections were cited by 

Kükenthal (1936) for Cyperus swartzii and its 

varieties: Guánica: Sintenis 3808 (B). Peñuelas: 

Sintenis 4742 (B). ST. CROIX: Benzon s.n. (B). 

These historical collections, all at Berlin (B), are 

probably now destroyed. No recent collections 

have been made from Puerto Rico or the Virgin 

Islands. 

39. Cyperus tenuis Sw., Prodr. 20. 1788; Mariscus 

tenuis (Sw.) C. B. Clarke in Urban, Symb. 

Antill. 2: 48. 1900. Type: Jamaica. Swartz s.n. 

(lectotype: S-Sw. R-1386; isolectotypes: B- 

Willd. 1390, C, M, S). designated by G. C. 

Tucker, Syst. Bot. 11: 15. 1986. 

Mariscus flabelliformis Kunth in Humb., Bonpl., 

& Kunth, Nov. Gen. Sp. 1 [quarto ed.]: 215. 

1816. Type: Venezuela; Caracas. Humboldt & 

Bonpland s.n. (holotype: B-Willd. 1433) . 

Cyperus platystachyus Griseb., Fl. Brit. W. I. 567. 

1864. Type: Jamaica. Collector unknown 


Cyperus hartii Boeck., Beitr. Cyper. 1: 9. 1888. 

Type: Jamaica. Hart s.n. (holotype B, 

destroyed). 

Caespitose perennial, 10-60 cm tall; rhizomes 

short, bulbous-thickened, 1-5 mm thick. Culms 

trigonous, often obtusely so, coarsely ribbed and 

channeled, slender but stiff, smooth, 0.7-1.8 (-2.2) 

mm wide, sheathing bases 2-5 mm wide. Leaves 

3-6, basal and lower cauline; sheaths palestramineous, 

the lowermost stained with red or 

purple-brown; ligule absent; blades V-shaped 

proximally, flattened distally, 5-35 cm × (1-) 1.8-


3 (-4.2) mm, smooth, antrorsely scabrous on the 

margins and abaxial midvein, narrowly acuminate 

at apex. Inflorescence a contracted or sometimes 

open, simple, umbel-like corymb with ascending 

rays, 2-5 (-6.5) × 3-8 (-11) cm; involucral bracts 

5-11, leaf-like, the lowermost to 42 cm long, 

spreading, ascending to horizontal; rays (2-) 5-12 

(-14), elongating to 5 cm; secondary rays absent; 

spikes (2-) 3-11, loosely ovoid to oblong-ovoid, 

(10-) 15-25 (-35) × (8-) 10-25 mm, broadly 

rounded to subtruncate at apex, with (10-) 15-50 

(-70) somewhat loosely disposed, spicately 

arranged spikelets; spikelets linear, 5-15 (-22) × 

0.7-1.1 mm, compressed-quadrate, acuminate at 

apex, cuneate at base, with (2-) 4-11 (-20) florets; 

rachilla broadly hyaline-winged; scales narrowly 

elliptic, oblong-elliptic, or ovate-oblong, dorsally 

obtuse, 2.4-3.4 × (1-) 1.3-1.8 mm, membranous, 

laterally 3-7 nerved on each side above the broadly 

scarious involute margins, greenish white or pale 

stramineous, often reddish brown on sides, carina 

3-nerved, green, prolonged beyond the broadly 

rounded apex as a short mucro. Stamens 3, the 


3-branched. Achene trigonous with concave to 

barely convex faces, linear to narrowly oblong, 

slightly curved adaxially, (1.4-) 1.6-1.8 (-2) × 0.4- 

0.5 mm, subacute at apex, apiculate, cuneate at 

base, short-stipitate, punctate, brown to reddish 

brown. 

General distribution: West Indies, Mexico, 

Central America, South America, and tropical 

Africa. 


Islands: Roadsides and open disturbed areas. 

Dorado, Maricao, Mayagüez, and San Germán; St. 

Croix. 


Dorado: Bo. Higuillar, Rd. 693, 2.1 km S of 

Dorado, Proctor 42459 (US). Maricao: Maricao 

State Forest, Liogier & Liogier 36778 (US); Bo. 

Maricao Afuera, vicinity, Proctor & McKenzie 

43836 (US). 

40. Cyperus unifolius Boeck., Linnaea 36: 374. 

1870. Type: St. Croix, U.S. Virgin Islands. 

Ravn s.n. (holotype: B, destroyed; isotype: 

probably at C). 

Cyperus calcicola Britton, Bull. Torrey Bot. Club 

42: 387. 1915. Type: Puerto Rico; Caja de 

Muertos. N. L. Britton et al. 4976 (holotype: 

NY!; isotypes: UPR!, US!). 

Caespitose perennial, (3-) 6-40 (-45) cm tall; 

rhizome short, nodose; roots aromatic. Culms 

tufted, firm, filiform, subcompressed, obtusely 

triquetrous, finely ribbed, smooth proximally, 

antrorsely scabridulous distally at apex, 0.3-0.8 

mm wide, sheathing base of culm 0.5-1.5 mm 

wide. Leaves 1-3, basal; sheaths elongate, pale 

reddish green to red with dark red veins, the 

lowermost often bladeless or with short blades; 

ligule absent; blades short, narrowly to broadly 

U-shaped proximally, involute distally, adaxially 

cellular-reticulate and spongy-thickened, often 

with a medial ridge, glossy, whitened or greenish, 

abaxially green, the midvein indistinct, 0.4-18 cm 

× 0.3-1.3 (-1.5) mm, antrorsely scabrous on the 

margins, attenuate to apex. Inflorescence a single, 

pseudolateral spike, or rarely 1-2 smaller, lateral, 

secondary ones present near its base, (5-) 6-14 

(-20) mm in diam.; involucral bracts (1-) 2-3, leaflike, 

the lowest well-developed, erect and 

appearing as a continuation of the culm, exceeding 

the inflorescence, the upper bracts setaceous, often 

divergent to reflexed; rays essentially absent; 

spikes 1 (-3), small, with (1-) 4-36 (-43) densely 

disposed spikelets; spikelets linear-subquadrate, 

flexuose, 3-10 (-13) × 0.8-1.5 mm, acuminate at 

apex, cuneate at base, with 3-12 florets; rachilla 

flexuose, broadly hyaline-winged; scales widely 

ovate-elliptic, dorsally acute, concavely curvatekeeled, 

1.6-2.5 × 1.4-2 mm, subherbaceous, with 

7-9 fine and indistinct nerves on each side above 

the scarious margins which clasp the achene at 

maturity, stramineous or yellowish or reddish 

brown, carina 3-nerved, greenish, prolonged 

beyond the subobtuse to acute apex as a short 


apiculate; styles 3-branched. Achene trigonous 

with plane faces, curvate on adaxial face, narrowly 

ellipsoid to ellipsoid-obovoid, 1.1-1.5 × 0.5-0.8 

mm, sub-rounded at apex, apiculate, short-cuneate 

at base, estipitate, puncticulate, brown to dark 





Islands: Primarily in the southwest region of 

Puerto Rico on limestone in dry scrub forest, 

thickets, and beaches along the coast. Cabo Rojo, 

Caja de Muertos, Guánica, Guayama, Guayanilla, 

Lajas, Mona Island, Ponce, San Juan, and Yauco; 

Anegada, Guana Island, St. Croix, and Tortola.

292 


Common name: Puerto Rico: Pajón de costa. 


Cabo Rojo: Bo. Boquerón, N end, Balneario 

Público de Boquerón, Proctor & McKenzie 43752 

(US).Guayanilla: Guánica State Forest, Proctor 

43386 (US). Guayama: Caja de Muertos, N.L. 

Britton et al. 4976 (UPR). Mona Island: Sardinera, 

Otero & Chardón 941 (US); Along E side of 

island, Proctor & Haneke 43112 (US). Yauco: Bo. 

Barina, Guánica Forest Reserve, Axelrod et al. 

10897 (UPRRP). TORTOLA: Guana Island, Proctor 

43700 (US). 

41. Cyperus unioloides R. Br., Prodr. 216. 1810; 

Pycreus unioloides (R. Br.) Urb., Symb. 

Antill. 2: 164. 1900. Type: Australia; 

Queensland. R. Brown 5900 (holotype: BM; 

isotype: K). 

Cyperus bromoides Willd. ex Link, Jahrb. 

Gewächsk. 3: 85. 1820. Type: South America. 

Humboldt s.n. (holotype: B-Willd. 1353). 

Cyperus angulatus Nees in Wight, Contr. Bot. 

India. 73. 1834; Pycreus angulatus (Nees) 

Nees, Linnaea 9: 283. 1835. Type: Nepal. 

Wallich 3324 a,b (holotype: K). 

Cyperus pseudobromoides Boeck., Linnaea 35: 

464. 1868. Syntypes: Nepal; Hispaniola. Not 

located. 


tall; rhizome short, 3-8 mm thick; roots coarse, 

yellowish or orangish. Culms triquetrous or 

subtrigonous proximally, the sides flat to concave, 

smooth, 1.3-3.5 mm wide; sheathing bases 3-8 mm 

wide. Leaves (1-) 2-5 (-6), basal; sheaths elongate, 

becoming fibrillose with age, brown to dark 

reddish brown, the inner band finely veined; ligule 

absent; blades flat or folded, often spiraling, 10- 

60 (-90) cm × 2.5-5 mm (unfolded), antrorsely 

scabrous on margins and midvein abaxially 

towards apex, smooth to remotely antrorsely 

scabrous adaxially, long-acuminate to triquetrous 

apex. Inflorescence a simple to rarely compound 

umbel-like corymb with ascending rays, 4-12 × 

3-10 cm; involucral bracts 1-4, leaf-like, 7-40 

(-60) cm long; rays (1-) 2-5 (-7), unequal, patent, 

up to 11 cm long, the tubular prophylls dark 

reddish brown, at least near base; spikes 2-4 × 2.2- 

3.5 cm, with 3-17 (-30) spicately disposed 

spikelets; spikelets lanceolate to ovate-lanceolate 

or oblong-lanceolate, flattened, 7-22 (-27) × 3-5 

mm, acute at apex, obtuse at base, with 7-25 (-33) 

florets; rachilla quadrangular, unwinged, the scales 

and achenes deciduous from the rachilla at 

maturity; scales ovate to ovate-lanceolate, dorsally 

acute, 3-5 × 2-2.6 mm, light-brown to yellowish 

brown on sides, pale brown and often reddish 

brown lineolate on either side of carina, sometimes 

tinged with dark brown basally, 1-nerved on each 

side paralleling carina above the entire to distally 

narrowly scarious, involute margins, carina 3nerved, 

greenish, prolonged beyond the acute apex 

as a short mucro. Stamens 3, the anthers 2-2.8 mm 

long, rounded-apiculate at apex; styles 2-branched. 

Achene biconvex, obovate to suborbicular, 1.1- 

1.3 × 0.8-1.1 mm, rounded to a short-beaked apex, 

short-cuneate at base, estipitate, outer epidermal 

layer shiny, smooth, silvery-gray, reticulate when 

mature, the inner surface minutely papillose, dull, 

blackish. 

General distribution: Pantropical. Greater 

Antilles, Mexico, Central America, South 

America, Africa, Malesia, and Australia. 

Distribution in Puerto Rico: In swamps or 

marshy areas; known only from Río Abajo State 

Forest and vicinity in northwestern Puerto Rico. 

Arecibo, Lares, and Utuado. 


Arecibo: Bosque de Río Abajo, Acevedo-Rdgz. et 

al. 2259 (NY, US); Río Abajo Forest Reserve, area 

S of Rt. 521, ca. 1.2 km W of Reserve office, 

Axelrod et al. 10915 (UPRRP, US). Lares: Sargent 

3250 (US). Utuado: Bosque de Río Abajo, Saltillo, 

Acevedo-Rdgz. 332 (SJ). 

42. Cyperus urbanii Boeck., Beitr. Cyper. 1: 10. 

1888; Mariscus urbanii (Boeck.) C. B. Clarke 

in Urban, Symb. Antill. 2: 47. 1900. 

Lectotype: Puerto Rico; Aibonito. Sintenis 

2929 (NY!), here designated. 


short; roots fine. Culms ascending, often arched, 

triquetrous, soft, flexuous, antrorsely scabrous on 

margins, finely ribbed, 0.3-0.9 mm wide. Leaves 

3-5, ascending, basal and lower cauline; sheaths 

short, subinflated, thinly herbaceous, light brown 

to brown, often reddish tinged, with dark red veins; 

ligule absent; blades narrowly linear, V-shaped to 

folded proximally, flattened distally, 4-40 (-53) cm 

× (1-) 1.2-3 mm, herbaceous, finely and closely 

veined, semi-glossy and cellular-reticulate 

adaxially, at least proximally, margins and abaxial 

midvein with both antrorsely and retrorsely set


barbs, attenuate to triquetrous tip. Inflorescence a 

single, terminal, or a terminal and 1-2 lateral 

essentially sessile spikes at the summit of the culm, 

(4-) 4.5-8 (-11) × 4-7 (-9) mm; involucral bracts 

3-5, the lowermost leaf-like, 1-14 cm long, 

uppermost short, linear-setaceous, ascending to 

divergent at maturity; spikes ovoid to oblongovoid, 

with 16-40 (-80) spikelets; spikelets oblongovoid, 

compressed-quadrate, (1.7-) 2-3.5 (-4) × 

0.7-1 mm, acuminate at apex, short-cuneate at 

base, with 2-4 florets; rachilla hyaline-winged; 

scales widely ovate-elliptic to subrounded, 1.4- 

1.7 × 1.1-1.5 mm, dorsally acute, submembranous, 

whitish stramineous with blackish longitudinal 

lineations, 5- to 9-nerved on sides, margins 

scarious proximally, rounded to an abruptly acute 

to short-acuminate apex, carina 3-nerved, green, 

ending in a excurved cusp at apex. Stamens 3, the 

anthers 0.4-0.5 mm long, apiculate; style 3branched. 

Achene trigonous, ellipsoid-obovoid, 

with the adaxial face plane to slightly concave and 

abaxial faces slightly convex, 1-1.4 × 0.6-0.7 mm, 

obtuse at apex, apiculate, short-cuneate at base, 

estipitate, finely puncticulate with an often faintly 

mottled surface, light brown to brown. 


Distribution in Puerto Rico: In upland 

habitats (150-830 m) on rocky shaded slopes, 

ridges, hillsides, and pastures. Aibonito, Cabo 

Rojo, Coamo, Salinas, and Vieques. 


Coamo: Bo. Santa Catalina, Rte. 155, Axelrod et 

al. 1567 (US); Bo. Cuyón, N slopes of Las Piedras 

Chiquitas, Proctor & Carrasquillo 42486 (US); 

Las Piedras Chiquitas, Bo. Cuyón, along spur from 

house at Rt. 717, km 2.1 to ridge of peaks, Axelrod 

& Sastre 5596 (US). 

43. Cyperus virens Michx., Fl. Bor.-Amer. 1: 28. 

1803. Type: United States; “Carolina”. 

Michaux s.n. (holotype: P; photo at GH). 

Cyperus formosus Vahl, Enum. Pl. 2: 327. 1805; 

Cyperus surinamensis var. formosus (Vahl) 

Kük., Repert. Spec. Nov. Regni Veg. 32: 74. 

1933. Type: “Habitat in Louisiana et 

Caribaeis” Collector unknown (holotype: P- 

Juss.; isotype C-Vahl). 

Caespitose, often coarse perennial, (20-) 55- 

140 (-170) cm tall; rhizome short, thick and 

hardened, 0.5-1 cm thick; roots coarse. Culms 

erect, triquetrous with sharply acute angles, firm 

but subspongy, predominantly antrorsely scabrous 

on angles, sometimes retrorse or extrorse barbs 

intermixed, (2-) 3-7 (-10) mm wide; sheathing 

bases 6-16 (-30) mm wide. Leaves 4-12, basal and 

lower cauline; sheaths relatively short, crossveined, 

subspongy-thickened proximally, pale 

brown, tinged with red proximally; ligule absent; 

blades linear, V-shaped proximally, subflattened 

or subplicate distally, (10-) 30-130 cm × 4-14 mm, 

conspicuously cross-veined abaxially, margins and 

abaxial midvein scabrous with fine antrorse, 

retrorse or extrorse barbs often intermixed, longacuminate 


simple to compound, somewhat open or often 

contracted, umbel-like corymb with ascending 

rays, 4-10 (-16) × 4.5-16 cm; involucral bracts 4- 

14, leaf-like, ascending to spreading, 13-70 (-100) 

cm long; primary rays 6-12, short to elongate, up 

to 12 (-14) cm long, secondary rays often present, 

shorter, several tertiary rays sometimes present; 

spikes ovoid, 1-2.5 cm in diam., with 8-40 

digitately disposed, densely clustered spikelets; 

spikelets narrowly ovate to oblong-lanceolate, 4- 

8 (-10) × 2-3.2 mm (excluding denuded rachilla), 

compressed, obtuse at apex, broadly acute at base, 

with 10-22 florets; rachilla unwinged; scales 

lanceolate-elliptic, 1.6-2.3 (-2.8) × 1-1.4 mm, boatshaped, 

dorsally 2-keeled, herbaceous or thickly 

so medially, glossy with a finely cellular-reticulate 

or -pebbled surface, pale reddish brown or 

greenish, dark reddish lineolate adaxially, with 1 

distinct or sometimes indistinct nerve on each side 

above the scarious margins, carina 3-nerved, the 

lateral nerves forming two pronounced keels 

proximally creating a declivity between each 

lateral keel and the midnerve, all three nerves 

converging into a single one at the acute apex, there 

prolonged as a short mucro, greenish or brownish, 

glabrous or occasionally scabrellate distally. 

Stamens 1-2, the anthers 0.8-1.4 mm long, with a 

linear-prickly appendage at tip; style 3-branched. 

Achene trigonous, with flat or slightly convex or 

concave faces, ovoid-ellipsoid or ellipsoid, 1-1.5 

× 0.3-0.6 mm, gradually tapered to an acute apex, 

subulate-beaked, short-cuneate at base, slightly to 

distinctly short-stipitate, finely puncticulate, 

brown to grayish brown. 


States, Mexico, Central America, West Indies, and 

South America.

294 


Distribution in Puerto Rico: In wet or swampy 

areas along river banks, pond borders, pastures, 

and ditches. Bayamón, Fajardo, San Juan, and Toa 

Baja. 


Bayamón: Finca Santa Ana, Hioram 230 (US). Toa 

Baja: Rd. 866, km 7.2, to Punta Salinas, González- 

Más 1144 (US). 


Cyperus cuspidatus Kunth in Humb., Bonpl., & 

Kunth., Nov. Gen. Sp. 1 [quarto ed.]: 204. 

1816. Reported for Anegada by Britton and 

P. Wilson (Bot. Porto Rico 5: 82. 1923). This 

record is based on a misidentification of a 

specimen of Cyperus squarrosus from 

Anegada. 

Cyperus ferrugineus Poir. in Lamarck, Encycl. 7: 

261. 1806. Cited by C. B. Clarke (in Urban, 

Symb. Antill. 2: 18. 1900) for St. Thomas as 

Pycreus ferrugineus (Poir.) C. B. Clarke based 

on a specimen (no collector given) at the 

British Museum (BM). Kükenthal (1936: 374) 

cites this, but treats it as Cyperus filicinus 

Vahl. Cyperus ferrugineus is strictly an 

African species and C. filicinus has not been 

recorded outside of the United States. No 

specimens have been seen of either taxon from 

Puerto Rico or the Virgin Islands in the 

preparation of this treatment. 

Cyperus hermaphroditus (Jacq.) Standl., Contr. 

U.S. Natl. Herb. 18: 88. 1916. The specimen 

record for this species (Sintenis 1256b from 

Fajardo, Puerto Rico) was treated as Mariscus 

elatus Vahl by C. B. Clarke (in Urban, Symb. 

Antill. 2: 46. 1900), Mariscus incompletus 

(Jacq.) Urb. by Urban (Symb. Antill. 4: 13. 

1903), and Cyperus coriifolius Boeck. by 

Kükenthal (in Engler, Pflanzenr. IV, 20 (Heft 

Eleocharis R. Br., Prodr. 224. 1810. 

6. ELEOCHARIS 

101): 492. 1936) who treated this latter taxon 

as distinct from C. hermaphroditus. However, 

Kükenthal (l.c. 492) doubts its occurrence in 

the West Indies. This specimen was not seen 

by McLaughlin (1944) nor located by us in 

order to confirm its identity. 

Cyperus humilis Kunth, Enum. Pl. 2: 23. 1837. 

Reported for Anegada by D’Arcy in “Anegada 

Island: Vegetation and Flora”, Atoll. Res. Bull. 

No. 188: 15. 1975, based on D’Arcy 4906A 

(MO!). This specimen is Cyperus squarrosus 

L. 

Cyperus retroflexus Buckley, Proc. Acad. Nat. Sci. 

Philadelphia 14: 9. 1862. Reported for Puerto 

Rico by Kartesz (1998) in his “A 

Synonymized Checklist of the Vascular Flora 

of the United States, Puerto Rico, and the 

Virgin Islands” (http://www.csdl.tamu.edu/ 

FLORA/b98/check98.htm). However, no 

specimen record exists to confirm its identity 

(Kartesz, pers. comm., 2001). 

Cyperus luzulae (L.) Rottb. ex Retz., Observ. Bot. 

4: 11. 1786. C. B. Clarke (in Urban, Symb. 

Antill. 2: 28. 1900) cited this taxon for Puerto 

Rico based on a collection of Bertero s.n. This 

taxon was later reported for Puerto Rico by 

Britton and P. Wilson (Bot. Porto Rico 5: 84. 

1923) based on Bertero s.n. (near Bayamón) 

and Hioram s.n.; Kükenthal (in Engler, 

Pflanzenr. IV, 20 (Heft 101): 170. 1936) 

without any citation given; and McLaughlin 

(1944: 51) based on Fredholm 4722. The 

Hioram and Fredholm specimens are Cyperus 

surinamensis Rottb. Neither the Bertero 

specimen, or any other collection of this taxon 

was seen by us in order to confirm its 

occurrence in Puerto Rico. 

Perennials or occasionally annuals, rhizomatous, stoloniferous, or tufted; culms usually cylindrical, 

sometimes trigonous, rarely quadrangular, solid or hollow, sometimes transversely septate, smooth, 

green, glabrous. Leaves reduced to bladeless sheaths; sheaths closed, tubular, the orifice truncate to 

obliquely truncate, dorsally carinate, the carina often prolonged as a short apiculum, sometimes with a 

puckered, scarious appendage above the apex, glabrous. Inflorescence a single, terminal spikelet at the 

summit of the culm; spikelets ovoid to lanceoloid, rarely obovoid, obtuse to acute, several- to many-


flowered, with spirally imbricate or sometimes subdistichous scales, the lowermost 1-2 sterile; scales 

oblong, oblong-ovate to lanceolate, or obovate, sometimes subrounded, firm or membranous, usually 

with hyaline margins, carinate or sometimes indistinctly so, nerved or nerveless on sides, glabrous. 

Flowers bisexual; perianth bristles present or rarely absent, (3-) 6 (-12), retrorsely barbed, rarely smooth; 

stamens (2) 3, the anthers elliptic to linear; styles capillary, 2- or 3-branched, with expanded base. 

Achene trigonous or biconvex, obovate or elliptic, smooth, cancellate, or cellular-reticulate; style base 

expanded, triangular or conical, sometimes spongy-thickened, persistent at the apex of the achene. A 

cosmopolitan genus with approximately 250 species. 

LECTOTYPE: Eleocharis palustris (L.) Roem. & Schult. (≡ Scirpus palustris L.), designated by Britton, 

Bull. Dept. Agric. Jamaica 5, Suppl. 1: 10. 1907. 

References: Svenson, H. K. 1929. Monographic studies in the genus Eleocharis. Rhodora 31: 224- 

242; Svenson, H. K. 1939. Monographic studies in the genus Eleocharis. Rhodora 41: 1-77, 93-110. 

Key to the species of Eleocharis 

1. Spikelets linear-cylindrical; spikelet scales firm, subcartilaginous (at least medially), rounded-obovate 

or subrounded, generally ca. as long as broad................................................................... 2 

2. Culms appearing nodose with conspicuous transverse septae ........................ 7. E. interstincta 

2. Culms appearing smooth, without conspicuous transverse septae........................................... 3 

3. Culm sharply trigonous, rarely obtusely so; bristles retrorsely barbed their entire length 

...................................................................................................................... 10. E. mutata 

3. Culms terete or essentially so; bristles smooth or sometimes several retrorse barbs present 

at apex ............................................................................................ 1. E. cellulosa 

1. Spikelets ovoid, lanceoloid, or obovoid; spikelet scales thin, membranaceous or herbaceous, ovate, 

ovate-lanceolate, ovate-elliptic, oblong-ovate, oblong-elliptic, or obovate to spatulate, generally longer 

than broad............................................................................................................................. 4 

4. Culms appearing nodose with conspicuous transverse septae................................................. 5 

5. Culms (1.3-) 2-10 mm wide; spikelets (3-) 4-9 mm wide; scales obovate to spathulate with 

an oblong limb, abruptly broadly obtuse or rounded to apex; achenes 1-1.5 × 0.5-0.7 mm; 

style base lanceolate, 0.7-1 mm long; bristles 6-8 .......................... 3. E. elegans 

5. Culms 1-2.5 mm wide; spikelets 2-4 mm wide; scales ovate to ovate-lanceolate, tapering to 

slightly dialated apex; achenes 0.9-1.1 × 0.6-1 mm, style base deltate to depressed-deltate, 

0.3-0.4 mm long; bristles 4-6 ......................................................... 9. E. montana 

4. Culms appearing smooth, without conspicuous transverse septae........................................... 6 

6. Style 2-branched; achenes biconvex................................................................................. 7 

7. Mature achenes finely reticulate, dull, yellowish ....................................... 4. E. fallax 

7. Mature achenes smooth and shiny, black or brown................................................... 8 

8. Summit of sheath conspicuously whitish hyaline and cross-puckered above the 

orifice; mature achene brown ..................................................... 5. E. flavescens 

8. Summit of sheath firm, or hyaline with a slender green carina dorsally, not 

conspicuously cross-puckered above the oblique or subtruncate orifice; mature 

achenes black.................................................................................................... 9 

9. Caespitose annual or short-lived perennial not forming basal spikelets; spikelets 

subglobose or ovoid, 2.5-4 mm long; fertile spikelet scales herbaceous, elliptic, 

1.5-2 mm long, dark green to brown with pale margins; wide-ranging, lowland, 

coastal plain species .......................................................... 6. E. geniculata 

9. Caespitose perennial, often with horizontally creeping rhizomes bearing sessile 

basal spikelets; spikelets ovoid-ellipsoid to ellipsoid-lanceoloid, 3.5-7 (-9) 

mm long; fertile spikelet scales membranous, ovate to oblong-ovate, 2-3 mm 

long, stramineous, often purple to purplish brown on sides medially; upland, 

montane species, restricted to Sierra de Luquillo and Sierra de Naguabo, 

450-1065 m ........................................... 2. E. debilis

296 


6. Style 3-branched; achene trigonous or compressed-trigonous....................................................... 10 

10. Achenes terete or nearly so, narrowly oblong or oblong-obovoid, the surface with longitudinal 

ridges separated by numerous cross-walls (trabeculae) ............... 13. E. radicans 

10. Achenes trigonous or compressed-trigonous, obovoid, the surface smooth or cancellate. 11 

11. Spikelet narrowly obovoid or obconical ................................................ 12. E. pachystyla 

11. Spikelet ovoid to oblong-ovoid, ovoid-ellipsoid, or lanceoloid.................................. 12 

12. Medium-sized perennial, not forming basal spikelets; culms (0.5-) 0.7-1.8 (-2.2) mm 

wide; style base nearly confluent with summit of achene, seemingly a continuation of 

it but of a slightly different texture .......................... 15. E. rostellata 

12. Slender, delicate annuals or perennials, often forming sessile spikelets at culm bases; 

culms 0.2-0.4 (-0.5) mm wide; style base triquetro-pyramidal. ................... 13 

13. Achenes conspicuously cancellate; style base decurrent on shoulders of achene 

................................................................................................... 14. E. retroflexa 

13. Achenes smooth or faintly reticulate; style base articulate with apex of achene. 

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

14. Spikelet scales dark purplish brown medially; achenes (including style base) 

0.7-1 × 0.4-0.6 mm; bristles 3-6 or rudimentary ....... 11. E. oligantha 

14. Spikelet scales stramineous, often faintly tinged with yellow-brown; achenes 

(including style base) 0.5-0.6 × 0.3-0.4 mm; bristles absent, rarely several 

rudimentary ones present ....................................... 8. E. minutiflora 

1. Eleocharis cellulosa Torr., Ann. Lyceum Nat. 

Hist. New York 3: 298. 1836. Type: United 

States; Mississippi. Ingalls s.n. (holotype: 

NY!; isotype: GH). 

Coarse stoloniferous perennial 30-80 (-125) 

cm tall; stolons elongate, stout, 2-3 mm thick, 

glossy, brown to chocolate-brown; roots coarse. 

Culms strongly ascending to erect, 3-5 mm wide, 

terete or obtusely angled, rarely trigonous, soft, 

spongy, easily compressed, with network of 

internal air cavities, very finely and shallowly 

ribbed, finely and bluntly antrorsely scabrous 

along ribs distally, greenish or stramineous. 

Proximal sheaths inflated or subinflated, 

submembranous, light green to stramineous, often 

tinged with brown or purple-brown distally; distal 

sheaths herbaceous, light green to light brown or 

stramineous, oblique at orifice, acute to broadly 

acute at apex with a long-subulate apiculum 

dorsally. Spikelets cylindrical, 1.5-4 (-5) cm × 3.2- 

4.5 (-5) mm, acute to obtuse or sub-rounded at 

apex, cuneate at base, many-flowered, the florets 

spirally imbricate; fertile scales obovate, 

subrounded, or widely ovate-rhombic, 3.5-4.7 

(-5) × 2.7-4 (-4.5) mm, dorsally obtuse to rounded, 

obtuse at apex, thickly herbaceous to 

subcoriaceous medially, stramineous with a 

conspicuous or sometimes inconspicuous brown 

border and narrow white scarious margins, reddish 

lineolate and dotted adaxially, finely veined 

abaxially, carina and lateral nerves indistinct; 

sterile basal scales 1, similar to fertile, the lower 

margins broadly clasping base of spikelet. Stamens 

3, the anthers 1.7-2.2 mm long, apiculate; style 3branched. 

Achene biconvex, elliptic-obovate or 

widely elliptic, 1.5-2 × 1.3-1.5 mm, abruptly 

narrowed to a truncate, low rim-like apex that 

extends distally on margins of style base, cuneate 

at base, cellular-reticulate with 17-21 longitudinal 

rows of often transversely oriented rectangular 

cells, glossy, light brown to brown; style base 

trigonous, thickened medially, 0.5-0.8 mm long, 

0.5-0.8 mm wide at base, subspongy, subconfluent 

with the thickened rim-like apex of achene; bristles 

6-7, closely appressed to achene body, subflattened 

proximally, light brown, smooth to sometimes a 

few antrorse barbs present at tips, equaling or 

slightly longer than achene body. 


States, Mexico, Central America, and the West 

Indies. 

Distribution in Puerto Rico: Occurring in 

north-central Puerto Rico only in shallow water 

or wet areas of lake margins, swales, and marshes. 

Manatí, San Juan, and Vega Baja. 

Common name: Puerto Rico: Junco fino.



Manatí: Bo. Tierras Nuevas Saliente, area just S 

of Laguna Tortuguero, Proctor 42687 (US). 

2. Eleocharis debilis Kunth, Enum. Pl. 2: 143. 

1837. Type: Brazil; Rio de Janeiro. Sellow s.n. 


Eleocharis shaferi Britton, Mem. Torrey Bot. Club 

16: 59. 1920. Type: Cuba; Oriente. Shafer 

3414 (holotype: NY!; isotype: NY!). 

Eleocharis yunquensis Britton in Britton & P. 


Puerto Rico; Luquillo Mountains. Britton & 

Bruner 7619 (holotype: NY!; isotypes: GH, 

UPR! US!). 

Eleocharis atropurpurea sensu Urban, Symb. 

Antill. 4: 116. 1903, non (Retzius) J. Presl & 

C. Presl, 1828. 

Eleocharis sintenisii of authors, non Boeckeler, 

1888. 

Rhizomatous perennial, 4-40 cm tall, often 

forming spikelets at culm bases; rhizomes slender, 

horizontally creeping or forming a dense mat; roots 

fine. Culms crowded along rhizome, often in 

dense, spreading clumps, erect to ascending, 

filiform, coarsely 4- to 6-ribbed, 0.3-0.7 mm wide. 

Sheaths subinflated, proximal ones membranous, 

purple to purple-black proximally, stramineous 

distally, distal sheaths thinly herbaceous, green to 

stramineous, often angled with 3-4 green marginal 

nerves, dorsally septate-nodulose distally, the 

orifice oblique to subtruncate, membranous, 

dorsally acuminate with green carina. Spikelets 

ovoid-ellipsoid to ellipsoid-lanceoloid, 3.5-7 (-9) 

× (1-) 1.3-2.7 (-3) mm, acute at apex, cuneate at 

base, 4- to 20-flowered with spirally imbricate 

florets, the scales spreading; fertile scales ovate 

to oblong-ovate, 2-3 × 1-1.5 mm, boat-shaped, 

acutely to obtusely keeled dorsally, stramineous, 

often purple to purplish brown on sides, finely 

striate, with a slender green carina ending short of 

the obtuse to acute apex, lateral nerves indistinct; 

sterile basal scales 1-2, ovate-elliptic, similar in 

size to fertile scales, herbaceous, obtuse to rounded 

at apex, the carina 3- to 5-ribbed, thickened 

distally. Stamens 2-3, the anthers 0.4-0.7 mm long, 

apiculate; style 2-branched. Achene biconvex, 

narrowly obovate, 1-1.4 × 0.6-0.8 mm, black, 

shining, finely and faintly transversely rugulose 

to essentially smooth; style base conical to conic- 

subulate, 0.4-0.6 mm long, 0.3-0.4 mm wide at 

base; bristles 6, unequal, whitish to light brown, 

retrorsely barbed from tip to near base, shorter than 

to equaling or exceeding the achene body. 

General distribution: Cuba and Puerto Rico, 

Central America, and South America. 

Distribution in Puerto Rico: Wet areas of 

montane forest, generally in higher elevations of 

the Sierra de Luquillo and Sierra de Naguabo, 450- 

1065 m; forest edges and openings, river banks, 

stream edges, creek and river beds, and roadside 

ditches. Ceiba, Naguabo, and Río Grande. 


Ceiba/Naguabo: Caribbean National Forest, 

Axelrod 8352 (US). Naguabo: Bo. Río Blanco, 

Caribbean National Forest, Axelrod & Chávez 

4122 (US); Sierra de Naguabo, vicinity of La 

Florida, Shafer 3138 (US). Río Grande: Sierra de 

Luquillo, Proctor 43192, 43200 (US); El Yunque, 

McKee 10643 (US); Luquillo Forest, top of El 

Yunque, González-Más 1532 (US); Luquillo 

Forest, Road towards El Yunque mountain, 


3. Eleocharis elegans (Kunth) Roem. & Schult., 

Syst. Veg. 2: 150. 1817; Scirpus elegans 


Sp. [quarto ed.] 1: 226. 1816. Type: 

Venezuela. Humboldt & Bonpland s.n. 

(holotype: P-HBK). 

Eleocharis constricta Schult., Mant. 2: 87. 1824; 

Scirpus constrictus (Schult.) Griseb., Syst. 

Veg. Karaiben: 122. 1857. Type: Brazil. Wied- 

Neuwied s.n. (holotype: P). 

Eleocharis geniculata sensu C. B. Clarke in Urban, 

Symb. Antill. 2: 74. 1900 and sensu Britton 

& P. Wilson, Bot. Porto Rico 5: 92. 1923, non 

(Linnaeus) Roem. & Schult., 1817. 

Coarse, rhizomatous perennial, (10-) 30-150 

(-170) cm tall; rhizome short, rarely elongated, 

ascending to horizontal, ligneous, 5-10 mm thick. 

Culms shortly ascending to erect, crowded along 

rhizome, (1.3-) 2-10 mm wide, terete, easily 

compressed, prominently transversely septate, the 

intervals between the septae 6-30 (-40) mm long 

proximally, becoming increasingly shorter distally, 

glaucous. Sheaths thinly herbaceous, finely 

veined, truncate at orifice, often with an 

inconspicuous subulate mucro dorsally, reddish to 

reddish purple, red-dotted, finely black-maculate

298 


at apex. Spikelets lanceoloid to lanceoloidellipsoid, 

7-30 × (3-) 4-9 mm, narrowly obtuse to 

acute at apex, truncate to broadly rounded at base, 

densely many-flowered, the florets spirally 

imbricate; fertile scales membranous, obovate to 

spathulate with an oblong limb, 2-3 × 0.7-1.5 mm, 

dorsally obtuse to rounded, pale, opaque brown 

to yellow-brown medially, margins broadly 

scarious, red-brown lineolate, obscurely 3-nerved 

medially, carina indistinct, ending short of the 

obtuse to sub-rounded apex; sterile basal scales 

2-3, sub-rounded to deltoid, 1.5-2.5 × 2-3 mm, 

obtuse at apex, green, glaucous, with pale scarious 

margins. Stamens 3, the anthers 1-1.2 mm long, 

apiculate; style 2- or 3-branched. Achene biconvex 

to slightly trigonous, obovate-elliptic to elliptic, 

1-1.5 × 0.5-0.7 mm, finely and shallowly reticulate 

with irregularly disposed, oblong or somewhat 

isodiametric cells, semi-glossy, yellow to yellowbrown; 

style base laterally compressed, lanceolate 

or deltate-lanceolate, 0.7-1 mm long, 0.3-0.5 mm 

wide at base, annular-thickened at base, articulate 

with apex of achene, brown to grayish brown; 

bristles 6-8, unequal, forming a short stipe at base 

of achene, subulate, brown, closely retrorsely 

spinulose nearly to base, exceeding achene, 

equaling to slightly overtopping apex of style base. 

General distribution: Central Mexico, Central 


Distribution in Puerto Rico: In wet habitats, 

generally at middle to higher elevations, 100-1050 

m; forest edges, marshy areas, margins of pools, 

springy depressions, edges of creeks, streams, 

riverbanks, meadows, pasture, cane fields, 

roadside ditches, and disturbed areas. Adjuntas, 

Aguadilla, Aguas Buenas, Aibonito, Añasco, 

Arecibo, Bayamón, Caguas, Cayey, Ceiba, 

Corozal, Fajardo, Humacao, Jayuya, Lares, 

Luquillo, Maricao, Mayagüez, Naguabo, 

Orocovis, Patillas, Ponce, Rincón, Río Grande, 

San Juan, San Lorenzo, Utuado, and Vega Baja. 

Common name: Puerto Rico: Junquillo. 


Adjuntas: Bo. Limaní, Axelrod & Montalvo 11600 

(UPRRP). Aibonito: Sintenis 2915 (US). Añasco: 

Heller 4534 (US). Arecibo; Río Abajo State 

Forest, Acevedo-Rdgz. 10680 (US). Bayamón: 

Hioram 311 (US). Caguas: Underwood & Griggs 

304 (US). Cayey: Rd. 184, Guavate, km 5.7, 

González-Más 1578 (US). Maricao: Indiera Fría, 

near Maricao, N.L. Britton et al. 4534 (US). 

Mayagüez: N.L. Britton 2361 (US). Naguabo: 

Sierra de Naguabo, Mouth of Río Icaco, Shafer 

3164 (US). Patillas: Bo. Muñoz Rivera, off Rt. 

184, Axelrod & Weisel 11126 (UPRRP, US). Ponce: 

Hills above Ponce, Sargent 383 (US). Río Grande: 

Along Rt. 186 bordering Caribbean National 

Forest, just N of the El Verde Field Station, Strong 

et al. 380 (GMUF). San Juan: Río Piedras, Heller 

& Heller 170 (US). Utuado: Vicinity of Utuado, 

N.L. Britton & Cowell 416 (US). 

4. Eleocharis fallax Weath., Rhodora 24: 23. 

1922. Type: United States; Massachusetts. 

Fernald & Long 18025, (holotype: GH; 


Eleocharis nervosa Kük., Repert. Spec. Nov. 

Regni Veg. 23: 192, 1926. Type: Cuba; Santa 

Clara, Lagua la Grande, Ekman 18543 

(holotype: S; isotype: NY!). 

Eleocharis ambigens Fernald, Rhodora 37: 394. 

1935. Type: United States; Virginia. Fernald 

& Long 3765 (holotype: GH). 


tall; rhizomes elongate, horizontally creeping, 

slender, flexuous, reddish, 1-2.5 mm thick with 

dark red striate scales; roots medium to coarse. 

Culms in small tufts along rhizome, very slender, 

subterete or slightly compressed, 0.5-1.5 (-1.7) mm 

wide, shallowly ribbed, light green, often some 

cells with light reddish pigment giving the culms 

and sheaths a speckled appearance. Sheaths 

herbaceous, the lower purple-black (at least 

proximally) the upper light green, often tinged with 

red, orifice slightly oblique to subtruncate, shortmucronate 

dorsally, the margin slightly thickened, 

red-punctate when young, turning brownish with 

age. Spikelets ovoid-ellipsoid to lanceoloid, (6-) 

7-10 (-16) × 1.5-3 mm, acute to obtuse or subrounded 

at apex, obtuse at base, many-flowered, 

the florets spirally imbricate; fertile scales ovateelliptic 

or oblong-ovate to lanceolate, 2-3.7 (-4) × 

1-1.6 mm, obtuse to sub-rounded at apex, thinly 

herbaceous to submembranous, dark brown to dark 

red on sides with green mid rib and broad pale 

scarious apex and margins; fertile basal scales 1- 

2, ovate to broadly ovate, like the fertile. Stamens 

3, the anthers 0.6-1.5 mm long with a lanceolate 

appendage at apex bearing crystalline papillae; 

style commonly 2-branched, sometimes 3branched, 

the branches scaly-fimbriate. Achene


biconvex to obtusely trigonous, obovoid, 1.2-1.7 

× 0.8-1.1 mm, finely and shallowly reticulate, 

yellow to brown; style base obtusely pyramidal, 

with a collar-like ring at base, 0.4-0.6 mm long, 

0.2-0.6 mm wide at base, wider at base than point 

of attachment, grayish; bristles 3-4 (-5), subulate, 

flattened proximally, reddish, retrorsely barbed 

from apex to near base, unequal, shorter than to 

1-2 sometimes as long as achene body. 

General distribution: Eastern United States 

and Greater Antilles. 


Islands: Wet, muddy areas; lake borders, fields, 

and pastures. Guánica and Lajas; St. Croix. 


Lajas: Bo. Palmarejo, NW corner of village of 

Maguayo, Proctor 46385 (US). 

5. Eleocharis flavescens (Poir.) Urb., Symb. 

Antill. 4: 116. 1903; Scirpus flavescens Poir. 

in Lamarck, Encycl. 6: 756. 1804. Type: 

Puerto Rico. Ledrú s.n. (holotype: P; isotype: 

P-2). 

Scirpus flaccidus Rchb. ex Spreng. f., Tent. Suppl. 

3. 1828; Eleocharis albivaginata var. flaccida 

(Rchb. ex Spreng. f.) Boeck., Linnaea 36: 437. 

1869; Eleocharis ocreata var. flaccida (Rchb. 

ex Spreng. f.) Boeck. ex C. B. Clarke in 

Urban, Symb. Antill. 2: 63. 1900; Eleocharis 

flaccida (Rchb. ex Spreng. f.) Urb., Symb. 

Antill. 2: 165. 1900. Type: Surinam. Weigelt 


Eleogenus ocreatus Nees in Martius, Fl. Bras. 2: 

102. 1842. Eleocharis ocreata (Nees) Steud., 

Syn. Pl. Glumac. 2: 79. 1854; Scirpus ocreatus 

(Nees) Griseb., Pl. Wright. 2: 535. 1862. Type: 

Brazil. Macrae s.n. (syntype: CGE); Macrae 

s.n. (syntype: CGE); von Martius s.n. 

(syntype: M); and Beyrich s.n. (syntype: B, 

destroyed). 

Eleocharis albivaginata var. humilis Boeck., 

Linnaea 36: 438. 1870; Eleocharis ocreata 

var. humilis (Boeck.) C. B. Clarke in Urban, 

Symb. Antill. 2: 64. 1900. Type: Puerto Rico. 

Collector unknown (holotype: B-Willd. 

1162). 

Stoloniferous perennial, (1-) 4-40 (-50) cm 

tall; rhizome horizontal to ascending, 0.6-1.2 mm 

thick. Culms caespitose or spaced singly along the 

rhizome, erect to ascending, subcompressed, 0.4- 

1 mm wide, prominently ribbed. Sheaths subloose, 

coarsely veined, membranous, green, red-tinged 

near base, with a prolonged, inflated, puckered, 

scarious, appendage above the apex, the apex 

short-mucronate dorsally. Spikelets ovoid to 

ovoid-ellipsoid, 3-7 × 1.5-3 mm, subacute, broadly 

cuneate at base, with many spirally imbricate 

florets; fertile scales boat-shaped, elliptic to 

narrowly-elliptic, 1.2-2.1 × 0.8-1.1 mm, broadly 

obtuse dorsally, pale green to stramineous, carina 

narrow, 3- to 5-nerved, subthickened at tip, ending 

short of the obtuse to subacute apex; sterile basal 

scale 1, narrowly-ovate, 1.4-2.2 × 1-1.2 mm, with 

broad finely and closely nerved, prominent, green 

carina which is thickened at apex. Stamens 3, the 

anthers 0.7-1 mm long, apiculate; styles 2branched, 

the branches minutely scaly, the 

unbranched portion smooth. Achene biconvex, 

obovate, 0.8-1 (including style base) × 0.6-0.7 mm, 

smooth to finely cancellate, lustrous, olive brown 

to dark reddish brown; style base conic, 0.2-0.3 

mm long, 0.2 mm wide at base, light brown; 

bristles 6-8, rarely absent, whitish to tawny, 

retrorsely barbed, equaling or exceeding the style 

base. 

General distribution: United States 

(predominately southeastern), Mexico, Central 

America, West Indies, and northern and eastern 



Islands: In wet mud or peat of marshy or grassy 

areas, river banks, border of marshes, forest 

borders, depressions and trails in montane forest, 

pasture, roadside ditches and banks, and disturbed 

areas. Adjuntas, Aibonito, Añasco, Arecibo, 

Barranquitas, Bayamón, Caguas, Canóvanas, 

Cayey, Ceiba, Ciales, Dorado, Florida, Humacao, 

Jayuya, Juncos, Luquillo, Manatí, Maricao, 

Naguabo, Orocovis, Patillas, Río Grande, San 

Juan, San Lorenzo, San Sebastián, Utuado, Vega 

Alta, Vega Baja, and Villalba; St. Thomas. 


Adjuntas: Monte Cerrote, near Adjuntas, N.L. 

Britton & Brown 5405 (US). Añasco: Heller 4532 

(US). Arecibo: Río Abajo State Forest, Acevedo- 

Rdgz. 10732 (UPRRP, US). Bayamón: Sintenis 

1222 (US). Canóvanas: Sierra de Luquillo, Proctor 

& Taylor 46297 (US). Cayey: Rd. 184, El Guavate, 

km 9.6, González-Más 1575 (US). Juncos: Sintenis 

2656 (US). Luquillo: Luquillo Forest (Mt. Britton), 

González-Más 1508 (US). Manatí: Sintenis 6685

300 


(US). Maricao: Sintenis 176 (US). Naguabo: Sierra 

de Naguabo, Loma Icaco, Shafer 3414 (US). 

Orocovis: above Divisoria, N.L. Britton & E.G. 

Britton 10137 (US). Río Grande: Luquillo 

Mountains: El Yunque, Stimson 1956 (US). San 

Juan: Martín Peña, Stevenson & Johnston 1678 

(US). Vega Alta: Bo. Sabana, ca. 2 km due NE of 

jct. Rds. 690 & 691, Proctor 43215 (US). Villalba: 

Toro Negro Forest Reserve, González-Más 2001 

(US). 

6. Eleocharis geniculata (L.) Roem. & Schult., 

Syst. Veg. 2: 150. 1817; Scirpus geniculatus 

L., Sp. Pl. 48. 1753; Eleocharis capitata R. 

Br., Prodr. 225. 1810, nom. illeg. Lectotype: 

Jamaica. Collector unknown. Herb. Clifford: 

21, Scirpus 1 (BM), designated by Furtado, 

Gard. Bull. Straits Settlements 9: 299. 1937. 

Scirpus caribaeus Rottb., Descr. Pl. Rar. 24. 1772; 

Eleocharis caribaea (Rottb.) S.F. Blake, 

Rhodora 20: 24. 1918. Type: St. Croix, U.S. 

Virgin Islands. Collector unknown (holotype: 

C-Rottb.). 

Scirpus geniculatus var. minor Vahl, Enum. Pl. 2: 

251. 1805; Eleocharis geniculata var. minor 

(Vahl) Roem. & Schult., Syst. Veg. 2: 150. 

1817. Type: Jamaica. Collector unknown 

(holotype: probably at C-Vahl). 

Eleocharis sintenisii Boeck., Beitr. Cyper. 1: 16. 

1888. Lectotype: Puerto Rico; Bayamón. 

Sintenis 1220 (NY!), here designated. 

Scirpus capitatus sensu Grisebach, Fl. Brit. W. I. 

570. 1864, non Linnaeus, 1753. 

Fig. 50. A-E 

Densely tufted annual or short-lived 

perennial, 2-60 cm tall. Culms obscurely angled, 

subcompressed, 0.3-1 mm wide, prominently 

ribbed. Sheaths finely many-veined, smooth, 

green, reddish to purplish proximally, oblique at 

summit, with a concave to truncate orifice, 

prolonged dorsally into a short, acuminate, 

cuspidate, thickened tip, slightly shorter than the 

acute to acumiante apex. Spikelets ovoid, ovoidellipsoid, 

of globose-ovoid, rarely ovoidlanceoloid, 

2-4.5 (-6) × 1.3-3.3 mm, obtuse, obtuse 

or sub-rounded at base, many-flowered, the florets 

spirally imbricate; fertile scales herbaceous, 

elliptic, 1.3-2 × 1-1.3 mm, boat-shaped to 

shallowly so, dorsally broadly rounded, dark green 

to brown with pale margins, carina obscurely 

nerved, ending short of the obtuse apex, the basal 

scales elliptic to widely elliptic; sterile basal scale 

1, 1.2-1.5 × 1.0-1.5 mm, 1-nerved, light green to 

straw-colored, smooth and shiny, with pale green 

mid rib thickened distally, tipped by an expanded 

callose. Stamens 3, the anthers 0.5-0.6 mm long, 

apiculate; styles 2-branched. Achene biconvex, 

obovate, 0.9-1.1 (including style base) × 0.6-0.8 

mm, smooth, lustrous, dark purplish black to black; 

style base depressed-conic, saucer-shaped, 0.2-0.3 

mm long, 0.3-0.5 wide at base, whitish; bristles 

6-8, rarely absent, whitish to tawny, retrorsely 

barbed, equaling or exceeding the style base. 

General distribution: In tropical and 

subtropical regions worldwide. 


Islands: Often on wet, sandy or gravelly soils of 

river banks, river beds, rock crevices and ledges 

along rivers, pond and lagoon margins, springy 

seepage areas, savannas, marshy and swampy 

areas, borders of mangroves, swales, pastures, 

roadsides ditches, and disturbed areas. Aibonito, 

Añasco, Arecibo, Arroyo, Bayamón, Cabo Rojo, 

Caguas, Carolina, Cataño, Cayey, Ceiba, Coamo, 

Dorado, Guánica, Guayama, Isabela, Lajas, Loíza, 

Manatí, Maricao, Mayagüez, Morovis, Naguabo, 

Patillas, Quebradilla, Río Grande, San Juan, San 

Sebastián, Toa Baja, Trujillo Alto, Vega Alta, Vega 

Baja, Vieques, and Villalba; Anegada, St. Croix, 

St. John, St. Thomas, and Tortola. 

Note: Small plants with acute spikelets, 

appearing transitional between this species and 

Eleocharis bahamensis Boeck., occur on Puerto 

Rico and Anegada, e.g., Proctor 43067 (SJ!, US!) 

and D’Arcy 5144 (MO!) respectively. None of the 

characters cited by Correll & Correll (1982) as 

differentiating the two species were consistent in 

material examined for the flora area. 


Añasco: Sintenis 5617 (US). Bayamón: Sintenis 

1219 (US), 1220 (NY). Cabo Rojo: (Joyuda), road 

102, González-Más 570 (US). Cataño: Back of 

Cataño, Heller 6412 (US). Cayey: Sintenis 2344 

(US). Coamo: Sintenis 1968, 3041, 3145 (US); 

Underwood & Griggs 528 (US). Dorado: Dorado 

to Cerro Gordo road, km 13, González-Más 294 

(US). Guánica: González-Más 954 (US). 

Guayama: Puerto Jobos, road 707, González-Más 

655 (US). Loíza: Bo. Loíza Aldéa, just NE of town 

and E of the Río Grande de Loíza, Proctor 43067 

(US). Mayagüez: Limón Road, González-Más 

1272 (US). Naguabo: Rd. 31, km 29.6, Río


Blanco, González-Más 1377 (US). Río Grande: 

Rd. 187, km 145, González-Más 1446 (US). San 

Juan: Santurce, Heller & Heller 585 (US); Laguna 

San José, N.L. Britton & E.G. Britton 7179 

(US).Toa Baja: Palo Seco, Hioram 815 (US). Vega 

Baja: Tortuguero Lagoon, González-Más 1054 

(US). Vieques Island: Resolución to Punta Arenas, 

Shafer 2893 (US). ST. JOHN: Reef Bay Quarter; 

along Fish Bay Gut, Acevedo-Rdgz. 2497 (US, 

VINPS). ST. THOMAS: Jumbee Gut, Eggers s.n. 

(NA). 

7. Eleocharis interstincta (Vahl) Roem. & 

Schult., Syst. Veg. 2: 149. 1817; Scirpus 

interstinctus Vahl, Enum. Pl. 2: 251. 1805. 

Type: West Indes. Martfelt s.n. (holotype: C- 

Vahl). 

Coarse, stoloniferous perennial, (18-) 40-120 

(-152) cm tall; rhizome with a short caudex, 

emitting coarse stolons, 2-4 mm thick; roots 

coarse, light brown or reddish brown. Culms erect, 

stout, (2-) 4-9 (-11) mm wide, terete, hollow, easily 

compressed, prominently and conspicuously 

transversely septate, the intervals between the 

septae 5-35 (-60) mm long, becoming approximate 

distally below the inflorescence. Sheaths often 

loose, membranous to herbaceous, pale green, 

faintly to strongly mottled or tinged with redbrown, 

smooth, glossy, the orifice obliquely 

truncate with a broadly triangular-acute to abruptly 

acuminate summit dorsally. Spikelets cylindrical, 

(15-) 25-50 (-70) × 3.5-6 (-7) mm, obtuse at apex, 

many-flowered, the florets spirally imbricate; 

fertile scales broadly oblong-ovate, broadly 

oblong-obovate, or subrounded, 4-6 × 2.3-5 mm, 

obtuse to broadly rounded at apex, dorsally obtuse 

to rounded, chartaceous, finely striate, stramineous 

or grayish green, obscurely 3- to 8-nerved 

medially, the margins broadly scarious; sterile 

basal scales 2-3, semi-circular to subdeltoid, 

subcoriaceous, with white-hyaline margins. 

Stamens 3, the anthers 1.8-3 mm long, with a 

subulate apiculum; style 2- or 3-branched. Achene 

thickly biconvex, obovate to elliptic-obovate or 

broadly obovate, (1.3-) 1.6-2 × 1-1.8 mm, 

subabruptly narrowed to a truncate, pronounced, 

annular-thickened apex, short-attenuate to obtuse 

at base, semi-glossy, with prominent transversely 

oriented linear cells arranged in longitudinal rows, 

yellowish brown or gray; style base lanceolate, 

flattened, subdeltate basally, 0.6-1.5 mm long, 0.8- 

1 mm wide at base, gradually tapering to the style, 

dark brown; bristles 7-8, stout, flattened, reddish, 

coarsely retrorsely spinulose nearly to base, 

exceeding the achene body, shorter than to 

exceeding tip of style base. 

General distribution: Widespread in 

subtropical and tropical America; southern United 

States (Florida, Alabama, Oklahoma, and Texas), 

Mexico, Central America, West Indies, and South 

America. 


Islands: In water or wet habitats; borders of 

lagoons, ponds, marshy and swampy areas, stream 

edges, pasture, roadside ditches, and disturbed 

areas. Añasco, Arecibo, Caguas, Cataño, Cidra, 

Corozal, Dorado, Florida, Guánica, Guayama, 

Guayanilla, Humacao, Luquillo, Manatí, 

Mayagüez, Naguabo, Peñuelas, Río Grande, San 

Juan, San Sebastián, Utuado, Vega Baja, and 

Yauco; St. Croix, and St. Thomas. 

Common names: Puerto Rico: Junco, Junco 

de aparejos, Junco de ciénaga, Junco de espiga. 


Arecibo: Bosque de Río Abajo, Los Puercos, 

Acevedo-Rdgz. 149 (SJ). Cataño: Rd. 165, km 5, 

González-Más 1163 (US). Guayama: Puerto Jobos: 

Rd. 707, González-Más 694 (US). Guayanilla: 

Guayanilla to Tallaboa, Shafer 1977 (US). Manatí: 

SE side of Laguna Tortuguero, Strong et al. 420 

(GMUF). Mayagüez: Las Mesas, Holm s.n. (US). 

Naguabo: Rd. 3, km 41, González-Más 1364 (US). 

Vega Baja: Laguna Tortuguero, González-Más 

1059 (US). ST. CROIX: Jealousy, J.J. Ricksecker 

252 (US). ST. THOMAS: Signalhill, Eggers s.n (US). 

8. Eleocharis minutiflora Boeck., Bot. Jahrb. 

Syst. 7: 274. 1886; Eleocharis nigrescens var. 

minutiflora (Boeck.) Svenson, Rhodora 39: 


Islands, Krum Bay. Eggers 546 (holotype: B, 

destroyed; isotypes: AMD, CAS, GOET, L). 

Eleocharis minima sensu C. B. Clarke in Urban, 

Symb. Antill. 2: 70. 1900 (in part), non Kunth, 

1837. 

Eleocharis microcarpa sensu Britton & P. Wilson, 

Bot. Porto Rico 5: 92. 1923, non Torrey, 1836. 

Delicate perennial, 2-12 cm tall, forming 

small clumps; rhizome short, ascending to nearly 

vertical, several-branched; roots capillary. Culms 

ascending to reflexed, filiform, 0.2-0.4 (-0.5) mm

302 


wide, finely ribbed, several-channeled, semiglossy. 

Sheaths short, membranous, few to absent, 

rarely more than one per culm, stramineous to 

reddish tinged, the orifice oblique, narrowed to 

an acute to acuminate scarious apex dorsally. 

Spikelets compressed, ovoid to ovoid-ellipsoid or 

oblong-ovoid, (1.5-) 1.7-3 × 1-1.5 mm, acute, 

rounded-cuneate at base, loosely several-flowered 

with spirally imbricate or subdistichous florets, the 

scales spreading; fertile scales ovate, boat-shaped, 

0.8-1 × 0.6-0.8 mm, thinly herbaceous to 

membranous, acutely to obtusely keeled dorsally, 

with 1- to 3-nerved green to pale green carina, 

stramineous, often faintly tinged with yellowbrown, 

short-mucronate at the acute to subobtuse 

entire or short-cleft apex; sterile basal scale 1, 

ovate, with broad green, 5-veined carina and 

broadly scarious margins, the apex obtuse. 


apiculate; style 3-branched. Achene trigonous with 

convex sides, narrowly obovate, 0.5-0.6 × 0.3-0.4 

mm, obtuse at apex, cuneate at base, distinctly 

costate on angles, finely and indistinctly cellularreticulate, 

essentially smooth, semi-glossy; style 

base triquetro-pyramidal, 0.1-0.2 long, 0.2-0.3 mm 

wide at base, shallowly articulate with apex of 

achene, brown; bristles absent or 1-2 short 

rudimentary ones present. 

General distribution: Mexico (Yucatan), 

Cuba, and St. Thomas. 

Distribution in the Virgin Islands: St. Thomas. 

Note: Eleocharis minutiflora consistently 

differs from its closest relative E. nigrescens in 

the lighter colored spikelet scales; distinctly 

costulate achenes with a smoother surface (finely 

and indistinctly reticulate); and more pyramidal 

style base (vs. depressed-pyramidal in E. 

nigrescens). 


Crum Bay, Eggers 767 (US). 

9. Eleocharis montana (Kunth) Roem. & Schult., 

Syst. Veg. 2: 153. 1817; Scirpus montanus 


Sp. [quarto ed.] 1: 226. 1816. Type: Colombia. 

Humboldt & Bonpland s.n. (holotype: P- 

HBK). 

Scirpus nodulosus Roth, Nov. Pl. Sp. 29. 1821; 

Eleocharis nodulosa (Roth) Schult., Mant. 2: 

87. 1824. Type: Brazil. Collector unknown 



tall; rhizome decumbent, stout, 4-8 mm thick. 

Culms erect, cylindrical, striate, conspicuously 

transversely septate or sometimes inconspicuously 

so, 1-2.5 mm wide, intervals between septa 1-5 

mm long. Sheaths elongate, thinly herbaceous, 

finely striate, purplish red proximally, stramineous 

or purple-tinged distally, red-dotted, the orifice 

truncate with blackish maculate horizontal band 

and a tooth-like mucro dorsally. Spikelets ovoidlanceoloid 

to oblong-lanceoloid, 7-15 (-24) × 2-4 

mm, acuminate or rarely obtuse at apex, manyflowered, 

the florets spirally imbricate; fertile 

scales ovate-lanceolate, (1.5-) 2-2.5 × 0.8-1.7 mm, 

dorsally rounded, membranous to thinly 

herbaceous medially, light brown, often dark 

brown medially, reddish lineolate, margins broadly 

scarious, carina slender, 1-nerved, indistinct, 

ending short of the obtuse or rounded, slightly 

dialated apex; sterile basal scale 1, subrounded, 

herbaceous, the margins broadly scarious. Stamens 

3, the anthers 0.8-1 mm long; style 2- or 3branched. 

Achene biconvex, obovate to broadly 

obovate, 0.9-1.1 × 0.6-1 mm, cuneate at base, 

rounded to subtruncate at apex, distinctly and 

finely pitted-reticulate, yellow to brown or 

olivaceous; style base flattened, deltate to 

depressed-deltate, 0.3-0.4 mm long, 0.4-0.7 mm 

wide at base, shallowly articulated with apex of 

achene body; bristles 4-6, unequal, forming a short 

stipe where commonly joined at base, minutely 

retrorsely spinulose or somewhat smooth, equaling 

or slightly longer than achene body, ferrugineous 

or brownish. 


Mexico, Central America, West Indies, and tropical 



Islands: In wet habitat; marshy or swampy areas, 

along forest trails, and roadside ditches. Aibonito, 

Naguabo, and Río Grande; cited for St. Croix by 

Fosberg, Rhodora 78: 86. 1976. 


Aibonito: N.L. Britton et al. 8252 (US). Naguabo: 

Rd. 191, km 21.1, González-Más 1397 (US). 

10. Eleocharis mutata (L.) Roem. & Schult., Syst. 

Veg. 2: 155. 1817; Scirpus mutatus L., Syst. 

Veg. ed. 10. 867. 1759. Lectotype: Jamaica. 

Browne s.n. (LINN-71.2), designated by 

Browning et al., S. Afr. J. Bot. 63: 172. 1997.


Coarse, stoloniferous perennial, (15-) 25-100 

(-120) cm tall; rhizome short, emitting coarse, dark 

brown stolons, 2-4 mm thick. Culms caespitose, 

erect, acutely trigonous to triquetrous, rarely 

obtusely trigonous, finely striate, (1.3-) 2-5 (-7) 

mm wide. Sheaths elongate, membranous, semiglossy, 

stramineous or light brown, tinged with 

red, often darkly so proximally, obliquely truncate 

at orifice, prolonged dorsally, the slender midvein 

excurrent and forming a cuadate appendage 

beyond the acute to acuminate apex. Spikelets 

cylindrical, (15-) 20-50 × 4-6 mm, obtuse, little 

wider than the culm, many-flowered, the florets 

spirally imbricate; fertile scales rounded-obovate 

to rounded, 2.7-5 × 2-4.5 mm, subcartilaginous 

medially, dorsally obtuse, often subcarinate, 

indistinctly finely and closely veined, stramineous 

medially, brown-lineolate, the margins broadly 

scarious, subtranslucent, carina 1-nerved, 

indistinct abaxially, distinct adaxially, ending short 

of the obtuse apex; sterile basal scales 0-1, like 

the fertile, the basal bract nearly encircling base 

of spikelet. Stamens 3, the anthers 1.3-2.3 mm 

long, with a lanceolate, reddish papillate apiculum; 

style 3-branched. Achene biconvex, broadly 

obovate, 1.5-2 × (1-) 1.2-1.5 mm, cuneate at base, 

rounded to an annular-thickened apex which 

merges gradually into the short style base, 

shallowly reticulate with 21-24 longitudinal rows 

of transversely oriented oblong cells, shiny, yellow 

to orange-brown; style base short, compresseddeltate, 

0.2-0.6 mm long, 0.6-0.8 mm wide at base, 

annular-thickened at base; bristles 7-8, unequal, 

retrorsely spinulose with coarse, soft spinules, 

equaling to slightly overtopping achene, lustrous 

brown. 


(Texas), Mexico, Central America, West Indies, 

South America, and tropical Africa. 


Islands: In water or wet habitats, generally in 

coastal areas; marshes, swamps, swales, shallow 

pools, borders of lakes and ponds, estuaries, 

channels, borders of mangroves, pastures, and 

roadside ditches. Arecibo, Arroyo, Cabo Rojo, 

Cataño, Culebra, Dorado, Fajardo, Guánica, 

Guayama, Humacao, Isabela, Loíza, Luquillo, 

Manatí, Naguabo, Río Grande, Salinas, San Juan, 

Vega Baja, Vieques, and Yabucoa; Anegada, St. 

Croix, and Tortola. 


Arroyo: Estuary - Guilarte Beach, González-Más 

1323 (US). Cataño: Rd. 165, km 5, González-Más 

1164 (US). Culebra Island: N.L. Britton & Wheeler 

172 (US). Guánica: Guánica Lake, Sargent 39 

(US). Humacao: Pozal, playa de Humacao, Eggers 

s.n. (US). Río Grande: Bo. Zarzal, vicinity of Punta 

Picúa, Proctor 43544 (US). Salinas: Rd. 705, km 

1, González-Más 719 (US). San Juan: Martín Peña, 

Stevenson 263 (US). Vega Baja: Tortuguero 

Lagoon, González-Más 1056 (US). Vieques Island: 

Playa Grande to La Mina, Shafer 2999 (US). 

Yabucoa: Sintenis 4942 (US). ST. CROIX: 

Bethlehem Gut, J.J. Ricksecker 414 (US); St. 

John’s, A.E. Ricksecker 210 (US). 

11. Eleocharis oligantha C. B. Clarke in Urban, 

Symb. Antill. 2: 69. 1900. Type: Cuba. Wright 

3367 (syntype: K; isosyntype: G); 3368 

(syntype: K; isosyntype: G, NY). 

Scirpus retroflexus sensu Grisebach, Cat. Pl. Cub. 

239. 1866, non Poiret, 1805. 

Delicate, densely matted, often proliferous 

annual, 1-15 cm tall, often forming spikelets at 

culm bases. Culms often recurved, hair-like, finely 

capillary, quadrangular-sulcate, minutely punctate, 

0.1-0.3 mm wide. Sheaths membranous, reddish 

proximally, stramineous distally, the orifice 

hyaline, subinflated, oblique, with a dorsal green 

carina which ends short of the obtuse apex; 

spikelets sublaterally compressed, ovate, 1.8-5.5 

× 1-2 mm, acute to acuminate, obtuse at base, with 

3-15 florets, the florets spirally imbricate or 

subdistichous; fertile scales spreading with 

developing achenes, ovate to ovate-lanceolate, 1.2- 

2.2 × 0.6-1 mm, submembranous, light brown, 

dark purplish brown medially, finely and closely 

nerved, broadly hyaline on margins, carina 1nerved 

or closely 3-nerved, green, equaling or 

shortly prolonged beyond the acute to obtuse apex 

as a mucro; sterile basal scales 1, hyaline except 

for the carina, oblong-ovate, slightly shorter than 

the fertile, broadly rounded at apex. Stamens 2-3, 

the anthers 0.4-0.7 mm long, apiculate; style 3branched. 

Achene trigonous, indistinctly costate 

on angles with slightly convex sides, obovoid, 0.7- 

1 (including style base) × 0.4-0.6 mm, sub-rounded 

at apex, short-cuneate at base, faintly reticulate, 

shiny, pale, becoming gray to dark olive-brown at 

maturity; style base pyramidal to depressedpyramidal, 

acute, 0.1-0.2 mm long, 0.3-0.5 mm 

wide at base, somewhat 3-pronged at base, light

304 


brown; bristles 3-6 or rudimentary, finely 

retrorsely spinulose, shorter than to as long as the 

achene body. 

General distribution: Central and western 

Cuba, Jamaica, and Puerto Rico. 


from a single locality; muddy border of a small 

pond in silica-sand area of north-central Puerto 

Rico; Vega Alta. However, according to G. Proctor 

(pers. comm., 2003), it grows in a pond in Vega 

Baja as an aquatic extreme previously thought by 

him to be Eleocharis confervoides (see excluded 

species at the end of Eleocharis descriptions). 


Vega Alta: Bo. Sabana, silica-sand area NE of 

Regardera, 1.7-2.2 km due NE of jct. Rds. 690 

and 691, Proctor 44145, 44572, 45796 (US). 

12. Eleocharis pachystyla (C. Wright) C. B. 

Clarke in Urban, Symb. Antill. 2: 72. 1900; 

Scirpus pachystylus C. Wright in Sauvalle, 

Anales Acad. Ci. Méd. Habana 8: 79. 1871, 

as a new name for Scirpus melanocarpus 

Griseb., Cat. Pl. Cub. 239. 1866, non Torrey, 

1836. Type: Cuba; Pinar del Río. Wright 3373 

(holotype: GOET; isotype: GH). 


rhizome short, horizontal or branched-ascending; 

roots medium. Culms ascending to sub-erect , 0.7- 

2 (-3) mm wide, terete to flattened-sulcate when 

dry, rather soft, shallowly ribbed, finely cellularreticulate, 

light green, glossy. Sheaths tight, 

purple-red, often light green to stramineous 

distally or only at orifice, thinly herbaceous, 

thickened-membranous at the oblique orifice, 

acute at apex dorsally. Spikelets obovoid, 5-7 (-8) 

× 2-3.5 mm, obtuse at apex, often clavate at base, 

the florets spirally imbricate; fertile scales 

numerous, ovate to oblong-ovate or oblongelliptic, 

2-3 × 1-1.7 mm, thinly herbaceous, 

dorsally obtuse, stramineous, brown-lineolate 

proximally, margins scarious, carina indistinct, 

green or green-dotted distally, not prolonged 

beyond the obtuse to subtruncate apex; sterile basal 

scales 3-10 or more, the uppermost like the fertile, 

the lowermost 1-2 broadly ovate or subrounded, 

thickly herbaceous, with broad green to dark green 

carina. Stamens 3, the anthers 0.6-0.8 mm long; 

style 3-branched. Achene trigonous, narrowly 

obovoid or obconical, 0.8-1.1 × 0.6-0.8 mm, 

rounded at apex, short-acuminate at base, finely 

costate on margins, essentially smooth, shiny, light 

brown to yellowish brown; style base flattenedpyramidal, 

sulcate on sides, sometimes subfalcate 

and decurrent on shoulders of achene, 0.5-0.8 mm 

long, 0.5-0.6 mm wide at base, light brown to 

brown; bristles 6, filiform, retrorsely spinulose, 

brown or reddish, slightly shorter than to equaling 

or slightly exceeding achene body. 



Distribution in Puerto Rico: Low elevation, 

wet, often peaty, marshy habitats of silica-sand 

areas in the vicinity of Laguna Tortuguero; Dorado 

and Vega Alta. 


Vega Alta: Bo. Sabana, silica-sand area NE of 

Regardera (1.7-2.2 km due NE of jct. Rds. 690 

and 691, Proctor 42892, 43282 (US). Dorado: 

Vicinity of Dorado, N.L. Britton et al. 6674 (US). 

13. Eleocharis radicans (A. Dietr.) Kunth, Enum. 

Pl. 2: 142. 1837; Scirpus radicans Poir. in 

Lamarck, Encycl. 6: 751. 1805, non Schkuhr, 

1793; Eleogiton radicans A. Dietr., Sp. Pl. 2: 

97. 1833; Eleocharis acicularis var. radicans 

(A. Dietr.) Britton, J. New York Microscop. 

Soc. 5: 105. 1889. Type: Puerto Rico. Ledrú 

s.n. (holotype: probably at P). 

Eleocharis acicularis var. lindheimeri C. B. Clarke 

ex Britton, J. New York Microscop. Soc. 5: 

105. 1889; Eleocharis lindheimeri (C. B. 

Clarke ex Britton) Svenson, Rhodora 31: 199. 

1929. Type: United States; Texas. F. 

Lindheimer 1315 (holotype: K, isotype: GH). 

Slender and delicate, caespitose perennial, 2- 

9 (-13) cm tall, often forming dense mats; rhizomes 

capillary, horizontal to ascending. Culms borne 

singly or in tufts along rhizome, ascending, 

compressed-trigonous, soft and spongy, finely 

ribbed, dark green, 0.3-0.6 (-1) mm wide. Sheaths 

membranous, tightly surrounding culm, oblique 

at orifice, dorsally with a narrow, thickened tip at 

the subacute apex. Spikelets ovoid to lanceoloid 

or ovoid-ellipsoid, 2-4 × 0.8-1.7 mm, with 6-15 

florets, the florets spirally imbricate; all scales 

fertile, ovate to narrowly ovate-elliptic or ovatelanceolate, 

1.4-2.3 × 0.6-1.1 mm, membranous, 

stramineous with broad scarious margins, carina 

green, 5- to 7-nerved, converging and ending in a


thickened tip short of the scarious, obtuse apex. 

Stamens 2 or 3, the anthers 0.3-0.5 mm long, 

apiculate; style 3-branched. Achene subterete, 

narrowly ellipsoid-obovoid, 0.7-1.1 × 0.3-0.5 mm, 

narrowly rounded at apex, cuneate at base, 

longitudinally ribbed with 3 coarser ribs on angles 

and cross-lined with fine close trabeculae; 

yellowish; style base narrow, 0.2-0.4 mm long, 0.2 

mm wide at base, conic-triangular, articulate at 

base with apex of achene; bristles 4-7, capillary, 

whitish, barbed, usually exceeding the achene and 

style base. 

General distribution: United States, West 

Indies, Mexico, Central America, and temperate 


Distribution in Puerto Rico: Recorded at 

present only in the Botanic Garden University of 

Puerto Rico, San Juan. It is cited on the specimen 

label as being cultivated there, but it is very likely 

adventive. Previously known only from the type. 


San Juan: Liogier & Liogier 29030 (UPR; photo 

at US). 

14. Eleocharis retroflexa (Poir.) Urb., Symb. 

Antill. 2: 165. 1900; Scirpus retroflexus Poir. 

in Lamarck, Encycl. 6: 753. 1805. Type: 

Puerto Rico. Ledrú s.n. (holotype: P; isotype: 

P). 

Cyperus depauperatus Vahl, Enum. Pl. 2: 305. 

1805. Eleocharis depauperata (Vahl) Kunth, 

Enum. Pl. 2: 140. 1837. Type: West Indies. 

Rohr s.n. (holotype: probably at C-Vahl). 

Eleocharis chaetaria sensu C. B. Clarke in Urban, 

Symb. Antill. 2: 68. 1900, non Roemer & 

Schultes, 1817. 

Caespitose annual, 1-20 cm tall, often forming 

spikelets at culm bases. Culms usually recurved, 

filiform, star-shaped or subflattened, sometimes 

bluntly scabrous on margins, obscurely punctate 

0.2-0.4 mm wide. Sheaths subloose and inflated, 

cross-puckered, membranous, stramineous to 

reddish tinged, scarious at obliquely truncate 

orifice, prolonged dorsally as a lanceolate 

appendage, the slender carina excurrent beyond 

the acuminate apex. Spikelets ovate or elliptic, 2.5- 

4 (-4.3) × 1.2-1.8 mm, sublaterally compressed, 

obtuse to acute, cuneate at base, 2- to 8-flowered, 

the florets subdistichously disposed, often 

spreading with maturing achenes; fertile scales 

ovate to ovate-lanceolate, 1.7-2.5 × 1-1.7 mm, 

membranous, obtuse to acute at apex, boat-shaped, 

dorsally acute, often with chestnut to reddish or 

purplish brown medially on sides, margins broadly 

scarious, carina pale green, 3-nerved; sterile basal 

scales 1, ovate-elliptic, like the fertile. Stamens 3, 

the anthers 0.4-0.6 mm long, apiculate; style 3branched. 

Achene trigonous, obovoid to urceolate, 

0.6-1.1 × 0.4-0.6 mm, subattenuate to base, with 

prominent costate angles and convex sides, pittedreticulate 

with 7-10 longitudinal rows of 

isodiametric cells on each side, whitish or 

stramineous, maturing olive-yellow; style base 

pyramidal-acuminate, the angles decurrent on 

shoulders of achene, with slightly upturned tips, 

0.3-0.4 mm long, 0.3-0.5 mm wide at base, light 

brown; bristles 6, subulate, retrorsely spinulose 

distally, whitish to pale, shorter than to 1 or 2 

sometimes equaling or slightly exceeding style 

base. 


(Alabama), tropics and subtropics of Mexico, 

Central America, West Indies, South America, 

Asia, Malesia, and Australia. 


Islands: In wet mud or sand of forested areas, 

generally at middle to higher elevations, on eroded 

slopes, river and creek banks, stream beds, trails, 

pastures, roadside banks, and ditches. Adjuntas, 

Arecibo, Bayamón, Caguas, Carolina, Cayey, 

Fajardo, Guayama, Mayagüez, Naguabo, Patillas, 

Río Grande, San Juan, and San Sebastián; cited 

by C. B. Clarke (1900) and Britton (1918) for St. 

Thomas. 



Britton & Brown 5406 (US). Arecibo: Río Abajo 

State Forest, along limon trail, Acevedo-Rdgz. 

10667 (US). Bayamón: Sintenis 1209 (US). 

Carolina: Sabana Abajo, N.L. Britton & E.G. 

Britton 9199 (US). Cayey: Rd. 184, Guavate, km 

5.7, González-Más 1577 (US). Fajardo: Colonia 

San Miguel, N.L. Britton & Shafer 1630 (US). 

Mayagüez: Near Hacienda Alicia, Liogier 9926 

(US). Naguabo: Sierra de Naguabo, Bo. de 

Maizales, N.L. Britton et al. 2108 (US). Río 

Grande: Sierra de Luquillo, Sintenis 1403 (US). 

San Juan: Río Piedras, Hioram 919 (US). San 

Sebastián: Sargent 242 (US). 

15. Eleocharis rostellata (Torr.) Torr., Fl. New 

York 2: 347. 1843; Scirpus rostellatus Torr.,

306 


Ann. Lyceum Nat. Hist. New York 3: 318. 

1836. Type: United States; New York. 

Sartwell s.n. (lectotype: US!; isolectotypes: 

GH, NY), here designated. 

Coarse rhizomatous perennial forming 

tussocks, 10-60 (-80) cm tall; rhizome short and 

thickened, erect; roots coarse. Culms erect to 

ascending, some often arching or reclining and 

rooting at tip, (0.5-) 0.7-1.8 (-2.2) mm wide, 

flattened, sulcate. Sheaths coriaceous, subloose, 

drab to chocolate brown proximally, light brown 

to stramineous or light green distally, dorsally 

obtuse to subacute, midcosta abruptly dialated and 

thickened near apex, the orifice suboblique to 

nearly truncate, maculate, with a subthickened, 

reddish brown or blackish margin. Spikelets ovoid 

to ovoid-lanceoloid or fusiform, 6-20 × 2-3.5 mm, 

acute at apex, many-flowered, the florets and 

subtending scales often arranged in a subloose 

spiral on the rachis, this causing the spikelet to 

appear as if twisted on its axis; fertile scales ovate 

to ovate-lanceolate, 3-5 × 1.8-3 mm, thickly 

herbaceous, stiffish, drab or brown, margins 

thickly scarious, obtuse at apex; sterile basal scales 

2 or sometimes all scales of spikelet fertile, similar 

to the fertile, widely ovate or subrounded, often 

with widely scarious margins and rounded apex. 

Stamens 3, the anthers 2-2.7 mm long, bluntly 

apiculate; style 3-branched (on all specimens 

studied). Achene obtusely trigonous, obovoid to 

ellipsoid-obovoid, 1.4-2 × 1-1.5 mm, narrowed at 

summit into the confluent pyramidal style base, 

obtuse at apex, acute to subcuneate at base, 

yellowish brown to olive brown, smoothish, finely 

and indistinctly cellular-reticulate, lustrous; style 

base obtusely pyramidal to lance-pyramidal, 

seemingly a continuation of the achene apex but 

of a slightly different texture; bristles 6, subulate, 

closely retrorsely spinulose, sometimes appearing 

subplumose, reddish, equaling to exceeding tip of 

style base. 

General distribution: Southern Canada, 

United States, Mexico, West Indies, and Andean 


Distribution in Puerto Rico: Wet, marshy or 

boggy habitats in white silica sand areas of northcentral 

Puerto Rico. Arecibo, Manatí, and Vega 

Baja. 


Arecibo: Bo. Islote, border of Ciénaga, Proctor & 

Rivera 46411 (US). Vega Baja: Bo. Algorrobo, 

white sand area just S of Laguna Tortuguero, 

Proctor et al. 47747 (US). Manatí: Tortuguero, 

Woodbury s.n. (US); Bo. Tierras Nuevas Saliente, 

Woodbury s.n. (US). 


Eleocharis atropurpurea (Retz.) J. Presl & C. Presl 

in C. Presl, Reliq. Haenk. 1: 196. 1828. 


Island: Vegetation and Flora”, Atoll. Res. Bull. 

No. 188: 15. 1975, based on D’Arcy 5144 

(MO!), but later identified by F.J. Menapace 

as E. bahamensis Boeck. C. B. Clarke (1900) 

also reported this species for Puerto Rico 

based on Sintenis 1220. Both D’Arcy 5144 

and Sintenis 1220 are Eleocharis geniculata 

(L.) Roem. & Schult. No specimens of 

Eleocharis atropurpurea or E. bahamensis 

from Puerto Rico or the Virgin Islands have 

been seen in the preparation of this treatment. 

Eleocharis confervoides (Poir.) G. C. Tucker: Cited 

as Websteria confervoides (Poir.) S.S. Hooper 

by Liogier & Martorell (2000: 247) as 

collected by G. Proctor at Vega Baja. No 

specimens of this taxon have been seen nor 

has it been recently collected in Puerto Rico. 

According to G. Proctor (pers. comm. 2003) 

the Vega Baja plant represents an aquatic 

extreme of Eleocharis oligantha C. B. Clarke. 

Eleocharis pachycarpa E. Desv. in C. Gay, Fl. 

Chil. 6: 174. 1853. Reported for Puerto Rico 

by Kartesz (1998) in “A Synonymized 

Checklist of the Vascular Flora of the United 

States, Puerto Rico, and the Virgin Islands” 

(http://www.csdl.tamu.edu/FLORA/b98/ 

check98.htm). However, no specimen record 

exists (Kartesz, pers comm., 2001) to confirm 

this record.


7. FIMBRISTYLIS 

Fimbristylis Vahl, Enum. Pl. 2: 285. 1805, nom. conserv. 

Perennials or annuals, rhizomatous or tufted with fibrous roots. Culms cylindrical, or subcompressed 

to flattened, finely to coarsely ribbed, smooth, glabrous. Leaves basal; sheaths glabrous or pubescent, 

closed at summit, ventrally splitting with age, the apex entire; ligule absent or present; blades flattened 

or inrolled, sometimes folded, narrowly linear to filiform, glabrous or pubescent, the margins smooth or 

scabrous. Inflorescence an open, simple or compound anthela with ascending rays, or sometimes a 

head-like cluster with congested spikelets; involucral bracts leaf-like, usually smaller than the leaves, 

rarely overtopping the inflorescence; primary and secondary rays (when present) unequal, slender, finely 

ribbed, cylindrical to subflattened, smooth or scabrous; spikelets ovoid, oblong-ovoid, subglobose, or 

lanceoloid, borne singly at ray tips or sometimes clustered, many-flowered; scales all fertile except for 

the lowermost one, spirally imbricate, occasionally somewhat distichous, ovate to oblong-ovate, obtuse 

to acute, glabrous or pubescent, 1- to 5-nerved on sides, dorsally with an often 3-nerved carina, the 

nerves converging at apex and forming a mucro. Flowers bisexual; hypogynous squamellae or bristles 

absent; stamens 1-3, the anthers oblong or linear, sometimes apiculate; styles 2- or 3-branched, the 

unbranched portion flattened and fimbriate on margins (2-branched style) or slender, 3-angled basally, 

and glabrous (3-branched style), disarticulating from the summit of the achene. Achene biconvex, 

lenticular, or trigonous, obovate, elliptic-obovate, or oblong, smooth, cancellate, reticulate, verrucose, 

or warty. A genus of approximately 300 species in warm temperate and tropical climates worldwide, 

with the center of diversity in southeastern Asia. 

TYPE: Fimbristylis dichotoma (L.) Vahl [as “dichotomum”] (≡ Scirpus dichotomus L.), typ. conserv. 

Reference: Kral, R. 1971. A treatment of Abildgaardia, Bulbostylis, and Fimbristylis (Cyperaceae) 

for North America. Sida 4: 57-227. 

Key to the species of Fimbristylis 

1. Style 3-branched, or 2-branched in F. cymosa, the unbranched portion slender, entire, often 3-angled 

at expanded base.................................................................................................................. 2 

2. Leaves with a ligule of short dense hairs at adaxial junction of sheath and blade; spikelets 4-7 

(-9) mm long .................................................................................................. 1. F. complanata 

2. Leaves eligulate at adaxial junction of sheath and blade; spikelets 1.5-4 (-5) mm long............. 3 

3. Style 2-branched (in the flora area); achene obovate or pyriform, dorsiventrally compressed, 

faintly reticulate, low-papillate, opaque, maturing blackish .......... 2. F. cymosa 

3. Style 3-branched; achene obovoid, trigonous with convex sides or subrounded, cancellate, 

verrucose or tuberculate, translucent, maturing pale brown to gray brown............ 4 

4. Leaf sheaths laterally compressed, dorsally keeled, slightly spongy-thickened; blades 

folded and essentially unfacial; spikelets globose or ovoid-globose, rounded at apex; 

mucro shorter than to equaling the obtuse, rounded or emarginate apex of 

scale...................................................................................................... 6. F. littoralis 

4. Leaf sheaths dorsally compressed and rounded; blades not spongy-thickened, flattened; 

spikelets ovoid, acute at apex; mucro prolonged and slightly excurved beyond the 

acute to acuminate apex of scale .................................. 7. F. quinquangularis 

1. Style 2-branched, the unbranched portion flattened, fimbriate on margins........................................ 5 

5. Spikelet scales with dense, silvery, short-appressed hairs distally ................... 4. F. ferruginea 

5. Spikelet scales glabrous........................................................................................................... 6 

6. Spikelets 1.7-2.5 mm wide; spikelet scales 2.1-3 × 1.8-2.2 mm; achenes coarsely and deeply 

striate-reticulate with (5-) 7-9 (-12) longitudinal rows of horizontally oriented, rectangular 

cells ...................................................................................... 3. F. dichotoma 

6. Spikelets 2.5-3.5 (-4) mm wide; spikelet scales 2.7-4.2 (-5.5) × 2.4-3.8 mm; achenes finely 

and shallowly reticulate or pitted with 15-37 longitudinal rows of isodiametric or nearly 

isodiametric cells................................................................................................... 7

308 


7. Basal sheaths pale brown to brown; spikelet scales with longitudinally striate-nerved 

surface, semi-glossy, yellowish brown, finely reddish lineolate; achenes finely reticulate 

with 22-37 longitudinal rows of nearly isodiametric cells.......................... 

........................................................................................................... 5. F. inaguensis 

7. Basal sheaths dark brown or purple-black; spikelet scales with smooth surface, glossy, 

dark brown to dark brown or red-brown, the nerves indistinct and often faint; achenes 

with 15-20 longitudinal rows of shallow to pitlike isodiametric cells 

.............................................................................................................. 8. F. spadicea 

1. Fimbristylis complanata (Retz.) Link, Hort. 

Berol. 1: 292. 1827; Scirpus complanatus 

Retz., Observ. Bot. 5: 14. 1789; Cyperus 

complanatus (Retz.) Willd., Sp. P1. 1: 270. 

1797; Trichelostylis complanata (Retz.) Nees 

in Wight, Contr. Bot. India 103. 1834; 

Fimbristylis autumnalis var. complanata 

(Retz.) Kük., Bot. Jahrb. Syst. 59: 50. 1924. 

Type: India. Koenig s.n. (holotype: C). 

Caespitose or rhizomatous perennial, 20-60 

(-100) cm tall; rhizome short, ascending. Culms 

erect, compressed-trigonous proximally, flattened 

distally, firm, flexuous, sometimes twisted, 

antrorsely scabrid on margins (at least distally), 

0.7-2 mm wide. Leaves numerous, subdistichous, 

ascending to spreading; sheaths short, the 

uppermost dorsally keeled, brown to stramineous; 

ligule a horizontal band of dense hairs, 0.3-0.5 mm 

wide; blades linear, flattened, 5-70 cm × (1-) 1.3- 

4 mm, adaxially antrorsely scabrous at base, 

smooth otherwise, finely veined abaxially, margins 

finely and closely scabrous, abruptly narrowed to 

a flattened, slightly curved, acute tip. Inflorescence 

a terminal, compound, open or contracted anthela 

with ascending rays, 1.5-9 cm diam.; involucral 

bracts 4-6, the lowermost one leaf-like, often 

exceeding the inflorescence, the uppermost linearsetaceous; 

primary rays 4-12, unequal; spikelets 

15-100 or more, ovoid-lanceoloid to lanceoloid, 

4-7 (-9) × 1-1.5 (-2) mm, acute, obtuse at base, 

solitary at ray tips; scales ovate to ovate-lanceolate, 

1.4-2.5 × 1-1.5 mm, dorsally acute, brown, reddish 

lineolate, carina 3-nerved, green, prolonged 

beyond the acute apex as a short, slightly excurved 

mucro. Stamens 2 or 3, the anthers 0.7-1 mm long, 

apiculate; style 3-branched, the unbranched 

portion smooth, trigonous-dilated at base. Achene 

trigonous with shallowly convex faces, obovoid, 

0.7-1 × 0.5-0.7 mm, apiculate, obtuse at base, 

tricostate, finely striolate-reticulate, tuberculate to 

nearly smooth, chalk-white when immature, graywhite 

or pale brown at maturity. 

General distribution: Widespread in 

subtropical and tropical climates worldwide, often 

weedy. 


Islands: Moist or wet open areas of pond and 

stream borders, forest margins, grassy areas, sandy 

savannas, marshes, lake shores, roadsides, and 

disturbed habitats. Añasco, Bayamón, Cabo Rojo, 

Caguas, Cayey, Cidra, Dorado, Fajardo, 

Mayagüez, Moca, Naguabo, Río Grande, San 

Juan, San Sebastián, Vega Baja, and Yauco; St. 

Croix. 


Bayamón: Finca Santa Ana, Hioram 822 (US). 

Cayey: Guavate, Liogier 9772 (US). Naguabo: 

Sierra de Naguabo, mouth of Río Icaco, Shafer 

3181 (US). Río Grande: Sierra de Luquillo, 

Sintenis 1411 (US). San Juan: Río Piedras, 

Underwood & Griggs 254 (US). Yabucoa: Sierra 

de Yabucoa, Sintenis 2595 (US). Yauco: Río Loco, 

Sargent 665 (US). ST. CROIX: reduced form 

growing in a flower pot, Proctor 44990, SJ. 

2. Fimbristylis cymosa R. Br. , Prodr. 1: 228. 

1810. Type: Australia. Collector unknown 

(holotype: BM), fide, Koyama in A.C. Smith, 

Fl. Vitiensis Nova 1: 243. 1979. 

Scirpus glomeratus Retz., Observ. Bot. 4: 11. 1786, 

non Linnaeus, 1753; Fimbristylis glomerata 

Urb., Symb. Antill. 2: 166. 1900, non Nees 

von Esenbeck, 1834. Type: Sri Lanka. Koenig 

s.n. (holotype: C). 

Fimbristylis spathacea Roth, Nov. Pl. Sp. 24. 

1821; Fimbristylis cymosa subsp. spathacea 

(Roth) T. Koyama, Micronesica 1: 83. 1964. 

Type: India. Heyne s.n. (holotype: B, 

destroyed). 

Fimbristylis sintenisii Boeck., Bot. Jahrb. Syst. 7: 

276. 1886. Type: Puerto Rico. Sintenis 96b 


Scirpus obtusifolius sensu Grisebach, Fl. Brit. W. 

I. 571. 1864, non Lamarck, 1791.


Fimbristylis obtusifolia sensu C. B. Clarke in 

Urban, Symb. Antill. 2: 82. 1900, non 

(Lamarck) Kunth, 1837. 


rhizome short, thick. Culms erect, rigid, 

compressed-trigonous, coarsely ribbed and 

channeled, green to stramineous, 0.5-2 (-3) mm 

wide, the stout, sheathing bases hard and stiff, with 

fibrous remnants of old leaf bases. Leaves 

numerous; sheaths short, rigid, prominently 

nerved, glabrous, pale green to dark brown, the 

membranous inner band with a concave orifice; 

ligule absent; blades stiff, flattened to broadly Ushaped 

or somewhat involute, spreading, 

frequently falcate, 3-30 (-50) cm × 0.8-3 mm, to 4 

mm wide basally, obscurely veined, green to strawcolored, 

glabrous, antrorsely scabrous on margins, 

the apex with an acute, slightly curved tip. 

Inflorescence a contracted to congested, 

sometimes open, simple or compound anthela, 1- 

5 (-8) cm diam.; involucral bracts 2-3, leaf-like 

but reduced, linear to subulate, shorter than the 

inflorescence or the lowest one sometimes 

exceeding the inflorescence; rays compressed, to 

4 cm long, or wanting; spikelets 10-150 or more, 

ovoid, 2.4-4 × 1.8-2.5 mm, acute, sub-rounded at 

base, solitary at ray tips; scales ovate, slightly 

keeled, 1.2-1.6 × 0.8-1.2 mm, green to brown, with 

broad, scarious margins, carina 3-nerved, narrow, 

light green, emucronate at the obtuse to emarginate 

apex, lateral nerves wanting or indistinct. Stamens 

2, the anthers 0.5-0.7 mm long, apiculate; styles 

2-branched (in the flora area) or 3-branched to 

below the middle, the unbranched portion 

glabrous, entire, 3-angled at expanded base. 

Achene unequally biconvex, turgid on both faces, 

obovoid, 0.7-0.9 × 0.5-0.7 mm, broadly rounded 

to truncate at apex, bluntly apiculate, cuneate at 

base, faintly reticulate, low-papillate, opaque, light 

brown, becoming dark brown to essentially black 

at maturity. 



Islands: Typically wet or dry coastal habitats, on 

beaches and in sandy savannas, marshy or swampy 

areas, swales, borders of mangroves, scrub forest, 

thickets, roadsides, and disturbed areas. Occurring 

sparingly inland on serpentine or limestone 

substrate. Arecibo, Arroyo, Bayamón, Cabo Rojo, 

Camuy, Carolina, Cataño, Cayey, Cayo Icacos, 

Cayo Lobos, Cayo Ramos, Dorado, Fajardo, 

Guánica, Guayama, Guayanilla, Hatillo, 

Humacao, Isabela, Isleta Marina, Juana Díaz, 

Juyuya, Lajas, Loíza, Manatí, Maricao, Mayagüez, 

Moca, Mona Island, Naguabo, Ponce, Rincón, Río 

Grande, Salinas, San Juan, Vega Alta, Vega Baja, 

Vieques, and Yabucoa; Anegada, St. Croix, St. 

John, St. Thomas, and Tortola. 


Arroyo: Playa Guilarte, González-Más 1311 (US). 

Bayamón: Bo. Guaraguao Abajo, Road 174, km 

6.9, Proctor & Foy 42329 (US). Cabo Rojo: 

Boquerón, González-Más 931 (US). Carolina: 

Boca de Cangrejos, González-Más 1016 (US). 

Cataño: Bo. Palmas, Axelrod & Díaz 12610A 

(UPRRP). Guánica: Sargent 64 (US). Guayama: 

Rd. 707, González-Más 671 (US). Guayanilla: Rd. 

336; Bahia, González-Más 819 (US). Juana Díaz: 

Pastillo, González-Más 752 (US). Manatí: Bo. 

Nuevas Tierras Saliente, Axelrod & Thomas 11324 

(UPRRP). Mayagüez: N.L. Britton 2363 (US). 

Mona Island: Camino del Diablo, Acevedo-Rdgz. 

et al. 4277 (FTG, MO, NY, SJ, US). Ponce: Bo. 

Cañas, along dirt road to Punta Cucharas, Axelrod 

& Chávez 7329 (US). San Juan: Las Marias, near 

Santurce, Hioram 810 (US). Vega Baja: 

Tortuguero, Rd. 687, González-Más 1035 (US). 

Vieques Island: Puerto Ferro Peninsula, Fosberg 

57498 (US). ST. CROIX: Caledonia Gut, Fosberg 

56790 (US). ST. JOHN: Coral Bay Quarter; trail to 

Fortsberg, Acevedo-Rdgz. 4087 (JBSD, MO, NY, 

UPR, US, VINPS). 

3. Fimbristylis dichotoma (L.) Vahl, Enum. Pl. 

2: 287. 1805; Scirpus dichotomus L., Sp. Pl. 

50. 1753. Lectotype: Sri Lanka. Herbarium 

Hermann 40, v. 2, 63, No. 40 (BM), designated 

by T. Koyama in A.C. Smith (ed.), Fl. Vit. 

Nova 1: 244. 1979. 

Scirpus annuus All., Fl. Pedem. 2: 277. 1785; 

Fimbristylis annua (All.) Roem. & Schult., 

Syst. Veg. 2: 95. 1817. Type: Italy. Collector 

unknown (holotype: TO). 

Scirpus diphyllus Retz., Observ. Bot. 5: 15. 1789; 

Fimbristylis diphylla (Retz.) Vahl, Enum. Pl. 

2: 289. 1805; Fimbristylis annua var. diphylla 

(Retz.) Kük., Repert. Spec. Nov. Regni Veg. 

23: 196. 1926. Type: India. Koenig s.n. 

(holotype: C). 

Scirpus villosus Poir. in Lamarck, Encycl. 6: 781. 

1804; Fimbristylis villosa (Poir.) Roem. & 

Schult., Syst. Veg. 2. 98. 1817. Type: Puerto

310 


Rico. Ledrú s.n. (holotype: probably B-Willd. 

1264 or at P). 

Fig. 50. F-J 

Tufted perennial, (4-) 7-80 (-110) cm tall; 

rhizome short, ascending, thickened and nodose. 

Culms compressed-trigonous, coarsely ribbed and 

channeled, green, glabrous, 0.5-2 mm wide. 

Leaves 5-7 (-10) per culm; sheaths ligulate, finely 

nerved, glabrous or sometimes pilose, light reddish 

brown, the inner band pilose distally, ciliate along 

margins; ligule a horizontal band of dense 

appressed hairs at adaxial base of blade, ca. 0.5 

mm wide; blades flattened to somewhat involute, 

3-40 (-70) cm × 0.8-4 mm, to 5 mm wide basally, 

distinctly veined on both sides, glaucous, glabrous 

or pilose, antrorsely scabrous and sometimes 

ciliate on margins, the apex with a rounded curved 

tip. Inflorescence an open to contracted, simple to 

compound anthela with ascending rays, 1.5-10 

(-15) cm diam.; involucral bracts 4-6, the 

lowermost ones, ciliate and pubescent proximally, 

to 10 cm long, usually equaling to overtopping 

the inflorescence, the upper ones linear to filiform; 

primary rays remotely scabrous on margins, to 7 

(-11) cm long, secondary rays shorter; spikelets 

(3-) 10-60, ovoid to ovoid-lanceoloid or oblonglanceoloid, 

4-8 (-15) × 1.7-2.5 mm, acute, subrounded 

at base, solitary at ray tips, sometimes 

fasciculate; scales ovate to broadly ovate or 

oblong-ovate, 2.1-3 × 1.8-2.2 mm, glabrous, shiny, 

pale brown to reddish brown, usually dark brown 

distally, carina 3-nerved, pale green, prolonged 

beyond the acute to obtuse apex as a mucro, lateral 

nerves 3-4, indistinct. Stamens 1 or 2, the anthers 

0.6-1 mm long, apiculate; styles 2-branched, the 

unbranched portion flattened, ca. 0.5 mm wide, 

fimbriate along margins. Achene biconvex, 

obovate, 1-1.3 × 0.8-1 mm, rounded and apiculate 

at apex, short-stipitate at base, whitish to 

stramineous or brownish, coarsely and deeply 

striate-reticulate with (5-) 7-9 (-12) longitudinal 

rows of horizontally oriented rectangular cells on 

each side, the longitudinal ribs between the cells 

prominent, smooth or sometimes sparsely 

tuberculate. 

General distribution: Cosmopolitan, 

primarily in warm-temperate and tropical regions. 


Islands: Damp often sandy soils in a wide variety 

of successional habitats including sandy savannas, 

forest edges, river banks, streams, ditches, 

agricultural fields, roadsides, grassy areas, lawns, 

waste places, and disturbed areas. Adjuntas, 

Aguada, Aguas Buenas, Aibonito, Añasco, 

Arecibo, Bayamón, Cabo Rojo, Caguas, Carolina, 

Cataño, Cayey, Ceiba, Coamo, Comerío, Corozal, 

Dorado, Fajardo, Guayama, Guaynabo, Gurabo, 

Humacao, Isabela, Jayuya, Juncos, Lares, Las 

Marías, Las Piedras, Loíza, Luquillo, Maricao, 

Maunabo, Mayagüez, Moca, Naguabo, Naranjito, 

Orocovis, Patillas, Peñuelas, Ponce, Quebradillas, 

Río Grande, Salinas, San Germán, San Juan, San 

Lorenzo, San Sebastían, Trujillo Alto, Utuado, 

Vega Alta, Vega Baja, Vieques, Villalba, Yabucoa, 

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


Common name: Puerto Rico: Junquito. 


Adjuntas: Rd. 10, km 32, González-Más 877 (US). 

Aguas Buenas: Bo. Sumidero, González-Más 1177 

(US). Aibonito: Liogier et al. 31445 (US). Añasco: 

Heller 4531 (US). Arecibo: Sargent 680 (US). 

Bayamón: Heller & Heller 411 (US). Cabo Rojo: 

Boquerón, Rd. 4, km 2.4, González-Más 938 (US). 

Caribbean National Forest, along hwy. 988, Boom 

7064 (US). Cataño: Bo. Palo Seco, Axelrod & Díaz 

12709 (UPRRP). Guayama: Puerto Jobos, Rd. 707, 

González-Más 653 (US). Guaynabo: P.R. Tropical 

Island Sonadora, Woodbury et al. s.n. (US). 

Gurabo: Substation Gurabo, Woodbury et al. s.n. 

(US). Humacao: km 27.1, González-Más 72 (US). 

Juncos: Sintenis 2505 (US). Maricao: Bo. Indiera 

Fría, Along track near Río Lajas below El Salto 

de Curet, Proctor & Padrón 45678 (US). 

Mayagüez: Sintenis 96 (US). Moca: Sargent 234 

(US). Naguabo: Rd. 3; Daguao River, González- 

Más 227 (US). Patillas: Rd. 3, km 144, Patillas to 

Maunabo, González-Más 1334 (US). Ponce: Road 

from Ponce to Adjuntas, Underwood & Griggs 757 

(US); Reserva Forestal de Toro Negro, Stimson 

1514 (US). Río Grande: Along Rt. 186 bordering 

Caribbean National Forest, just N of the El Verde 

Field Station, Strong et al. 392. (GMUF); Luquillo 

Forest, Top of El Yunque Mt., González-Más 1534 


San Lorenzo: Sierra de Cayey, Carite Forest 

Reserve, Proctor & Milety 43964 (US). San 

Sebastián: Bo. Cidral, González-Más 925 (US). 

Utuado: Road from Utuado to Lares, Underwood 

& Griggs 68 (US). Vega Baja: Tortuguero, Rd. 687, 

González-Más 1045 (US). Vieques Island: Vicinity 

of Isabel, Shafer 2502 (US). Yauco: Río Lares, 

Sargent 665 (US). ST. JOHN: Bethamia to


Rosenberg, N.L. Britton & Shafer 277 (US). ST. 

THOMAS: Crown, N.L. Britton & Marble 1345 

(US); Signalhill, Eggers s.n. (NA). TORTOLA: Road 


(US). 

4. Fimbristylis ferruginea (L.) Vahl, Enum. Pl. 

2: 291. 1805; Scirpus ferrugineus L., Sp. Pl. 

50. 1753. Lectotype: Jamaica. Herb. A. van 

Royen, sheet no. 902.77-420 (L), designated 

by C. D. Adams in Cafferty & C. E. Jarvis 

(ed.), Taxon 53: 180. 2004. 

Scirpus debilis Lam., Tabl. Encycl. 1: 141. 1791; 

Scirpus ferrugineus var. debilis (Lam.) Poir. 

in Lamarck, Encycl. 6: 780. 1804. Type: South 

America. Richard s.n. (holotype: probably at 

P). 

Fimbristylis ferruginea var. compacta Kük., 

Repert. Spec. Nov. Regni Veg. 23: 196. 1926. 

Type: Cuba; Pinar del Río. Ekman 17829 



tall; rhizomes short, stout. Culms compressedtrigonous 

proximally, compressed to subflattened 

distally, 0.7-3 mm wide, to 4 mm wide at base, 

coarsely ribbed, glabrous. Leaves 3-6 per culm; 

sheaths ligulate, finely and prominently nerved on 

back, pale green, glabrous, the inner band 

ferrugineous; ligule a horizontal band of pale, 

appressed hairs at adaxial base of blade, 0.3-0.4 

mm wide; blades involute, 1-25 cm × 0.7-1.5 (-2) 

mm, light green to brown, glabrous, antrorsely 

scabrous on margins, abaxial surface distinctly 

nerved, the apex acute, often with a slightly curved 

tip. Inflorescence a simple or rarely compound, 

dense or sometimes somewhat head-like anthela 

with ascending rays, 1-4 (-8) cm diam.; involucral 

bracts (2-) 3-6, leaf-like, but reduced, flattened to 

somewhat involute, to 7 cm long, 0.8-1.4 mm 

wide, shorter the inflorescence or the lowermost 

one exceeding it; rays compressed, short, finely 

ribbed, the primary ones to 5 cm long; spikelets 

(1-) 3-14, ovoid to narrowly ovoid, 6-12 × 3-5 mm, 

solitary at ray tips; scales widely ovate to roundedovate, 

or oblong-ovate, boat-shaped, dorsally 

obtuse, 3-4 × 2.8-4 mm, pale reddish brown, with 

dense, silvery, short-appressed hairs and ciliate 

margins distally, lateral nerves indistinct, carina 

1-nerved, green or grayish green, prolonged 

beyond the obtuse to rounded apex as a short, 

thickened mucro. Stamens 3, the anthers 0.9-1.3 

mm long, bluntly apiculate at apex; styles 2branched, 

the unbranched portion flattened, 0.4- 

0.5 mm wide, fimbriate on margins. Achene 

biconvex, obovate, 1.4-1.7 × 1-1.2 mm, obtuse to 

truncate at apex, bluntly apiculate, cuneate at base, 

short-stipitate, straw-colored to pale brown, finely 

and shallowly reticulate with 25-35 longitudinal 

rows of isodiametric cells. 



Islands: Wet, brackish or saline coastal habitats, 

in marshes, estuaries, borders of mangroves, 

swampy areas, swales, pasture, and drainage 

canals. Aguadilla, Añasco, Arecibo, Arroyo, 

Bayamón, Cabo Rojo, Cataño, Cayo Icacos, Cayo 

Ramos, Coamo, Culebra, Fajardo, Guánica, 

Guayama, Humacao, Isabela, Isleta Marina, 

Jayuya, Loíza, Luquillo, Mayagüez, Ponce, 

Quebradillas, Río Grande, Salinas, San Juan, Santa 

Isabel, Toa Baja, Vega Baja, and Vieques; 

Anegada, St. Croix, St. John, St. Thomas, Tortola, 



Añasco: Sargent 633 (US). Arecibo: Bo. 

Garrochales, Cienaga Tiburones, Proctor et al. 

46139 (US). Bayamón: Sintenis 1234 (US). Cabo 

Rojo: Joyuda Road 102, González-Más 613 (US). 

Cataño: Punta Salinas, Liogier et al. 32494 (NY). 

Cayo Ramos: Woodbury & Martorell 1 (US). 

Coamo: Sintenis 3318 (US). Culebra Island: N.L. 

Britton & Wheeler 250 (US). Fajardo: Bo. 

Quebrada Vueltas, between Rt. 3 and the sea, 

Axelrod & L. Pérez 9074 (HAC). Guánica: Sargent 

192 (US). Guayama: Bo. Jobos, area S of oil 

refinery and E of port, Axelrod & Pérez 8640 (US). 

Humacao: Sargent 3004 (US). Mayagüez, Heller 

4367 (US). Ponce: Bo. Cañas, along dirt road to 

Punta Cucharas, Axelrod & Chávez 7328 (US). 

Salinas: Rd. 3, km 156, González-Más 1295 (US). 

San Juan: Stone quarry 2 mi. E of Santurce, Heller 

& Heller 821 (US). Toa Baja: Bo. Sabana Seca, 

area behind Naval Station, N of Rt. 867, Axelrod 

et al. 10214 (NY, US). Vega Baja: Bo. Cabo 

Caribe, between Los Naranjos & Caño Las Pozas, 

Proctor et al. 45595 (US). ST. CROIX: Midland 

road, J.J. Ricksecker 244 (US). ST. THOMAS: Long 

Cay, Eggers s.n. (NA, US). 

5. Fimbristylis inaguensis Britton, Torreya 13: 

216. 1913. Type: Bahamas; Inagua. Nash & 

Taylor 1019 (holotype: NY!).

312 



rhizome short, knotty, often with old bulbous culm 

bases bearing fibrous remnants of leaf sheath 

bases. Culms solitary or 2-3 together, erect, rigid, 

subterete to terete proximally, subflattened distally, 

smooth, 1-2 mm wide. Leaves 7-13 per culm, 

spreading and often subdistichous; sheaths rigid, 

ligulate, often appressed-puberulent, pale brown 

to brown; ligule a band of pale or whitish coarse 

hairs, ca. 0.5 mm long, extending out along upper 

margin of the deeply V-shaped orifice; blades 

linear, stiffened, 10-60 cm × 0.8-3 mm, flattened 

or subinvolute, finely veined and green abaxially, 

smooth and whitish stramineous adaxially, 

appressed-puberulent basally above ligule, smooth 

to finely and densely antrorsely scabrous on 

margins (at least distally), glabrous, the apex 

abruptly narrowed to a flattened, acute or subrounded 

tip. Inflorescence an open or contracted, 

simple or compound, anthela with ascending rays, 

4-8 cm diam.; spikelets 3-30, ovoid to ovoidlanceoloid 

or ellipsoid, 6-13 (-15) × 2.5-3.3 mm, 

acute, sub-rounded at base, solitary at ray tips; 

scales ovate to widely ovate or oblong-ovate, 2.7- 

3.5 × 2.4-2.8 mm, dorsally obtuse to rounded, 

closely and often distinctly 7- to 9-nerved on each 

side above the narrowly scarious, often distally 

ciliate margins, yellowish brown, finely reddish 

lineolate, dull to semi-glossy, carina 3-nerved, 

included or slightly prolonged beyond the acute 

to obtuse apex as a short mucro. Stamens 3, the 

anthers 1.2-1.6 mm long, with a short, papillate 

apiculum; style 2-branched, flattened, the margins 

of the unbranched portion densely fimbriate. 

Achene unequally biconvex, obovate to broadly 

so or obpyriform, broadly rounded or truncate at 

apex, short-cuneate at base, 1.1-1.4 × 0.8-1.2 mm, 

pale to dark brown, finely reticulate with 22-37 

longitudinal rows of horizontally oriented, 

rectangular to nearly isodiametric cells. 

General distribution: Endemic to the 

Bahamas and Greater Antilles. 

Distribution in the Virgin Islands: Coastal 

sands, in openings of strand vegetation along 

beaches. Only known in the flora area from 

Anegada. 

Selected specimens examined: ANEGADA: West 

End, N.L. Britton & Fishlock 966 (US); near West 

End, D’Arcy 4811 (US). 

6. Fimbristylis littoralis Gaudich. in Freycinet, 

Voy. Uranie 413. 1829. Type: Marianas, 

Molucca, or Timor Islands. Collector 

unknown (holotype: probably at P). 

Scirpus miliaceus L., Syst. Nat. ed. 10, 2: 868. 

1759, nom. reject. (see note below); 

Fimbristylis miliacea (L.) Vahl, Enum. Pl. 2: 

287. 1805. 

Densely tufted annual or short-lived 

perennial, 7-75 (-100) cm tall. Culms slender, erect 

to ascending, flexuous, firm distally, cross-shaped 

to somewhat 4-angled proximally or subflattened, 

0.5-2 mm wide, to 3 mm wide at base, light green, 

glabrous, smooth or sparsely scabrous just below 

inflorescence, clothed with 2-3 bladeless sheaths 

at base. Leaves primarily basal, distichous, 

numerous, spreading and often fan-like; sheaths 

laterally flattened, 3-5 mm wide, soft and spongy, 

dorsally keeled, stramineous or pale green, 

glabrous, tapering gradually to blades; ligule 

absent; blades narrowly linear, soft, laterally 

compressed and essentially unifacial, 10-30 (-60) 

cm × 0.7-3 (-4) mm, glabrous, smooth proximally, 

spinulose-scabrous on margins distally, tapering 

evenly from base to attenuate, subacute, slightly 

dialated, pale tip. Inflorescence a compound or 

decompound anthela with ascending rays, 2-12 × 

1.3-6 (-9) cm, bearing numerous spikelets; primary 

involucral bracts 2-6, setaceous, scabrous on 

margins, shorter than the primary rays 3-7, 

unequal, ascending, uppermost often divergent, 1- 

6 cm long, scabrous; spikelets solitary at ray tips, 

globose to globose-ovoid, 1.5-3 (-5) × 1.5-3 mm, 

with both ends rounded; scales ovate, 1-1.3 × 0.7- 

1 mm, cucullate at maturity, membranaceous, 

broadly white hyaline on the margins, carina 3nerved, 

green or yellowish, rarely prolonged 

beyond the obtuse, rounded or emarginate apex. 

Stamens 1-2, the anthers 0.5-0.8 mm long, 

apiculate; style 3-branched, the unbranched 

portion slender, smooth or with some fimbriolae 

distally, pyramidal at base. Achene trigonous with 

convex sides, obovoid, 0.6-0.8 × 0.3-0.5 mm, 

rounded to an apiculate apex, cuneate to base, 

conspicuously cancellate with 5-7 longitudinal 

rows of fine, transversely oriented linear-oblong 

cells on each side, the longitudinal ribs frequently 

prominent and sparsely verruculose, translucent, 

pale brown to gray-brown.



Mexico, Central America, West Indies, South 

America, Africa, Southeast Asia, Malesia, 

Australia, and Pacific islands and atolls, often 

weedy. 

Distribution in Puerto Rico: In wet or damp 

sandy soils of river banks, marshy areas, swales, 

ditches, agricultural fields, pasture, roadsides, and 

disturbed areas. Caguas, Carolina, Cayey, 

Humacao, Naguabo, Patillas, San Juan, and 

Yabucoa. 

Note: The name Fimbristylis miliacea has 

been used in two different senses based on two 

specimens of type material side by side in the 

Linnaean herbarium (LINN) which are different 

taxa. The first sheet, 71:40 is the species with ovoid 

spikelets, otherwise known as Fimbristylis 

quinquangularis and the second, 71:41, is the 

species with globose spikelets circumscribed by 

Vahl as Fimbristylis miliacea. Rejection of the 

name Scirpus miliaceus (Fimbristylis miliacea) 

was necessary to stabilize the nomenclature and 

eliminate the name as a source of confusion (see 

Strong, Taxon 53: 1069-1070. 2004). 


Caguas: Bo. Cañabón, along W side of Río 

Cagüitas just S of Rd. 156, Proctor 46150 (US). 

Carolina: Sabana Abajo, P.R. 26, González-Más 

1238 (MAPR, NY, US). Cayey/Patillas: Rte. 179, 

ca. 2-3 mi. N of rte 742, Axelrod & Taylor 9574 

(UPRRP). Humacao: Entrada al pueblo, Carr. 3, 

km 68.8-9, Montaño & González-Más 100 

(MAPR). Naguabo: relict coconut plantation, ca. 

0.8 km ESE of highway 3 bridge at mouth, Proctor 

& Torres 50152 (SJ). San Juan: Bo. Hato Rey 

Norte, just S of Caño de Martín Peña, Proctor & 

Cintrón 43106 (SJ). Yabucoa: Sintenis 4947 (NY, 

US). 

7. Fimbristylis quinquangularis (Vahl) Kunth, 

Enum. Pl. 2: 229. 1837; Scirpus 

quinquangularis Vahl, Enum. Pl. 2: 279. 1805; 

Trichelostylis quinquangularis (Vahl) Nees in 

Wight, Contr. Bot. India 104. 1834. Type: 

India. Koenig s.n. (holotype: C-Vahl). 

Caespitose annual or short-lived perennial, 

(10-) 20-60 (-90) cm tall. Culms erect, slender, 

flexuous, cross-shaped or subflattened, antrorsely 

scabrous on margins just below inflorescence, 0.7- 

2 mm wide. Leaves 3-5 per culm, primarily basal, 

spirally arranged, the lowermost ones bladeless 

or essentially so; sheaths loose, thinly herbaceous, 

dilated at base, gradually narrowing to blade; 

ligule absent; blades linear, 3-40 cm × 1.3-3 (-4) 

mm, flattened to conduplicately folded, margins 

antrorsely scabrous (at least distally), blade and 

sheath surface adaxially scabrous proximally, 

otherwise glabrous, acuminate to a flattened, 

slightly curved tip. Inflorescence a compound to 

decompound anthela with ascending rays, 2.5-13 

× 2-7 cm; involucral bracts 4-9, linear to linearsetaceous, 

shorter than the inflorescence; primary 

rays 7-16, unequal, several often reduced and 

unispiculate; spikelets (12-) 20-170 (-230), ovoid 

to ovoid-lanceoloid, 2.5-4 (-6) × 1.2-1.8 mm, 

acute, obtuse to sub-rounded at base; fertile scales 

ovate to broadly ovate, 1.1-1.5 × 0.8-1.2 mm, 

thinly herbaceous, dorsally acute to obtuse, 

curvate-keeled, brown to red-brown, glabrous, 

margins entire, broadly scarious, carina 3-nerved, 

greenish, prolonged beyond the acute to obtuse 

apex as a short slightly excurved mucro. Stamens 

1-2, the anthers 0.5-0.6 mm long, apiculate; style 

3-branched, the unbranched portion slender, 

trigonous, smooth. Achene obtusely trigonous to 

subterete, obovoid or pyriform, 0.6-0.8 × 0.4-0.6 

mm, obscurely 3-costate, rounded at apex, 

apiculate, acute at base, short-stipitate, pale, finely 

cancellate-tuberculate with 3-6 vertical rows of 

horizontally oriented linear cells on each side. 

General distribution: West Indies, South 

America, and Old World tropics, often weedy. 


sandy soils of ditches, river banks, marshy areas, 

swales, agricultural fields, pasture, roadsides, and 

disturbed areas. Aguada, Añasco, Carolina, Ceiba, 

Dorado, Gurabo, Humacao, Luquillo, Maunabo, 

Mayagüez, Naguabo, Río Grande, and Toa Baja. 


Aguada: Sargent 561 (US). Carolina: 65 Inf. road, 

Río Grande Loíza banks, González-Mas 1230 

(US). Ceiba: Bo. Guayacán, Roosevelt Roads 

Naval Reservation, Axelrod et al. 10863 (US). 

Humacao: Bo. Punta Santiago, beside Hwy 3 at 

Río Anton Ruiz bridge, Proctor 44203 (US). 

Luquillo: Bo. Mata de Platano, in back of old 

coconut plantation, Axelrod & Axelrod 6544(US). 

Mayagüez: College campus, González-Mas 2197 

(US). Naguabo: Road 13, km 4.1, González-Más 

1366 (US).

314 


8. Fimbristylis spadicea (L.) Vahl, Enum. Pl. 2: 

294. 1805. Scirpus spadiceus L., Sp. Pl. 51. 

1753. Lectotype: Jamaica. Sloane, Voy. 

Jamaica 1: t. 76, f. 2. 1707, designated by 

McVaugh, Fl. Novo-Galiciana 13: 380. 1993. 

Fimbristylis castanea of authors, non (Michaux) 

Vahl, 1805. 

Coarse, caespitose perennial, 60-200 cm tall; 

rhizome ascending or horizontal, ligneous. Culms 

erect, slender, compressed or subterete proximally, 

half-terete or obtusely trigonous distally, stiff, 

glabrous, 1.3-3 (-4) mm wide. Leaves numerous, 

ascending to erect; sheaths coriaceous, dark brown 

or purple-black proximally, smooth, the inner band 

reddish brown with a U-shaped orifice; ligule 

present on basal sheaths, a horizontal band of hairs, 

ca. 0.3 mm wide, often extending out along upper 

margins of sheath orifice, essentially absent on 

uppermost sheaths; blades narrowly linear, 

substiffened, crescentform to strongly involute, 

10-100 cm × 0.5-1.5 (-2) mm, distinctly veined 

abaxially, smooth adaxially, light green to 

stramineous, antrorsely scabrous on margins, the 

apex with an acute, slightly curved tip. 

Inflorescence a terminal, open or contracted, 

compound anthela with ascending rays; involucral 

bracts 3-5, the lowermost one leaf-like, stiffly 

erect, equaling or exceeding the inflorescence; 

primary rays to 9 (-12) cm long; spikelets 10-60, 

ovoid-lanceoloid, lanceoloid, or narrowly 

ellipsoid, cylindrical, 6-17 (-20) × 2.7-3.5 (-4) mm, 

acute, short-cuneate at base, solitary at ray tips; 

scales broadly ovate, 2.8-4.2 (-5.5) × 2.5-3.8 mm, 

dorsally rounded, closely and faintly 7- to 9-nerved 

on each side above the narrowly scarious, often 

distally erose margins, smooth, dark brown to dark 

8. FUIRENA 

Fuirena Rottb., Descr. Icon. Rar. Pl. 70, t.19. 1773. 

brown or red-brown, semi-glossy, carina 3-nerved, 

indistinct, pale, shortly prolonged beyond the 

broadly acute to obtuse apex as a short mucro. 

Stamens 2-3, the anthers 1.5-2 mm long, with a 

subulate, prickly apiculum; style 2-branched, the 

unbranched portion strongly flattened and 

fimbriate. Achene lenticular, obovoid or pyriform, 

1.4-1.9 × 0.8-1.1 mm, rounded to subtruncate at 

apex, cuneate at base, finely cancellate, with 15- 

20 longitudinal rows of shallow to pitlike 

isodiametric cells, pale to deep brown. 




Islands: Wet, sandy soils of coastal areas, in salt 

marsh, borders of mangroves, brackish marshland, 

swales, canals, and salt ponds. Arecibo, Arroyo, 

Cabo Rojo, Cataño, Guánica, Guayama, 

Guayanilla, Peñuelas, Ponce, Salinas, San Juan, 

Santa Isabel, and Yauco; St. Croix and St. Thomas. 


Island: Vegetation and Flora”, Atoll. Res. Bull. No. 

188: 15. 1975, based on D’Arcy 4811 and 4836 

(both at MO). 


Arroyo: Guilarte Beach, González-Más 1325 (US). 

Cabo Rojo: Bo. Pedernales, ca. 0.5 km E of Laguna 

Guanaquilla, Proctor & McKenzie 43924 (US). 

Cataño: Palo Seco, Rd. 165, González-Más 1166 

(US). Guánica: Sintenis 3871 (US). Guayama: 

Puerto Jobos, Rd. 707, González-Más 691 (US). 

Guayanilla: Sargent 161 (US). Peñuelas: Rd. 2, 

km 10.9, González-Más 809 (US). Ponce: Sintenis 

4875 (US). Salinas: Rd. 705, km 1, González-Más 

722 (US). Santa Isabel: Rd. 1, km 110.5, González- 

Más 755 (US). ST. CROIX: Limetree Bay, J.J. 


Annual or perennial herbs, often pubescent; rhizomes of perennial species horizontal, scaly, producing 

roundish cormous offshoots in some species. Culms trigonous or terete, rarely pentagonous, coarsely or 

finely ribbed, green. Leaves basal and cauline, the lowermost often reduced to short-bladed or bladeless 

sheaths; sheaths well-developed, finely veined, pubescent; ligule a flange of membranous tissue, ciliate 

on margin with brownish hairs; blades linear, finely veined, often pubescent abaxially, smooth or 

sometimes pubescent adaxially, green. Inflorescence paniculate, composed of a terminal and series of 

lateral partial panicles with 2-many digitate, subdigitate or corymbose clusters of spikelets from the 

upper leaf-like bracts, or a single terminal head or cluster of spikelets; peduncles and branches pubescent;


spikelets ovoid to ovoid-lanceoloid or ovoid-ellipsoid, many-flowered, with subloosely spirally imbricate 

scales, the lowermost 1-3 scales occasionally empty; scales obovate to oblong-obovate, dorsally obtuse 

to rounded, pubescent or sometimes glabrous, with broad scarious margins, often reddish brown or 

grayish, the carina often prolonged at apex as a short or elongate, excurved mucro. Flowers bisexual; 

hypogynous bristles and/or bladed bristles 3 or 6, rudimentary, or wanting, when present, in one or two 

whorls, the outer whorl of 3 retrorsely barbed or smooth simple bristles, the inner whorl of 3 bladed 

bristles, rarely simple ones; stamens 1-3; style 3-branched. Achene trigonous with prominent angles, 

broadly ellipsoid or obovoid, slender-beaked at apex, often stipitate at base, the surface smooth, finely 

striated, or rarely tuberculate or cancellate. Approximately 30 species in warm temperate and tropical 

regions with the centers of distribution in Africa and the Americas. 

TYPE: Fuirena umbellata Rottb. 

References: Kral, R. 1978. A synopsis of Fuirena (Cyperaceae) for the Americas north of South 

America. Sida 7(4): 309-354. Muasya, A.M. 1998. A synopsis of Fuirena (Cyperaceae) for the flora of 

tropical East Africa. Kew Bull. 53(1): 187-202. 

Key to the species of Fuirena 

1. Leaf margins with ascending to appressed, stiff trichomes (at least proximally); spikelets 1.8-2.8 mm 

wide; achene bearing a whorl of 3 subulate bristles and a whorl of 3 bladed bristles; blades of bladed 

bristles narrowly oblong, thickened, obscurely 3-nerved, slightly incurved-apiculate or mucronate 

...................................................................................................................... 1. F. robusta 

1. Leaf margins glabrous; spikelets 1.3-2.2 mm wide; achene bearing 3 bladed bristles only; blades of 

bladed bristles broad, oblong, obovate, or turbinate, thinly herbaceous, conspicuously 3- to 5-nerved, 

incurved-awned, the awn often coiled apically ........................................... 2. F. umbellata 

1. Fuirena robusta Kunth, Enum. Pl. 2: 185. 1837. 

Type: Brazil. Sellow s.n. (holotype: B, 

destroyed). 

Coarse, rhizomatous perennial, 80-170 (-225) 

cm tall; rhizomes short, horizontal, stout, 7-10 mm 

thick, producing cormous offshoots. Culms often 

rooting from lowermost nodes, erect, 

subcaespitose or crowded in a row, pentaquetrous, 

subspongy and easily compressed proximally, firm 

distally, glabrous, with conspicuous pale cells, 2.3- 

12 mm wide, to 18 mm wide (flattened) at base. 

Leaves 7-16 below inflorescence; sheaths 

elongate, subinflated, smooth, green or the 

lowermost brownish; ligule a thin, membranous, 

light brown, short-ciliate flange of tissue, 1-3 mm 

long; blades short proximally, elongate midculm 

and distally, lanceolate to linear-lanceolate or 

linear-oblong, 2-25 (-35) cm × 7-23 (-30) mm, 

essentially smooth except for hispid veins on both 

sides distally, margins with ascending to 

appressed, stiff trichomes (at least proximally), 

abruptly narrowed at apex and inrolled to a 

narrowly acute to acuminate tip. Inflorescence a 

terminal and series of 1-3, subcontiguous to 

remote, simple or compound, lateral partial 

corymbs from the upper leaf-like bracts; corymbs 

sublax, 1.5-7 cm diam., the spikelets in short, often 

lobate, fasciculate clusters at branch tips; spikelets 

ovoid to ovoid-lanceoloid, 5-8 (-10) × 1.8-2.8 mm, 

to 4 mm wide in fruit, acute to narrowly so, subrounded 

at base; scales ovate-elliptic, 2.5-3.3 × 

1.7-2.5 mm, puberulent, reddish brown or greenish 

brown, finely cellular-striate, obscurely 2- to 4nerved 

on sides, often coarsely beset distally, and 

along carina, with coarse tubercle-based 

translucent hairs, ciliate on margins, carina 3- to 

rarely 5-nerved, prolonged beyond the rounded or 

retuse apex as a narrowly lanceolate, antrorsely 

scabrous, erect awn, 0.5-2 mm long. Hypogynous 

segments 6, represented by a whorl of 3 subulate, 

antrorsely barbed to smoothish bristles which are 

as long as achene or slightly longer and a whorl of 

3 bladed bristles which have ovate to narrowly 

ovate or ovate-elliptic clawed blades, 1-1.5 mm 

long, thickened, minutely cellular roughenedreticulate, 

obscurely 3-nerved, ciliate on margins 

distally, the acute apex with a slightly incurved 

scabridulous awn-like appendage. Stamens 3, the 

anthers 1-2 mm long, apiculate. Achene rhomboid 

with flat to slightly convex sides, 1-1.7 (including 

beak) × 0.6-0.9 mm, acute at apex with a trigonous-

316 


subulate beak, abruptly attenuate to a stipitate base, 

pale brown to brown with wire-like angles, 

smooth, lustrous, the beak antrorsely barbed 

distally. 

General distribution: Mexico, Panama, Cuba, 

Puerto Rico, and South America. 

Distribution in Puerto Rico: Low, wet areas 

in marshy or swampy habitats. Known only from 

the Municipality of Florida from a collection made 

by Woodbury s.n. on 19 May 1968 (UPR; photo at 

US). 

Note: Previously cited for Puerto Rico by 

Urban (1903) based on Heller 247. However, the 

US and NY duplicates of that collection are F. 

umbellata Rottb. 


Florida: Woodbury s.n. (UPR; photo at US). 

2. Fuirena umbellata Rottb., Descr. Icon. Rar. 

Pl. 70. 1773. Type: Surinam. Rolander s.n. 

(holotype: C-Rottb.). 

Fig. 51. A-E 


rhizomes 4-6 mm thick, producing cormous 

offshoots. Culms erect or ascending, 

pentaquetrous, rib-angled, soft and easily 

compressed proximally, firm distally, glabrous or 

hispid near apex, 3-10 mm wide. Leaves 7-11 

below inflorescence; sheaths elongate, subinflated, 

smooth to hirtellous or hispid; ligule a thin, 

membranous, brownish flange of tissue at junction 

of sheath and blade, short-ciliate; blades short 

proximally, elongate midculm and distally, linearlanceolate, 

4-30 cm × (4-) 7-20 (-25) mm, 

abaxially smooth, adaxially smooth to hispidulous, 

margins smooth proximally, antrorsely scabrous 

distally, narrowly acute to attenuate at apex. 

Inflorescence a terminal and series of 1-3, 

subcontiguous to remote, simple or compound, 

lateral partial corymbs from the upper leaf-like 

bracts; corymbs dense to sublax, 2-5 cm diam., 

the spikelets in dense, often lobate, spike-like 

clusters at branch tips; spikelets ovoid, ovoidellipsoid, 

ellipsoid-lanceoloid, or cylindrical, 4-8 

(-10) × 1.3-2.2 mm, acute, sub-rounded at base; 

scales oblong to obovate, 2.2-3 × 1-1.3 mm, 

puberulent, strigillose, or rarely smooth, greenish 

brown to dark brown, finely cellular-striate, often 

coarsely beset distally, and on awn basally, with 

coarse tubercle-based translucent hairs, carina 3- 

nerved, prolonged beyond the rounded or retuse 

apex as an elongate, linear-lanceolate, antrorsely 

scabrous, excurved awn. Hypogynous segments 

represented by a single whorl of 3 bladed bristles 

with crimped claws, the blades oblong, obovate, 

or turbinate, 1.3-2 × 0.4-1 mm, obtuse, rounded, 

or emarginate, thinly herbaceous, 3- to 5-nerved, 

incurved-awned, the awn often coiled apically. 

Stamens 2 or 3, the anthers 0.6-0.9 mm long, 

apiculate. Achene broadly ovoid or broadly 

ellipsoid-obovoid with concave sides, 0.9-1.5 × 

0.5-0.8 mm (including beak), stipitate, brownish, 

with pale, wire-like angles, faintly striatecancellate, 

the beak smooth or papillate distally. 


America, West Indies, South America, Africa, 

Asia, Malesia, Australia, and Pacific Islands. 

Distribution in Puerto Rico: Low, wet areas 

in marshy or swampy places, forest openings, 

riversides, drainage canals, roadside ditches, 

pastures, agricultural fields, and disturbed areas. 

Aguada, Arecibo, Bayamón, Cataño, Ceiba, Cidra, 

Dorado, Fajardo, Guayama, Isabela, Lares, 

Luquillo, Manatí, Mayagüez, Naguabo, Patillas, 

Río Grande, San Juan, San Sebastián, and Vega 

Baja. 



10746 (UPRRP, US). Cidra: Pueblo Viejo, Hioram 

92 (US). Lares, Sargent 3252 (US). Manatí: Bo. 

Tierras Nuevas Saliente, area 0.7-1.4 km E of road 

686 at Baldwin School (SE of El Pulguero), 

Proctor & Concepción 42205 (US). Mayagüez: 

Las Mesas, Holm 237 (US). Naguabo: Road off 

P.R. 191 to Pico del Este, Pfeifer et al. 2551 (US). 

Patillas: Bo. Muñoz Rivera, Axelrod & Weisel 

11125 (UPRRP). Río Grande: Along Rt. 186 

bordering Caribbean National Forest, Kelloff et al. 

366 (US). San Juan: 2 mi. E of Santurce, Heller & 

Heller 247 (US). San Sebastián, Sargent 247 (US). 

Sierra de Luquillo, Sintenis 1471 (US). 


Fuirena squarrosa Michx., Fl. Bor.-Amer. 1: 37. 

1803. Cited for western Puerto Rico by Urban 

(1903) and Britton & P. Wilson (1923) based 

on a determination by C. B. Clarke of a 

drawing done by Consul Krug. No historical 

specimens were seen during the preparation 

of this treatment, nor has it been collected 

recently on the island.


Fuirena simplex Vahl, Eclog. Amer. 2: 8. 1798. 

Cited by Kral (Sida 7: 334. 1978) for Puerto 

Rico. No historical specimens were seen 

9. KYLLINGA 

Kyllinga Rottb., Descr. Icon. Rar. Pl. 12. 1773, nom. conserv. 

during the preparation of this treatment, nor 

has it been collected recently on the island. 

Small, slender perennials or rarely annuals, with creeping, elongate or short knotty rhizomes, 

or tufted. Culms 3-angled or obscurely so, smooth, glabrous. Leaves basal and often 1-2 lower cauline; 

sheaths often short, finely veined, the inner band membranous, closed at summit, with a concave to 

truncate orifice; ligule absent; blades when present flat or somewhat pleated, the margins and abaxial 

midvein scabrous, green, glabrous. Inflorescence a subglobose head of 1-3 (-4), crowded, sessile spikes; 

spikes cylindrical, ovoid, or globose, densely-flowered with slender rachis; spikelets ovate to lanceolate, 

flattened, 1-flowered, disarticulating at base when mature, falling entire, densely disposed on the rachis; 

scales 4, 2-ranked, the fertile scale ovate, folded, hyaline or membranous, with a scabrous or smooth 

keel, sometimes winged, mucronate or mucronulate and often recurved at apex, the sterile scales 2, 

minute, basal, the apical scale similar to the fertile one, often sterile, rarely staminate. Flowers bisexual 

and staminate; hypogynous squamellae or bristles absent; stamens 1-3, the anthers oblong-elliptic to 

linear, apiculate at apex; style 2-branched, the unbranched portion uniform at base. Achene laterally 

flattened with one angle facing the rachilla, narrowly ovate to oblong or elliptic, obtuse to sub-rounded 

or subtruncate at apex, apiculate, cuneate to rounded at base, substipitate to distinctly stipitate, the 

surface puncticulate. A primarily pantropical genus of approximately 40 species, 8 of which occur in 

temperate, subtropical and tropical regions of the New World. 

TYPE: Kyllinga nemoralis (J. R. Forst. & G. Forst.) Dandy ex Hutchinson & Dalz. 

(≡ Thryocephalon nemorale J. R. Forst & G. Forst.), typ. conserv. 

Reference: Tucker, G.C. 1984. A revision of the genus Kyllinga Rottb. (Cyperaceae) in Mexico 

and Central America. Rhodora 86: 507-538. 

Key to the species of Kyllinga 

1. Spikelet scales obliquely shaped from the broadly winged keel with closely spinulose-scabrous margin, 

snowy white, becoming tawny or whitish brown at maturity, variegated with reddish brown 

.......................................................................................................................2. K. nemoralis 

1. Spikelet scales uniform, curvate-keeled, unwinged and regularly or remotely spinulose-scabrous on 

keel, or rarely entire, white, whitish brown, greenish, or pale brown. ............................................ 2 

2. Plants with horizontally creeping rhizomes, the culms forming in a row................................ 3 

3. Lowermost involucral bract erect, leaflike but appearing like a continuation of the culm 

..........................................................................................................................1. K. brevifolia 

3. Lowermost involucral bract ascending to reflexed, rarely erect....................................... 4 

4. Involucral bracts 3-7 (-20) mm long, subcucullate; spikes globose; denuded rachis 

spherical to hemispherical ...................................................................... 5. K. tibialis 

4. Involucral bracts 10-30 (-80) mm long, flattened; spikes subglobose to ovoid; denuded 

rachis cylindrical to conica1 ....................................................... 6. K. vaginata 

2. Plants with short rhizomes or rhizomes lacking, the culms caespitose or tufted, not forming in a 

row....................................................................................................................................... 5 

5. Perennial; culms loosely caespitose; basal sheaths short, brown, scarcely tinged with red; 

scales whitish with pale or light green keel; anthers 0.6-0.8 mm long; achene maturing 

reddish brown to dark brown with 0.2-0.3 mm wide, broadly short-stipitate base........... 

...................................................................................................................... 3. K. odorata

318 


5. Annual or short-lived perennial; culms in dense tufts; basal sheaths red-tinged with dark 

red veins; scales pale brown with conspicuous green or dark green keel; anthers 0.2-0.4 

mm long; achene maturing light brown with 0.1 mm wide, narrowly short-stipitate base 

.........................................................................................................................4. K. pumila 

1. Kyllinga brevifolia Rottb., Descr. Icon. Rar. 

Pl. 13. 1773; Cyperus brevifolius (Rottb.) 

Hassk. Cat. Hort. Bot. Bogor. 24. 1844. Type: 

East Indies. Koenig s.n. (holotype: C-Rottb.; 

isotype: C-Rottb.). 

Kyllinga monocephala sensu Thunberg, Fl. Jap. 

35. 1784; and Grisebach, Fl. Brit. W. I. 568. 

1864, non Rottbøll, 1773. 

Kyllinga cruciformis Schrad. ex Schult., Mant. 2: 


Islands. Ehrenberg 69 (holotype: B, 

destroyed; isotypes: C, HAL). 


tall; rhizome long-creeping, scaly, 2-3 mm thick. 

Culms loosely caespitose at first, later narrowly 

to widely spaced along the rhizome, forming in a 

row, soft, triquetrous, deeply channeled, 0.6-1.2 

(-1.5) mm wide, with 2-3 mm wide sheathing 

bases. Leaves 2-4; sheaths short, basal and lower 

cauline, the lowermost bladeless, basal ones 

brownish or at most slightly reddish tinged, the 

inner band reddish brown lineolate; blades 1-15 

(-20) cm × 1.5-3.5 (-4) mm, flattened. Involucral 

bracts 3 (-4), leaf-like, 3-15 (-20) cm long, the 

lowest erect, appearing as a continuation of the 

culm, the upper ones ascendent to horizontally 

spreading or reflexed; spike 1, globose or broadly 

ovoid-globose to cylindrical, 4-8 (-9) × (4-) 5-6 (- 

7) mm; spikelets (20-) 40-60 (-100), ellipticlanceolate, 

2-3 (-3.5) × 0.6-0.9 (-1.2) mm; fertile 

scales ovate-elliptic, 2-2.8 × 1-1.6 mm, 

submembranous, dingy to pale brown or greenish, 

variegated with reddish brown, laterally 2-3 

nerved, keel green, unwinged, spinulose-scabrous, 

often remotely so, rarely entire, prolonged at apex 

as a short, recurved cusp. Stamens (1) 2, the 

anthers 1-1.2 mm long. Achene elliptic to oblongelliptic 

or elliptic-obovate, 1-1.5 × 0.6-0.9 mm, 

broadly rounded to subtruncate at apex, shortcuneate 

at base, yellowish brown to brown. 


States, Mexico, Central America, West Indies, 

South America, and Old World Tropics. 


Islands: Wet to moist meadows, roadsides, ditches, 

gravel washes, river banks, lawns, pastures, pond 

margins, forest edges, and disturbed areas. 

Adjuntas, Aguas Buenas, Añasco, Arecibo, 

Arroyo, Bayamón, Cabo Rojo, Caguas, 

Canóvanas, Carolina, Cataño, Cayey, Ceiba, 

Dorado, Fajardo, Guayama, Humacao, Loíza, 

Luquillo, Manatí, Maricao, Mayagüez, Naguabo, 

Patillas, Peñuelas, Ponce, Río Grande, Salinas, San 

Germán, San Juan, San Sebastián, Toa Baja, 

Trujillo Alto, Vega Alta, Vega Baja, Villalba, and 

Yabucoa; St. Croix, St. Thomas, and Tortola. 


Adjuntas: Rd. 10, km 9, González-Más 869 (US). 

Arroyo: Playa Guilarte, González-Más 1314 (US). 


(US); Hato Tejas, Goll 247 (US). Cataño: Palo 

Seco, González-Más 2105 (US). Dorado: Rd. 693 

to Cerro Gordo, González-Más 337 (US). 


(US). Humacao: González-Más 1350 (US). 

Luquillo: Luquillo Forest, road towards top of El 

Yunque Mt, González-Más 1553 (US). Manatí: Bo. 

Tierras Nuevas Saliente, Axelrod et al. 10962 

(UPRRP). Maricao: Rd. 120, González-Más 427 

(US). Mayagüez: Sintenis 13 (US). Naguabo: Rd. 

3, Daguao River, González-Más 215 (US). San 

Juan: At Río Piedras Experimental Station, 

Woodbury s.n. (US). San Sebastián: entrance to 

town near baseball park, González-Más 1480 (US). 

Toa Baja: Bo. Sabana Seca, area S of Rt. 867, near 

Naval Station, Axelrod & Pérez 9828 (US).Vega 

Alta: Rd. 2, km 30.8, González-Más 1110 (US). 

Villalba: New road under construction to Jayuya, 

González-Más 2193 (US); Reserva Forestal de 

Toro Negro, Stimson 1473 (US).. Yabucoa: Rd. 3, 

km 96.2, González-Más 1344 (US). ST. CROIX: 

Spring garden, A.E. Ricksecker 479 (US). ST. 

THOMAS: St. Peter, Eggers s.n. (NA); Eggers s.n. 

(US). TORTOLA: Road Town to High Bush, N.L. 


2. Kyllinga nemoralis (J. R. Forst. & G. Forst.) 

Dandy ex Hutchinson & Dalz., Fl. W. Trop. 

Afr. 2: 486. 1936; Thryocephalon nemorale 

J. R. Forst & G. Forst., Char. Gen. Pl. 65, t.


65. 1775; Cyperus kyllinga Endl., Cat. Horti 

Vindob. 1: 94. 1842, nom. illeg. Lectotype: 

Society Islands; Otaheite. Forster & Forster 

s.n. (BM), designated by Nicolson, Forsters: 

184. 2004. 

Kyllinga monocephala sensu G. Forster, Fl Ins. 

Austr.: 6, n. 30. 1786, non Rottbøll, 1773 

(which is a nom. superfl. for Schoenus 

coloratus L., 1753). 

Kyllinga triceps sensu G. Forster, Fl. Ins. Aust.: 7, 

n. 31. 1786, non Rottbøll, 1773 (which is a 

nom. superfl. for Scirpus glomeratus L., 

1753). 

Rhizomatous perennial, 7-45 cm tall; rhizome 

long-creeping, scaly, 2-3 mm thick. Culms 

narrowly to widely spaced along the rhizome, firm 

but flexuous, triquetrous or trigonous proximally, 

0.7-1.1 mm wide, with 2-3 mm wide sheathing 

bases. Leaves 3-6, the basal ones often bladeless; 

sheaths lower cauline, pale brown to brown or 

purplish brown; blades 8-30 cm × 2-3.5 (-5) mm, 

flattened. Involucral bracts 3-4, leaf-like, to 30 cm 

long, horizontally spreading to reflexed, rarely the 

lowermost erect; spikes 1 (-3), globose, the 

terminal one 5-8 (-10) × 5-8 mm, lateral secondary 

spikes smaller; spikelets 100-200, obliquely ovateelliptic 

to ovate-lanceolate or elliptic-lanceolate, 

2.7-3.5 × 1-1.8 mm; fertile scales oblique from 

the broadly winged keel, ovate, 2.5-3.5 × 1-2 mm, 

submembranous, white to brownish white or pale 

brown at maturity, variegated with reddish brown, 

laterally 3-4 nerved, keel green, broadly winged 

with closely spinulose-scabrous margin, prolonged 

at apex as a short, recurved mucro. Stamens 3, the 

anthers 0.5-1 mm long. Achene elliptic-obovate 

to oblong-obovate, 1.2-1.5 × 0.5-0.8 mm, obtuse 

to sub-rounded at apex, short cuneate at base, 

yellowish brown to dark brown or blackish at 

maturity. 

General distribution: Tropics and subtropics 

of Africa, Asia, Malesia, Australia, and the Pacific 

region including Hawaii. Introduced in the 

Neotropics where it has been found in Dominica, 

Puerto Rico, and reported from Brazil. 

Distribution in Puerto Rico: Roadsides and 

roadbanks, pastures, lawns, weedy lots, disturbed 

areas, and waste places. Arecibo, Arroyo, Caguas, 

Carolina, Comerío, Fajardo, Guaynabo, Gurabo, 

Humacao, Manatí, Mayagüez, Naguabo, Ponce, 

Río Grande, San Juan, Trujillo Alto, and Utuado. 



10713 (US). Gurabo: Bo. Jaguas, Finca Laing, 1.3 

km due NNW of Escuela Jaguas, Proctor 45129 

(US). 

3. Kyllinga odorata Vahl, Enum. Pl. 2: 382. 1805. 

Type: South America. Rohr s.n. (holotype: C). 

Kyllinga sesquiflora Torr., Ann. Lyceum Nat. Hist. 

New York 3: 287. 1836; Cyperus sesquiflorus 

(Torr.) Mattf. & Kük. in Engler, Pflanzenr. IV. 

20 (Heft 101): 591. 1936. Type: United States, 

Florida. Chapman 12 (holotype: NY). 

Fig. 50. K-N 

Tufted perennial, 4-30 cm tall; rhizome short, 

nodose; plant fragrant with odor of citronella. 

Culms crowded along nodes of the rhizome, 

obscurely 3-angled, somewhat compressed, 

distinctly ribbed, 0.5-1.2 mm wide. Leaves 5-20; 

sheaths pale brown to brown, reddish tinged, the 

membranous inner band red-dotted; blades 3-18 

(-30) cm × 2-3 (-4) mm, many-veined, glabrous, 

the apex long-acuminate. Involucral bracts (1-) 3 

(-4), leaf-like, 1-8 (-12) cm × 1-3 mm, exceeding 

the inflorescence; spikes 1-3, ovoid to cylindrical, 

the terminal one 5-12 × 5-8 mm, lateral spikes 

generally smaller; spikelets ovate to ovatelanceolate, 

2-3 × 1-1.2 mm; fertile scales broadly 

ovate, 2-2.7 × 1.2-2 mm, curvate-keeled, 3-4 

nerved on each face, glabrous, whitish to pale 

brown at maturity, minutely red-lineolate, the keel 

recurved at apex, short mucronate, smooth or 

remotely spinulose-scabrous towards base. 

Stamens 2, the anthers 0.6-0.8 mm long. Achene 

oblong-ovate, 1.2-1.5 × 0.7-1 mm, broadly 

rounded to subtruncate at apex, sub-rounded at 

base, broadly stipitate, with a 0.2-0.3 mm wide 

whitish or stramineous stipe, reddish brown to dark 

brown. 

General distribution: Pantropical, 

southeastern United States to eastern Texas, 

Mexico, Central America, West Indies, South 

America, tropical Africa, Madagascar, southeast 

Asia, Malesia, and northern Australia. 


Islands: Wet ground in pastures, lawns, roadsides, 

and gravel washes. Adjuntas, Bayamón, Corozal, 

Loíza, Mayagüez, Salinas, San Juan, and Vega 

Baja; St. John and St. Thomas. 


Mayagüez: Bo. Miradero, González-Más 1498

320 


Fig. 50. A-E. Eleocharis geniculata. A. Habit and detail of culm showing sheath apex. B. Spikelet. C. Spikelet scale. D. Flower. 

E. Achene. F-J. Fimbristylis dichotoma. F. Habit, and detail of culm showing apex of sheaths. G. Inflorescence. H. Spikelet. I. 

Spikelet scale. J. Achene with style still attached. K-N. Kyllinga odorata. K. Habit. L. Spikelet. M. Flower. N. Achene. From 

Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New York Bot. Gard. 78.


(US). Vega Baja: Laguna Yeguada, near Vega Baja, 

Britton et al. 6772 (NY). ST. JOHN: Rosenberg, N.L. 

Britton & Shafer 304 (US). ST. THOMAS: Eggers 

s.n. (NA). 

4. Kyllinga pumila Michx., Fl. Bor.-Amer. 1: 28. 

1803; Cyperus densicaespitosus Mattf. & Kük. 

ex Kük. in Engler, Pflanzenr. IV. 20 (Heft 101): 

597. 1936. Type: United States; Illinois. 

Michaux s.n. (holotype: P: isotype: P). 

Kyllinga flexuosa Boeck., Beitr. Cyper. 2: 1. 1890. 

Type: Jamaica. Eggers 3753 (holotype: B, 

destroyed). 

Kyllinga odorata sensu Duss, Ann. Inst. Col. 

Marseille 3: 543. 1897, non Vahl, 1805. 

Densely tufted annual or short-lived perennial, 

7-40 (-65) cm tall. Culms firm but flexuous, 

triquetrous distally, often trigonous near base, finely 

and coarsely ribbed and deeply channeled, 0.7-1.3 

mm wide. Leaves 3-5, the basal ones often 

bladeless; sheaths elongate, the basal ones pink 

tinged or reddish with dark red veins; blades 4-20 

(-30) cm × 1.5-3 (-3.8) mm, flattened to V-shaped. 

Involucral bracts 3-5, leaf-like, slightly ascending 

to horizontal, rarely the lowermost erect; spikes 1- 

3, globose-ovoid to cylindrical, the terminal one 

5-8 (-10) × 4-8 mm, lateral secondary spikes 

smaller; spikelets 50-150, lanceolate to oblonglanceolate, 

(1.9-) 2.3-3 (-3.8) × 0.7-1.1 mm; fertile 

scales ovate, 1.8-3.1 (-3.4) × 1-1.7 mm, pale 

brownish to hyaline, laterally 2-4 nerved, the keel 

green, unwinged, with 3-10 antrorsely spinulosescabrous 

barbs, up to 0.2 mm long, rarely smooth, 

prolonged at apex as a short mucro. Stamens 2, the 

anthers 0.2-0.4 mm long. Achene ellipsoid-obovoid 

to oblong-obovoid, 1-1.2 (-1.4) × 0.5-0.7 mm, 

subtruncate at apex, cuneate to sub-rounded at base, 

light brown to yellow-brown. 

General distribution: Eastern United States 

south to Texas, Mexico, Central America, West 

Indies, South America, and tropical Africa. 

Distribution in Puerto Rico: Wet soils in 

meadows, pastures, roadsides and road banks, 

ditches, along trails in forested areas, gravel 

washes, river banks, irrigation ditches, lawns, and 

disturbed open areas. Adjuntas, Aibonito, Arroyo, 

Barranquitas, Caguas, Carolina, Cayey, Ceiba, 

Fajardo, Humacao, Jayuya, Luquillo, Maricao, 

Mayagüez, Naguabo, Orocovis, Ponce, Río Grande, 

San Juan, San Lorenzo, Santa Isabel, Utuado, and 

Villalba. 


Adjuntas: Sintenis 4612 (US). Barranquitas: 

Sargent 367 (US). Caguas: Mil. Road, 9 mi. pt., 

Heller & Heller 1401 (US). Humacao: Coto 

Nabú, km 27.1, González-Más 61 (US). Maricao: 

Rd. 109, Maricao to Mayagüez, González-Más 

1587 (US). Naguabo: Sierra de Luquillo, 

Caribbean National Forest, Rd. 191, km 20, 

Proctor & Thomas 43201 (US). Río Grande: 

Sierra de Luquillo, Caribbean National Forest, 

Proctor 42072 (US). San Juan: Río Piedras, 

Stevenson 1808 (US). Utuado: Vicinity of Utuado, 

N.L. Britton & Cowell 414 (US). Villalba: near 

Salto Doña Juana, González-Más 2190 (US). 

5. Kyllinga tibialis Poit. ex Ledeb. in Ledebour 

& Alderstam, Diss. Bot. Pl. Doming. 6. 

[May] 1805. Type: Haiti. Poiteau s.n. 

(holotype: LE). 

Mariscus aphyllus Vahl, Enum. Pl. 2: 373. [Oct.] 

1805; Kyllinga aphylla (Vahl) Kunth, Enum. 

Pl. 2: 127. 1837. Type: Senegal. Dupuis s.n. 

(holotype: C-Vahl). 

Kyllinga peruviana var. foliata Kük., Repert. 

Spec. Nov. Regni Veg. 12: 92. 1913; Cyperus 

peruvianus var. foliatus (Kük.) Kük. in 

Engler, Pflanzenr. IV. 20 (Heft 101): 587. 

1936. Type: Jamaica. Britton & Hollick 2705 

(holotype: B, destroyed; isotype: NY!). 


tall; rhizome horizontally-creeping, hardened, 

bearing finely and closely veined brown to 

reddish brown scales, 3-6 mm thick. Culms 

arising singly from each node of the rhizome, 

arranged in a row, narrowly spaced, rounded 

trigonous proximally to obtusely trigonous 

distally, hollow but thick-walled, somewhat firm 

but fairly easily compressed, finely longitudinally 

striate, finely to coarsely ribbed and channeled, 

1.5-3.5 mm wide, sheathing base 2-6 mm wide. 

Leaves 3-7, often bladeless; upper sheaths 

elongated, 3-12 cm long, finely veined, reddish 

lineolate or speckled adaxially, cinnamoncolored, 

those lacking blades with an obliquely 

truncate, conspicuously cinnamon-brown, 

scarious orifice, the inner band membranous, 

reddish lineolate or speckled; basal sheaths short, 

cataphylloid, pale brown, tinged with red; blades, 

when present, flattened to subinvolute, finely 

veined, 1-8 cm × 3-7 mm, margins scabrellate 

near apex. Involucral bracts 3-4, short, scarious-

322 


margined proximally, often antrorsely spinulosescabrous 

on margins distally, 3-7 (-20) mm long, 

subcucullate, often divergent to reflexed at 

maturity, the apex acute to acuminate with a stiff, 

thickened tip; spike 1, globose, 7-12 mm in diam., 

the rachis spherical to hemispherical; spikelets 

100-200, broadly to narrowly elliptic-lanceolate, 

(3-) 3.2-3.8 (-4.5) × 1-1.4 mm; fertile scales ovate, 

2.5-3.3 × 1.6-2 mm, submembranous, greenish or 

brownish white, semi-glossy, laterally 5- to 7nerved, 

keel whitish or greenish, unwinged, entire 

or with 1-3 spinulose teeth, 3-nerved, prolonged 

as a short mucro at apex. Stamens 3, the anthers 

0.8-1.3 mm long, apiculate at apex. Achene 

oblong-obovate, 1-1.5 × 0.7-0.9 mm, broadly 

rounded to subtruncate at apex, cuneate at base, 

orangish brown to reddish brown. 

General distribution: West Indies, Central 

America, the Caribbean coast of Colombia, and 

tropical West Africa. 

Distribution in Puerto Rico: Primarily coastal, 

on wet gravels and sands, in sand dunes, wet 

meadows, marshes, ditches, roadsides, and 

disturbed areas. Añasco, Cataño, Cayo 

Palominitos, Ceiba, Humacao, Loíza, Luquillo, 

Manatí, Mayagüez, Naguabo, Río Grande, San 

Juan, and Vega Baja. 


Añasco: Sargent 637 (US). Ceiba-Naguabo: 

Caribbean National Forest, Axelrod & Grose 11211 

(UPRRP). Humacao: Playa de Humacao, Eggers 

595 (US); to Ceiba, km 76.7, González-Más 76 

(US); González-Más 249 (US). Mayagüez: N.L. 

Britton 2359 (US); Sintenis 130 (US). Naguabo: 

Rd. 191, km 14.7, Sierra de Luquillo, El Yunque, 


6. Kyllinga vaginata Lam., Tabl. Encycl. 1: 148. 

1791; Kyllinga peruviana Lam., Encycl. 3: 

366. 1792, nom. illeg.; Cyperus peruvianus 

F. N. Williams, Bull. Herb. Boiss., ser. 2. 7: 

90. 1907. Type: Peru. Dombey s.n. (lectotype: 

P-Lam., photo at DUKE; isolectotype: C), 

designated by G. C. Tucker, Rhodora 86: 525. 

1984. 

Kyllinga pungens Link, Hort. Berol. 1: 326. 1827. 

Type: not located. 

Kyllinga obtusata J. Presl & C. Presl in C. Presl, 

Reliq. Haenk. 1: 183. 1828; Cyperus 

obtusatus (J. Presl & C. Presl) Mattf. & Kük. 

in Engler, Pflanzenr. IV. 20 (Heft 101): 585. 

1936. Type: Peru. Haenke s.n. (holotype: PR). 


rhizome horizontally creeping, knotty, hardened, 

bearing light brown to brown scales which often 

have reddish veins, 2.5-4.3 mm thick. Culms 

arising singly from each node of the rhizome, 

arranged in a row, narrowly to widely spaced, 

trigonous, hollow, easily compressed, finely and 

coarsely ribbed, (0.6-) 0.8-2 mm wide, the 

sheathing bases 1.8-4 mm wide. Leaves 3-5, basal 

and lower cauline, upper blade-bearing, lowermost 

bladeless; blade-bearing sheaths 2-10 cm long, 

with a concave to U-shaped orifice; upper 

bladeless sheaths prolonged dorsally, with an acute 

to short-acuminate, mucronate apex, light brown, 

the membranous inner band red- or blackspeckled; 

basal sheaths cataphylloid, pale brown, 

reddish lineolate; blades, when present, short, 1- 

12 cm × 2-4.5 mm, flattened, finely veined, semiglossy 

and finely cellular-reticulate adaxially, 

margins and abaxial midvein scabrellate, shortacuminate 

to acute at apex. Inflorescence a single, 

subglobose to ovoid spike at the summit of the 

culm, 7-11 mm in diam., the rachis cylindrical to 

conical with 70-200 (-250) spikelets; involucral 

bracts 2-4, leaf-like, 1-5 (-8) cm long, acuminate 

to a triquetrous tip, spreading horizontally, often 

divergent to reflexed at maturity; spikelets ellipticlanceolate, 

2.7-4 × 1-1.5 mm, flattened, acuminate 

at apex, short-cuneate at base; fertile scales ovate, 

2.4-3.4 × 1.6-2.2 mm, thinly herbaceous, whitish 

to pale reddish brown, laterally 3-4 (-5) nerved, 

keel whitish or greenish, unwinged, with 2-4 

spinulose teeth, prolonged as a short, slightly 

recurved mucro at apex. Stamens 3, the anthers 

0.9-1.3 mm long, apiculate at apex. Achene 

oblong-ellipsoid, 1-1.3 × 0.4-0.7 mm, broadly 

rounded at apex, cuneate at base, light brown. 


Central America, South America, and tropical 

Africa. 


Islands: Wet meadows, roadsides, and coastal 

sands. Añasco, Humacao, Mayagüez, and Río 

Grande; cited for St. Croix by Britton (1918) and 

Fosberg, Rhodora 78: 85. 1976. 


Añasco: Sargent 638 (US). Mayagüez: Sintenis 13 

(US).


10. LAGENOCARPUS 

Lagenocarpus Nees, Linnaea 9: 304. 1834. 

Rhizomatous, monoecious or dioecious perennials; rhizomes short and stout, often vertical, 

sometimes emitting scaly stolons. Culms central, erect, trigonous or subtriquetrous, stiff, subhardened. 

Leaves basal and often cauline, with well-developed blades or reduced to bladeless cataphylls; sheaths 

elongate, the inner band prolonged as a rounded-deltate contraligule at the orifice; ligule absent; blades 

linear, 1- to 3-costate, plicate or subflattened, thickly herbaceous, substiffened. Inflorescence a compound 

panicle often consisting of a terminal and series of cymose partial panicles from the upper leaf-like 

bracts, these remote to subcontiguous, the lower ones staminate and the upper pistillate in monoecious 

plants; spikelets unisexual, oblong-ellipsoid or ellipsoid-obovoid; staminate spikelets many-flowered, 

borne singly or in clusters or glomerules at branch tips; pistillate spikelets bearing a single terminal 

floret, with 4-6 empty (sterile) scales below, borne singly or rarely paired; scales often pubescent, 

mucronate or short-awned. Flowers spirally imbricate; hypogynous squamellae typically 3 or wanting, 

minute and often inconspicuous, scale-like or rarely bristle-like, long-pilose, ciliolate, or entire on margins; 

stamens 1 or 2 (-6); style 3-branched. Achene trigonous or obtusely so, flask-shaped or bottle-shaped, 

sometimes beaked, estipitate or sometimes short-stipitate, smooth or pitted, rarely pubescent, often 

trisulcate or tricostate on margins. Approximately 30 species with Neotropical distribution: Central 


TYPE: Lagenocarpus guianensis Nees 

References: Koyama, T. 1965. Cyperaceae tribe Lagenocarpeae. Mem. New York Bot. Gard. 12: 8- 

54. Nees von Esenbeck, C.G.D. 1842. Cyperaceae. Pp. 1-226. In: C. Martius, Fl. Brasiliensis 2 (1). 

1. Lagenocarpus guianensis Nees, Linnaea 9: 

304. 1834; Scleria guianensis (Nees) Steud., 

Syn. Pl. Glumac. 2: 177. 1855. Type: Guyana. 

possibly Schomburgk s.n. (holotype: CGE 

[edges of sheet trimmed off, no inscription 

on back], photocopy of CGE at US). 

Lagenocarpus portoricensis Britton, Bull. Torrey 

Bot. Club 50: 55. 1923. Type: Puerto Rico. 

Underwood & Griggs 946 (holotype: NY!; 


Lagenocarpus tremulus sensu Urban, Symb. 

Antill. 4: 126. 1903, non Nees von Esenbeck, 

1843. 

Fig. 51. F-K 

Dioecious perennial, 75-150 (-200) cm tall; 

rhizome vertical, short and stout, 2-3 cm thick, 

bearing fibrous remnants of old leaf sheath bases, 

sometimes emitting coarse scaly stolons. Culms 

solitary or loosely caespitose, subtriquetrous to 

trigonous proximally, obtusely trigonous to 

subterete distally, antrorsely appressedhispidulous, 

glabrescent, 3-12 mm wide near base; 

sheathing bases 1-3 cm wide. Leaves numerous, 

basal and cauline; sheaths distinctly veined, 

septate-nodulose, basal ones slightly spongythickened, 

glabrescent, pale brown to brown 

proximally, often tinged with red, contraligule 

rounded-deltate, stiff, ciliate on margin; blades 30- 

150 cm × 6-23 mm, plicate to subflattened, septatenodulose 

adaxially and abaxially, often 

inconspicuously so, antrorsely scabrous on 

margins, abaxial midcosta, and adaxial lateral 

costate, long-acuminate to triquetrous apex. 

Inflorescence paniculate, composed of a terminal 

and series of 5-9 lateral, cymose, trullate partial 

panicles from the upper leaf-like bracts, sometimes 

2 from a single node, 3-20 × 1-6 (-10) cm, the 

terminal one flexuous and often curving at apex; 

partial panicles staminate or pistillate, the 

staminate borne from proximal 1-5 nodes and the 

pistillate borne from distal 4-5 nodes; panicle axes 

appressed- to spreading-hispidulous, the 

branchlets ascending, often flexuose; spikelets 

unisexual, solitary or in fascicles of 2-5 from 

prophyllar nodes of branchlets, ellipsoid-obovoid, 

2.5-3.5 × 1.3-1.8 mm, the pistillate spreading to 2 

mm wide with maturing achene; scales ciliate on 

margins, puberulent distally, reddish brown; carina 

1- to 3-nerved, prolonged beyond the obtuse, 

truncate, or subemarginate apex as a prickly mucro 

or short awn; staminate scales numerous, oblong, 

2-2.5 × 0.7-1 mm; pistillate scales 6-7, only the 

terminal one fertile, lowermost 1-2 ovate, shortawned, 

uppermost 5 widely oblong-ovate, 1.8-2.2 

× 1.3-1.8 mm, mucronate. Hypogynous

324 


squamellae 3, minute, rounded-obdeltate, ca. 0.1 

mm long or less, eciliate or sparsely ciliate, 

sometimes shortly bristle-tipped, reddish, adherent 

to base of achene. Stamens 1, the anthers 1-1.3 

mm long, with a prickly apiculum. Achene bottleshaped, 

terete or obscurely trigonous, 2.5-3 (-4) × 

1.6-2 mm, sub-rounded at base, conical at apex 

with a truncate tip, smooth, minutely rugulosepapilliate, 

glabrous to sparsely puberulent, brown 

with a dark brown patch on each side. 

General distribution: Central America, West 

Indies, and South America. 

11. LIPOCARPHA 

Lipocarpha R. Br. in Tuckey, Narr. Exped. Zaire 4: 459. 1918, nom. conserv. 

Distribution in Puerto Rico: Savanna-like 

swales. Dorado and Vega Baja. Known only from 

the vicinity of Laguna Tortuguero and Dorado. 

Note: Puerto Rican plants belong to the more 

widely distributed typical subspecies, subsp. 

guianensis. 


Dorado: Sintenis 6845 (US). Manatí: Near rail road 

track from Manatí to Vega Baja, Underwood & 

Griggs 946 (US). Vega Baja: Bo. Algarrobo, just 

S of Laguna Tortuguero, Proctor 45652 (US). 

Annual or rarely perennial herbs, tufted. Culms rounded or broadly elliptical in cross section. Leaves 

crowded at base; sheaths short, closed; ligule absent; blades well-developed, flattened or _ inrolled. 

Inflorescence of 1-few clusters of ovoid or globose spikes, of many spirally arranged reduced 1-flowered 

spikelets, terminal or sometimes appearing lateral (pseudolateral), capitate; involucral bracts several, 

the lower well-developed and leaf-like, the upper smaller and resembling the spike bracts; spike bracts 

obovate to spathulate, often with an acute to acuminate apex; inflorescence prophylls present; spikelets 

with a single bisexual flower (plants hermaphroditic), densely spirally imbricated on a cylindrical axis; 

scales dorsally rounded or flattened, hyaline, sometimes notched at apex, often reduced or wanting, the 

margins free, sometimes enveloping the flower and at maturity the achene. Terminal flowers absent; 

hypogynous squamellae or bristles absent; stamens 1-3; style 2- or 3-branched, glabrous, the base not 

expanded. Fruit an achene, obovoid or ellipsoid-obovoid to narrowly subcylindrical, apiculate or entire, 

often with a constriction at base, minutely papillate or puncticulate, naked or sometimes enveloped or 

clasped by the fertile spikelet scale and falling together with it. Thirty-five species with its center of 

distribution in Africa. Also in the Americas, Madagascar, southeast Asia, Malesia, and Australia. 

TYPE: Lipocarpha senegalensis (Lam.) T. Durand & H. Durand (≡ Lipocarpha argentea R. Br., nom. 

illeg.; ≡ Hypaelyptum argenteum Vahl, nom. illeg.; ≡ Scirpus senegalensis Lam.). 

Reference: Goetghebeur, P. & A. Van den Borre. 1989. Studies in Cyperaceae 8. A revision of 

Lipocarpha, including Hemicarpha and Rikliella. Wagningen Agric. Univ. Pap. 89-1: 1-87. 

1. Lipocarpha micrantha (Vahl) G. C. Tucker, J. 

Arnold Arbor. 68: 410. 1987; Scirpus 

micranthus Vahl, Enum. Pl. 2: 254. 1805; 

Hemicarpha micrantha (Vahl) Pax in Engler 

& Prantl, Nat. Pflanzenfam. 2, Abt. 2: 105. 

1887. Type: French Guiana. Richard s.n. 

(holotype: C-Vahl; isotype: P). 

Scirpus subsquarrosus Muhl., Descr. Gram. 39. 

1817; Hemicarpha subsquarrosa (Muhl.) 

Nees in Martius, Fl. Bras. 2(1): 61. 1842. 

Type: United States; Pennsylvania. 

Muhlenberg s.n. (holotype: B-Willd. 1202). 

Fig. 52. A-E 

Tufted annual, 1-13 (-20) cm tall. Culms 

slender, spreading, often curved outwards, 

obtusely trigonous, finely ribbed, smooth, 0.3-0.5 

mm wide. Leaves 2-3, basal; sheaths loose, 5-25 

mm long, thinly herbaceous, finely veined with 

often reddish veins proximally, reddish brown or 

light brown, obliquely truncate at orifice, the inner 

band hyaline distally, splitting with age; blades 

lanceolate to narrowly linear, subinvolute, 1-20 × 

0.2-0.5 mm, obtuse at apex. Inflorescence a small 

pseudolateral head of 1-3 spikes, 2-7 mm diam.; 

involucral bracts 1-2 (-3), the lowermost erect, 

culm-like, 0.5-5 cm long, the uppermost shorter, 

leaf-like, divergent to reflexed; spikes ovoid to


Fig. 51. A-E. Fuirena umbellata. A. Habit. B. Spikelet. C. Flower. D. Spikelet scale, dorsal view and lateral views. E. Achene. 

F-K. Lagenocarpus guianensis. F. Habit. G. Inflorescence branch. H. Spikelet. I. Spikelet scale. J. Pistil. K. Achene. (A-E, from 

Proctor 48747; F-G, from Proctor 45602; H-K, from Proctor 39464).

326 


oblong-ovoid, 1-5 × 1.3-2.3 mm, obtuse to 

subacute, rounded to subtruncate at base, the rachis 

elongating and becoming denuded proximally with 

falling of mature achenes and bracts; spike bracts 

rhombic-obovate to obtrullate, cuneate at base, 

0.5-1 (-1.3) × 0.3-0.4 mm, scaberulous distally, 

yellowish to red-brown or chestnut brown on sides, 

2- to 3-nerved on each side, carina 3-nerved, 

greenish, converging at apex and forming an acute 

to acuminate tip; spikelet scale 1, oblanceolate, 

hyaline, often wanting. Stamens 1, the anthers 

elliptic, 0.2-0.3 mm long, blunt at apex; style 2branched. 

Achene obscurely trigonous or 

subrounded, obovoid to oblong-obovoid, 0.5-0.8 

12. MACHAERINA 

Machaerina Vahl, Enum. Pl. 2: 238. 1805. 

× 3-4 mm, rounded at apex, short-apiculate, 

cuneate at base, yellowish or purplish brown, with 

a silvery cast at maturity, puncticulate. 


Central America, West Indies, South America, and 


Distribution in Puerto Rico: White sand 

savanna. Vega Baja. Known only from the vicinity 

of Laguna Tortuguero, collected by Woodbury s.n., 

12 Mar 1960 (UPR; photo at US). 


Vega Baja: Laguna Tortuguero, Woodbury s.n. 

(UPR; photo at US). 

Large or medium-sized perennials; rhizomes hardened, woody, short or horizontally creeping, often 

emitting creeping, stout, scaly stolons, 0.5-2 cm thick; roots coarse. Culms central, solitary, compressed, 

lenticular to ancipital, frequently channeled along one edge distally, finely or sometimes coarsely ribbed, 

glabrous. Leaves numerous, 2-ranked, primarily basal, frequently 1-2 (-3) reduced cauline ones medially, 

the upper (3-) 5-11 basal ones bearing elongate blades, the lower with blades shorter than the sheaths or 

wanting; sheaths laterally folded, narrowed to summit with an indentation along margins at junction 

with blade, finely ribbed, septate-nodulose, essentially glabrous; ligule absent; blades unifacial, flattened 

to ancipital in cross section, linear to linear-falcate, curving to an acuminate tip, finely ribbed, green, 

glabrous, the margins entire. Inflorescence 1-3 remote to subcontiguous or contiguous, open or contracted, 

partial panicles; main axes and lateral branches of panicles frequently flexuose; bracts subtending panicles 

blade-bearing, the blades rarely exceeding length of subtending panicle, the uppermost ones often with 

an uncinate apex; spikelets numerous, in turbinate or spherical clusters or fascicles of 2-4 (-6), or 

sometimes solitary, laterally compressed, each subtended by a short or barely sheathing prophyllar 

bract, 2- to 7-flowered, with 0-4 basal scales empty, the scales distichous or sometimes nearly spirally 

arranged; scales laterally subcompressed, thin, boat-shaped, keeled, brown to brown-black, purple-black, 

or black, the margins frequently ciliate. Flowers bisexual; hypogynous bristles 6-3, subulate or flattened, 

antrorsely spinulose distally, or often undeveloped and represented only by very short rudiments at 

base; stamens 3, the anthers sagittate at base, the connective forming an acuminate, lanceolate-subulate 

appendage at apex; style 3-branched, the stigmatic branches minutely scaly or fimbrillate, 1 /3 to ½ length 

of style. Achene with a 3-winged, stipitate base and persistent, triquetrous or 3-winged style base, the 

wings thin and chartaceous or incrassate, decurrent and often with a distinct wire-like margin between 

base of wing and achene body; achene body globose, obovoid, turbinate, or ellipsoid, often with a 

constriction (sometimes indistinct) just above the stipitate cellular-reticulate base, faintly rugose or 

rugulose, finely transversely rugulose or wrinkled, or essentially smooth, the pericarp very thin and 

brittle; style base triquetrous or 3-winged, triangular to triangular-lanceolate, the angles often whitened 

or blackened, crinkled and blistered, scabrous, or beset with crystalline fimbrillae or scales. Approximately 

25 species (here excluding Baumea Gaudich.), Pantropical, many local or relictual: West Indies, South 

America, Africa, Asia, Malesia, and the Pacific Islands (including the Hawaiian Islands). 

TYPE: Machaerina restioides (Sw.) Vahl (≡ Schoenus restioides Sw.). 

Reference: Strong, M. T. 1997. Machaerina (Cyperaceae) in South America. Novon 7: 308-319.


1. Machaerina restioides (Sw.) Vahl, Enum. Pl. 

2: 238. 1805; Schoenus restioides Sw., Prodr. 

19. 1788; Cladium restioides (Sw.) Benth. ex 

C. B. Clarke in Urban, Symb. Antill. 2: 135. 

1900. Type: Montserrat, Ryan s.n. (holotype: 

BM; isotypes: B-Willd. 1149, C). 

Fig. 52. F-J 


cm tall; rhizomes scaly, ca. 1 cm thick. Culms 

lenticular, 1.8-6 (-8) mm wide, sheathing bases 8- 

30 mm wide just above rootstock, widely 

spreading above. Leaves primarily basal, 1-2 

cauline below inflorescence, 5-11, 2-ranked; 

sheaths elongate, light brown, tinged reddish to 

cinnamon-colored, essentially glabrous; blades 

unifacial, linear-falcate, stiffened, indistinctly 

septate-nodulose, (5-) 12-70 (-90) cm × (3-) 5-15 

(-18) mm. Inflorescence a terminal and often 1-2 

remote or subcontiguous, narrowly rhombic partial 

panicles from the upper leaf-like bracts, the 

terminal one largest, 6-17 × 1-4 cm; branches 

flexuose, bearing small turbinate clusters of (1-) 

2-6 spikelets, 3-8 mm in diam.; spikelets ovate, 

3.5-4.5 × 2-3 mm, with 2-5 florets; fertile scales 

2-ranked, often obscurely so, ovate-elliptic or 

oblong-ovate to oblong-lanceolate, dorsally obtuse 

to subrounded, 2.5-3.2 × 1.3-1.7 mm, chartaceous, 

brown-black to dark purple-black or black, with 

dark reddish brown scarious margins proximally, 

dull to sublustrous, minutely antrorsely scabroushispid 

on sides distally, margins ciliate-scabrous, 

lateral veins indistinct, the carina prominent, 

13. REMIREA 

Remirea Aubl., Hist. Pl. Guiane 44, t. 16. 1775. 

antrorsely scabrous (at least distally), sometimes 

extending as a very short mucro at the acute to 

subobtuse apex. Stamens 3, the anthers 1-1.8 mm 

long. Achene 1.8-2.5 × 0.6-0.9 mm (including 

stipitate base); achene body ellipsoid-obovoid, 

ovoid-ellipsoid, or rounded-ovoid, with convex 

sides, indistinctly rugulose to nearly smooth 

medially or sometimes venose, with a distinct 

constriction just above the stipitate base, narrowly 

winged, brown, with pale wing-angles, the base 

triquetrous, cellular reticulate; style base 

triquetrous, triangular-lanceolate to linearlanceolate, 

0.3-0.8 × 0.3-0.4 mm, minutely 

antrorsely scabrous-hispid with crystalline 

prickles, pale at base, reddish distally; bristles 6, 

antrorsely spinulose distally or essentially smooth, 

variable in length, shorter than to 1 or 2 exceeding 

achene body, often caducous, rarely rudimentary. 

General distribution: Endemic to the West 

Indies; Cuba, Hispaniola, Puerto Rico, and the 



from the summit of El Yunque in wet rocky places, 

800-934 m, growing in rock crevices on cliffs or 

open grassy areas. Río Grande. 

Common names: Puerto Rico: Cortadora, 

Cortadera, Cortadora de altura, Lambedora. 


Río Grande: El Yunque, Sargent 549 (US); 

Woodbury s.n. (US); top of El Yunque Mt., 

González-Más 1521 (US); Sierra de Luquillo, 

Hioram 366 (US); Sintenis 1382 (US-2). 

A monospecific genus represented by the species listed below. Pantropical. 

TYPE: Remirea maritima Aubl. 

Reference: Oteng-Yeboah, A.A. 1975. Morphology, anatomy and taxonomy of the genus Remirea 

Aublet (Cyperaceae). Boissiera 24a: 197-205. 

1. Remirea maritima Aubl., Hist. Pl. Guiane 45. 

1775. Type: French Guiana. Aublet s.n. 

(holotype: BM). 

Remirea pedunculata R. Br., Prodr. 236. 1810; 

Cyperus pedunculatus (R. Br.) J. Kern, Acta 

Bot. Neerl. 7: 798. 1958; Mariscus 

pedunculatus (R. Br.) T. Koyama, Gard. Bull. 

Singapore 30: 157. 1977. Type: Australia; 

Queensland. Brown s.n. (holotype: BM). 

Fig. 53. E-H 


rhizome long-creeping, branched, clothed with 

membranous, brownish sheaths and their fibrous

328 


Fig. 52. A-E. Lipocarpha micrantha. A. Habit. B. Inflorescence spike. C. Spikelet scale. D. Flower. E. Achene. F-J. Machaerina 

restioides. F. Habit. G. Inflorescence branch. H. Spikelet scale. I. Flower. J. Achene. (A, from Curtis 5240; B-E, from Kral 

7570; F, from Sargent 549; G-J, from Hioram 366).


remnants, rooting at the nodes, 1-3 mm thick. 

Culms solitary or loosely crowded at tips of 

rhizome, erect, ascending, or nearly prostrate, 

trigonous, rigid, smooth, 1-2.5 mm wide. Leaves 

numerous, primarily cauline, crowded from base 

to apex; sheaths short, basal ones fibrillose with 

age, light brown, the inner band membranous, 

reddish brown, truncate to U-shaped at orifice; 

ligule absent; blades 2-12 cm long, elongating to 

20 cm in sterile individuals, 2-7 mm wide basally, 

linear-lanceolate, rigid, canaliculate, often 

recurved, antrorsely scabrous on margins and 

abaxial midcosta distally, coriaceous, glaucousgreen, 

attenuate to a triquetrous apex, the tip 

pungent. Inflorescence a capitate group of (1-) 3- 

6 (-8) sessile short spikes at the summit of the 

culm, 1.2-2.5 cm diam.; involucral bracts 6-9, leaflike 

but reduced, subreflexed, the lowermost to 8 

cm long, overtopping the inflorescence; spikes 

ovoid or ellipsoid, 7-20 × 5-10 mm, obtuse to 

subrounded; spikelets densely crowded on rachilla, 

ovoid to ovoid-ellipsoid, 3.6-5 × 1.2-2 mm, acute 

at apex, obtuse at base, slightly compressed, 1flowered, 

with a single sterile scale below, 

disarticulating above the scale-like prophyll and 

bracteole, falling as a whole, terminated by a 

vestigial “corky organ”, strongly flattened, ca. 3 

mm long; rachilla of the uppermost internode 

becoming thick and corky at maturity, enveloping 

the achene; fertile scale ovate to broadly ovate or 

ovate-elliptic, 3.5-4.5 × 1.5-2.5 mm, boat-shaped, 

dorsally obtuse, 3- to 5-veined on each side above 

the scarious margins, carina 3-nerved, ending at 

or slightly prolonged beyond the acute apex as a 

blunt mucro; sterile scale like the fertile, but 

slightly smaller. Flower bisexual; hypogynous 

squamellae or bristles absent; stamens 3, the 

anthers 1.6-2.2 mm long, with a short, conical, 

reddish apiculum; style 3-branched. Achene 

tightly enclosed in the upper corky internode of 

the rachilla, trigonous, ovoid-ellipsoid to ellipsoid 

or oblong-ovoid, 2-2.5 × 0.7-1 mm, slightly 

dorsally compressed, short-beaked, obtuse at base, 

estipitate, puncticulate, shiny, dark brown to 

grayish brown or blackish. 


(Florida), Mexico, Central America, West Indies, 

South America, Africa, southern Asia, Malesia, 

Pacific Islands, and Australia. 

Distribution in Puerto Rico: In coastal sands 

of beaches, dunes, strand vegetation, and pastures. 

Arecibo, Camuy, Carolina, Hatillo, Isabela, Loíza, 

and San Juan. 

Common name: Puerto Rico: Junco de playa. 


Arecibo: West of Arecibo, Chase s.n. (US). 

Camuy: Sargent B16 (US). Loíza: N.L. Britton et 

al. 5727 (US). San Juan: Sintenis 1121 (US). 

14. RHYNCHOSPORA 

Rhynchospora Vahl, Enum. 2: 229. 1805 [as “Rynchospora”], nom. et orth. conserv. 

Perennials or sometimes annuals, vegetatively diverse; roots fibrous. Culms caespitose or borne 

singly, trigonous or obscurely so, occasionally cylindrical, shallowly to deeply channeled along one 

side or margin (at least distally) with channel edges often antrorsely scabrous, essentially smooth, 

sometimes scabrous distally, glabrous or sometimes hirsute. Leaves basal or basal and cauline, rarely 

strictly cauline; sheaths green, light brown or reddish brown, sometimes whitened at base, inner band 

membranous on basal sheaths, often purple-dotted, splitting with age, often membranous only at orifice 

on cauline sheaths, the orifice truncate or concave, sometimes U-shaped, rarely convex; ligule absent or 

sometimes present, often a narrow band of thickened tissue or trichomes at adaxial junction of sheath 

and leaf blade; blades flattened, V-shaped, or folded, occasionally involute or crescentform-capillary, 

linear, filiform, or sometimes capillary, herbaceous or occasionally stiff, glabrous, hirsute, or occasionally 

scabrous or scabridulous distally, rarely papillose or transversely rugulose, the margins and abaxial 

midvein usually antrorsely scabrous, ciliate, or with setose hairs. Inflorescence terminal or both terminal 

and with a series of lateral partial panicles from the upper leaf-like bracts, paniculate, corymbose, racemose 

or congested and head-like; involucral bracts in capitate species sometimes whitened at base; branches 

cylindrical, 3-angled, crescentform, or subcompressed, finely ribbed, scabrous, ciliate or smooth on 

margins, the spikelets solitary or in fascicles at branch tips; spikelets ovoid, ellipsoid, lanceoloid, or 

fusiform, cylindrical or subcompressed, primarily 1- to several-flowered, the scales spreading with

330 


maturing achenes; scales spirally imbricate, ovate to lanceolate, shallowly to deeply boat-shaped, often 

inrolled around flower, finely and indistinctly nerved on sides, with a single distinct or sometimes 

indistinct midcosta, light to dark brown or ferrugineous, sometimes whitish. Flowers bisexual above the 

(1-) 2-5 (-9) empty basal scales of spikelet, the terminal often staminate with a rudimentary ovary or 

reduced and empty; bristles (1-) 6 (-20), rudimentary, or absent, when present antrorsely or retrorsely 

barbed, sometimes smooth or plumose; stamens 1-3 (-12), the anthers linear, narrowly elliptic or oblong, 

often with minute crystalline papillae or lobes at base; styles subulate, 2-branched or undivided, often 

long-exserted beyond apex of subtending scale. Achene biconvex to subcylindrical, sometimes inrolled 

with winged or wavy margins, obovate, oblong-obovate or oblong-elliptic, deeply pitted, transversely 

rugulose or sometimes smooth, the expanded, usually triangular, or sometimes discoid style base, 

persistent at the summit. A genus of approximately 270 species with its greatest diversity in the Western 

Hemisphere, particularly warm-temperate North America and the Neotropics. 

TYPE: Rhynchospora alba (L.) Vahl (≡ Schoenus albus L.), typ. conserv. 

References: Gale, S. 1944. Rhynchospora section Eurhynchospora in Canada, the United States, 

and the West Indies. Rhodora 46: 89-134, 159-197, 207-249, 255-278. Guaglianone, R. 1979. Sobre 

Rhynchospora rugosa (Vahl) Gale (Cyperaceae) y algunas especies afines. Darwiniana 22: 255-311. 

Koyama, T. 1972. Cyperaceae-Rhynchosporeae and Cladieae. In: B. M. Maguire, The botany of the 

Guayana Highland-Part 9. Mem. New York Bot. Gard. 23: 23-89. Kükenthal, G. 1949-1951. Vorarbeiten 

zu einer Monographie der Rhynchosporideae. Bot. Jahrb. Syst. 74(3): 375-509. 1949, 75(1): 90-126. 

1950, 75(2):127-195. 1950, 75(3): 273-314. 1951. Thomas, W. W. 1984. The systematics of Rhynchospora 

section Dichromena. Mem. New York Bot. Gard. 37: 1-116. Thomas, W.W. 1992. A synopsis of 

Rhynchospora (Cyperaceae) in Mesoamerica. Brittonia 44: 14-44. 

Key to the species of Rhynchospora 

1. Inflorescence a single capitate cluster of essentially sessile spikelets at the summit of the culm; 

involucral bracts subtending spikelets (3-) 4-8, appearing whorled, elongate, often whitened basally, 

uniformly green in R. radicans subsp. microcephala........................................................ 2 

2. Involucral bracts uniformly green; spikelets 3-6 (-7) ..... 25. R. radicans subsp. microcephala 

2. Involucral bracts whitened basally, at least adaxially; spikelets 5-50...................................... 3 

3. Involucral bracts whitened at base adaxially only; rhizome lacking stolons; culms caespitose; 

anthers (1.7-) 2-3 mm long ........................................................... 5. R. ciliata 

3. Involucral bracts whitened at base both adaxially and abaxially; rhizome producing slender, 

scaly stolons to 20 cm long; culms solitary or rarely 2 together; anthers 1-2 mm long 

............................................................................................................. 6. R. colorata 

1. Inflorescence paniculate, terminal only or a terminal and series of 1-7 lateral, remote to contiguous, 

congested or open, small or large, simple to compound partial panicles from the upper sheathing 

bracts, at least some branching evident or spikelets pedicellate; sheathing or involucral bracts 

subtending each panicle or partial panicle 1 (-2), green.................................................................. 4 

4. Large, robust, stout perennials; leaf blades (5-) 8-23 (-25) mm wide at their widest point; achene 

margins irregular, thickened with constrictions and protuberances; style base spongy-thickened with 

2-lobed sagittate base, as wide as and as thick as or slightly wider and thicker than achene body at 

base with a longitudinal medial groove on both sides........................................................ 5 

5. Adaxial surface of leaf blades dark green, smooth and glossy, the abaxial surface green, septatenodulose; 

spikelets in globose clusters at branch tips ......................... 14. R. gigantea 

5. Adaxial and abaxial surfaces of leaf blades green, semi-glossy, the abaxial surface not septatenodulose; 

spikelets in hemispherical fascicles at branch tips ............ 8. R. corymbosa 

4. Small to medium-sized, slender perennials or annuals; leaf blades 0.2-8 (-10) mm wide at their widest 

point; achene margins entire or flaring at apex in one species; style base not spongy-thickened, 

brittle, at most shallowly lobed at base, often narrower and thinner than achene body with a plane 

surface on both sides................................................................................................... 6


6. Well-developed leaves cauline, 8-40, closely set (at least medially) with short intervals between 

the nodes, those at base and lower cauline reduced to short-bladed or bladeless sheaths. 

................................................................................................................................... 7 

7. Achenes transversely rugulose or indistinctly so; fertile scales 0.8-1.5 mm wide; style base 

conic-subulate; bristles absent .......................................................... 30. R. uniflora 

7. Achenes cellular-cancellate with 11-20 longitudinal rows of cells; fertile scales 1.6-2.8 mm 

wide; style base triangular-lanceolate or oblong-lanceolate; bristles rudimentary or 

sometimes 1-2. ................................................................................................................ 8 

8. Inner band of leaf sheath pilose to wooly at orifice, the surface emuriculate; culms 1.8- 

4 mm wide; leaf blades 2-7 mm wide; achene 0.7-1 mm wide; style base triangularlanceolate, 

shallowly 2-lobed at base ........................... 16. R. jamaicensis 

8. Inner band of leaf sheath short-pilose to glabrous distally, the surface muriculate at 

orifice; culms 0.8-2.3 mm wide; leaf blades 1.3-4 mm wide; achene 1-1.3 mm wide; 

style base oblong-lanceolate, mitriform .......................................... 24. R. racemosa 

6. Well-developed leaves primarily basal, cauline leaves 1-5 (not including upper sheathing bracts 

subtending partial panicles or fascicles of inflorescence), remote, with long intervals between 

the nodes.................................................................................................................. 9 

9. Bristles absent at base of achene.................................................................................... 10 

10. Spikelets (8-) 9-13 (-14) mm long ................................................... 13. R. filiformis 

10. Spikelets 1.5-8 (-9) mm long................................................................................. 11 

11. Spikelet scales white or dingy white; rhizome producing slender stolons; on 

serpentine and limestone soils in secondary forest, often shaded habitats........... 

........................................................................................................ 2. R. berteroi 

11. Spikelet scales light brown, yellowish brown, reddish brown, ferrugineous or 

copper-colored; rhizomes (when present) estoloniferous; on red clay or sandy or 

peaty soils in savannas and grassy or marshy areas, often open habitats in full 

sun............................................................................................................. 12 

12. Achene margins at apex flaring, prolonged into an ear-like projection on 

each side that is confluent with margins of the tounge-like style base.......... 

............................................................................................ 29. R. tenerrima 

12. Achene margins at apex not prolonged, rounded to broadly rounded or 

truncate..................................................................................................... 13 

13. Spikelets 3.2-8 (-9) mm long; fertile spikelet scales numerous, 9-45 or 

more.................................................................................................. 14 

14. Sheaths uniformly pilose; lower corymb branches of inflorescence 

panicle and spikelet pedicels often divergent or recurved at maturity, 

the spikelets often nodding; spikelets broadly ovoid to globose with 

obtuse or broadly acute apex..................................... 10. R. divaricata 

14. Sheaths essentially glabrous or sometimes pubescent proximally; 

lower corymb branches and spikelet pedicels of inflorescence panicle 

ascending; spikelets ovoid to oblong-ovoid or ovoid-ellipsoid with 

acute apex.......................................................... 15 

15. Fertile spikelet scales 2.1-3 mm long; achene 1-1.1 × 1 mm, 

lustrous, yellowish brown to golden brown, maturing to dark 

reddish brown or brown-black, the rugae often lighter in color; 

style base 0.2-0.6 mm long, 1 mm wide at base... 21. R. nitens 

15. Fertile spikelet scales 3.2-6.2 mm long; achene 1.5-1.7 × 1.3- 

1.6 mm, brown, yellowish brown, or dark brown; style base 

0.8-1.2 mm long, 1.2-1.4 mm wide at base............. 32. R. velutina 

13. Spikelets 1.5-3.2 mm long; fertile spikelet scales 2-5........................ 15

332 


16. Style base short bulbiform, 0.1-0.2 mm long, 0.3-0.4 mm wide at base 23. R. pusilla 

16. Style base depressed-trigonous or strap-like, 0.1-0.4 mm long, 0.5-1 mm wide at 

base................................................................................................................................. 17 

17. Style base emphatically depressed trigonous, oval-shaped from top view, not lobed, 

0.1 (-0.2) mm long, often 0.1 mm long or less .......................... 9. R. depressirostris 

17. Style base forming 2 strap-like lobes that are decurrent on shoulders of achene body 

to about the middle, abruptly narrowing distally into a short triangular-lanceolate tip, 

0.3-0.4 mm long ......................................................... 7. R. contracta 

9. Bristles present at base of achene................................................................................................... 18 

18. Bristles plumose for most of their length............................................................................. 19 

19. Bristles ½ length of achene or shorter; inflorescence a single spikelet or small, lax, 

corymbiform panicle of 2-3 spikelets; spikelets subflattened-fusiform, 7-9.5 (-10) mm 

long; rachilla anfractuose-elongate, spirally zig-zag above sterile basal scales; spikelet 

scales 7-10 .................................................................................................. 4. R. breviseta 

19. Bristles equaling to shortly exceeding achene and style base; inflorescence a single terminal 

small contracted corymbiform panicle of numerous spikelets, rarely a second lateral one 

below; spikelets ovoid to ovoid-ellipsoid, (2.5-) 2.8-5 mm long; rachilla not anfractuoseelongate; 

spikelet scales 5-6.................................................. 22. R. plumosa 

18. Bristles antrorsely or retrorsely barbed, sometimes setose at very base.............................. 20 

20. Bristles 10-12, retrorsely barbed ....................................................................... 1. R. alba 

20. Bristles (1-) 3-6, antrorsely barbed............................................................................... 21 

21. Spikelets tightly compacted in globose capitula at ray or branch tips; style base linearlanceolate, 

4-angled, often curved, 1.8-3.1 mm long, 0.2-0.4 (-0.6) mm wide at 

base.................................................................................... 15. R. holoschoenoides 

21. Spikelets in fascicles or solitary at branch tips; style base deltate, triangular, triangularlanceolate, 

lance-attenuate, or depressed-conical, flattened or compressed, not angled 

nor curved, 0.2-1.6 (-2.3) mm long, 0.5-1 mm wide at base...................... 22 

22. Spikelets lanceoloid to fusiform, 8-12 mm long, long-acuminate at apex; achene 

surface finely cellular-cancellate; style base lance-attenuate with concave margins, 

1.2-2.3 mm long........................................ 11. R. domingensis 

22. Spikelets ovoid to broadly ovoid, ovoid-globose, ovoid-ellipsoid, ovoid-lanceoloid, 

or ellipsoid, 2-8 mm long, acute to acuminate at apex; achene surface transversely 

rugose or rugulose, or smooth; style base deltate, triangular, triangular-lanceolate, 

or depressed-conical, 0.5-1 (-1.6) mm long................................... 23 

23. Adaxial surface of leaf blades finely transversely rugulose; achene planoconvex 

or unequally biconvex; plant of serpentine soils (particularly in the 

Maricao region) ................................................................ 17. R. lindeniana 

23. Adaxial surface of leaf blades finely cellular-reticulate or essentially smooth; 

achene biconvex; plants of sandy soils or deposits, damp or wet peats, red 

laterite or red clay soils............................................................ 24 

24. Achene surface essentially smooth............................................... 25 

25. Achenes brown with a light brown or yellowish brown tumid center; 

fertile scale 1, clawed at base; style base 0.5-0.7 mm wide at 

base................................................ 3. R. brachychaeta 

25. Achenes dark brown to brownish black or blackish with an often 

red or yellowish circular patch medially; fertile scales 2-4, not clawed 

at base; style base 0.7-1 mm wide at base.... 26 

26. Leaf blades 1-3 mm wide; achene obovate or suborbicular, dark 

brown to blackish with a yellowish brown to deep red circular 

patch medially; style base acute to short-acuminate at tip 

........................................................ 12. R. fascicularis


26. Leaf blades 0.2-1.3 (-2) mm wide; achene elliptic, dark brown to brownish black with a 

distinct or indistinct somewhat paler brown or reddish area medially; style base with a 

broad bill-like extension at tip ............................................................... 32. R. wrightiana 

24. Achene surface distinctly transversely rugulose or pitted-reticulate............................................ 27 

27. Leaf blades 0.2-1 mm wide; spikelets few, in open corymbiform partial panicles on elongate 

filiform pedicels .............................................................................................. 26. R. rariflora 

27. Leaf blades (0.7-) 1-8 (-10) mm wide; spikelets in congested corymbiform partial panicles, 

fasciculate, short-pedicellate................................................................................................ 28 

28. Style base triangular-lanceolate, 1.2-1.6 mm long; bristles often 3 times as long as achene 

.................................................................................................... 18. R. marisculus 

28. Style base triangular, deltate, or depressed-conical, 0.2-1.1 mm long; bristles shorter than 

to equaling achene or at most 2 times as long as achene. ...................................... 29 

29. Spikelets 5.6-8 mm long, acuminate at apex; achene shortly attenuate at base to a 0.5 

mm long glossy reddish brown stipe; stamens often marescent..... 21. R. odorata 

29. Spikelets 2.5-4.7 (-5) mm long, acute at apex; achene not stipitate; stamens 

evanescent............................................................................................................... 30 

30. Apex of fertile scales acuminate; bristles with crystalline trichomes at base...... 

.......................................................................................................... 28. R. rugosa 

30. Apex of fertile scales obtuse to emarginate; bristles lacking trichomes at base... 

........................................................................................................................... 31 

31. Achenes thinly biconvex, obovate to widely depressed-obovate; style base 

deltate, essentially confluent with the slightly rimmed apex of achene........ 

........................................................................................... 19. R. microcarpa 

31. Achenes thickly biconvex, broadly obovate to subrounded; style base 

depressed-conical or subtriangular with a constriction between its base and 

apex of achene .................................................................... 27. R. recognita 

1. Rhynchospora alba (L.) Vahl, Enum. Pl. 2: 236. 

1805; Schoenus albus L., Sp. Pl. 44. 1753. 

Lectotype: Europe. (LINN-68.17), designated 

by D. A. Simpson in Cafferty & C. E. Jarvis 

(eds.), Taxon 53: 179. 2004. 

Rhynchospora luquillensis Britton, Bull. Torrey 


Sierra de Luquillo. Hioram 364 (holotype: 

NY!; isotypes: NY!, US!). 

Fig. 53. A-D 

Caespitose perennial, often forming dense 

tussocks, 6-70 (-80) cm tall; rhizome short, 

inconspicuous. Culms erect to ascending, 0.3-1.5 

mm wide, triquetrous, firm but flexible, finely and 

coarsely ribbed, essentially smooth, green with 

larger ribs often pale green, glabrous, antrorsely 

scabrous on angles near apex. Leaves numerous, 

basal and cauline; sheaths short, herbaceous, finely 

veined, green to pale brown, glabrous; ligule a 

thickened narrow band of tissue at adaxial junction 

of sheath and blade; blades narrowly linear, 3-30 

(-70) cm × 0.5-2.5 mm, flattened proximally, V- 

shaped proximally to subplicate distally, attenuate 

to triquetrous apex, finely veined abaxially, 

smooth adaxially with epidermal cells evident, 

margins and midvein beneath essentially smooth 

proximally, antrorsely scabrous distally, green, 

glabrous. Inflorescence a terminal and series of 

1-2 lateral fascicles of spikelets from the upper 

sheathing bracts; fascicles turbinate, 7-16 × 4-23 

mm, the smaller lateral fascicles on slender erect 

to ascending peduncles; spikelets ovoid-ellipsoid, 

3.5-5 × 0.5-1.5 mm, acute to acuminate at apex, 

acute at base, with 4-5 (6) scales; fertile scales 2 

(-3), ovate-elliptic to ovate-lanceolate, 2.5-4 × 1.4- 

2.6 mm, slightly curvate-keeled, dorsally obtuse 

or subrounded, submembranous, finely cellularstriate, 

semi-glossy, whitish to pale rufous, 

glabrous, margins broadly scarious, midcosta 

slender, extended beyond the acute apex as a short 

mucro. Stamens 2, the anthers 1.5-2 mm long, 

apiculate, rounded at base; style 2-branched. 

Achene biconvex, obpyriform, 1.6-1.8 (-2) × 0.9- 

1.2 mm, rounded to a truncate apex, attenuate to 

base, faintly transversely rugulose, yellowish

334 


Fig. 53. A-D. Rhynchospora alba. A. Habit. B. Terminal inflorescence unit. C. Spikelet. D. Achene. E-H. Remirea maritima. E. 

Habit. F. Inflorescence. G. Spikelet. H. Achene. (A, from Proctor 40815; B-D, from Sargent 583; E, from Sargent B16 and 

Curtis 5385; F-H, from Chase s.n., 22 Nov 1913).


brown with a prominent pale medial disc and pale 

margins; style base triangular-lanceolate to 

lanceolate, 0.7-1.3 mm long, 0.3-0.5 mm wide at 

base, flattened, brittle, light brown, often reddish 

dotted or lineolate; bristles 10-12, flattenedsubulate, 

stiff, retrorsely barbed, slightly villous 

at base, pale yellowish brown or reddish, 

exceeding the achene, shorter than to slightly 

exceeding apex of style base. 

General distribution: Canada, United States, 


Distribution in Puerto Rico: Grows on 

pockets of organic soil in exposed rock crevices 

and ledges on the summit of El Yunque, ca. 1100 

m, in the Luquillo Mountains and along road to 

the summit of Cerro de Punta, in the Cordillera 

Central, 1200-1300 m. Naguabo, Ponce, and Río 

Grande. 


Luquillo: Catalina-Yunque Trail, N.L. Britton & 

Bruner 7626 (NY). Ponce: Toro Negro Forest 

Reserve, Axelrod et al. 8874 (UPRRP). Luquillo: 

top of El Yunque Mt., González-Más 1524 (NY, 

UPRRP); Luquillo Mts., El Yunque, González-Más 

& Woodbury 3455 (MAPR). Naguabo: Caribbean 

National Forest, Axelrod & Grose 11223 (UPRRP); 

Sierra de Luquillo, Proctor 40815 (US). Road to 

East Peak, Woodbury s.n. (NY). Río Grande: El 

Yunque, Gleason & Cook X-86 (NY); El Yunque, 

Sargent 583 (US). 

2. Rhynchospora berteroi (Spreng.) C. B. Clarke 

in Urban, Symb. Antill. 2: 119. 1900; 

Hypolytrum berteroi Spreng., Neue Entd. 1: 

241. 1820. Type: Guadeloupe. Bertero s.n. 

(holotype: TO; isotypes: M, MO). 

Schoenus pusillus Sw., Prodr. 20. 1788; 

Dichromena pusilla (Sw.) Kunth, Enum. Pl. 

2: 275. 1837; Rhynchospora pusilla (Sw.) 

Griseb., Abh. Königl. Ges. Wiss. Göttingen 

7: 271. 1857, non Chapman ex M. A. Curtis, 

1847. Type: Jamaica. Swartz s.n. (holotype: 

BM; isotype: M). 

Kyllinga brevifolia Roem. & Schult., Mant. 2: 134. 

1824, non Rottbøll, 1773. Type: Dominican 

Republic. Bertero s.n. (holotype: TO; 

isotypes: M, MO). 

Scleria microdiscus Steud., Syn. Pl. Glumac. 2: 

175. 1855. Type: Puerto Rico. Herb. Wydler 

s.n. (holotype: G). 

Dichromena filiformis C. B. Clarke in Urban, 

Symb. Antill. 2: 102. 1900; Rhynchospora 

subfiliformis H. Pfeiff., Repert Spec. Nov. 

Regni Veg. 49: 80. 1940, as a new name for 

Dichromena filiformis C. B. Clarke, non 

Rhynchospora filiformis Vahl, 1805; 

Rhynchospora nervosa var. subfiliformis (H. 

Pfeiff.) Kük., Bot. Jahrb. Syst. 75: 295. 1951. 

Type: Hispaniola. Poiteau s.n. (holotype: G). 

Stoloniferous perennial, 0.6-13 cm tall; 

stolons slender, 0.2-1 mm thick, stramineous to 

reddish brown; roots fine. Culms tenuous, erect 

to arching, 0.2-0.3 mm wide, obtusely triquetrous, 

delicate, finely ribbed, smooth, green, glabrous. 

Leaves 3-10 per culm, ascending to arching, basal 

and lower cauline; sheaths short, herbaceous, 

finely veined, pale green, glabrous to sparsely 

hirtellus; ligule absent; blades narrowly linear, 

(5-) 10-25 cm × (0.3-) 0.5-2 mm, flattened to 

subinvolute, attenuate to triquetrous tip, finely 

veined abaxially, finely cellular-reticulateroughened 

adaxially, long pilose-ciliate along 

margins, antrorsely scabrous on margins and 

abaxial midvein (at least distally), green, glabrous 

or sometimes hirtellus or sparsely so adaxially or 

abaxially (at least proximally). Inflorescence a 

small fascicle of 2-6 congested spikelets at the 

summit of the culm; involucral bracts leaf-like but 

reduced, often with scarious margins proximally, 

ciliate-scabrous (at least distally), the lowermost 

equaling or slightly overtopping the inflorescence, 

to 20 mm long; fascicles 4-8 × 2-5 mm; spikelets 

ovoid, 2.5-5 × 0.8-1.8 mm, acute to obtuse at apex, 

cuneate to sub-rounded at base, the scales 6-8 

(-10), spreading with maturing achenes; fertile 

scales 5-6 (-8), ovate to ovate-lanceolate, 2-3.2 × 

1-1.8 mm, boat-shaped, dorsally obtuse to 

rounded, thinly herbaceous to submembranous, 

white to dingy white, margins scarious, entire, 

midcosta fine, indistinct proximally, light green 

distally, extending only slightly beyond acute apex 

as short mucro. Stamens 3, the anthers 0.6-1 mm 

long, with a dark triangular-papillate appendage 

at apex, sub-rounded and minutely papillate at 

base; style 2-branched. Achene biconvex, obovate 

to broadly obovate, or somewhat obpyriform, 0.9- 

1 × 0.8-0.9 (-1) mm, truncate to slightly arching 

at apex, subabruptly narrowed at base, transversely 

rugulose, stramineous to orangish brown, 

sometimes with dark brown to blackish brown

336 


rugae; style base triangular to shallowly triangular, 

0.2-0.6 mm long, 0.5-0.7 mm wide at base, with 

concave angles, subflattened to thickened, 

crustose, dull to semi-glossy, light brown to brown 

or blackish; bristles absent. 


Guadeloupe, and Belize. 


Islands: On serpentine and limestone soils in often 

shaded, secondary forests; forested hillsides and 

slopes, banks, rocky areas, cliffs, ridges, and road 

banks at lower to middle elevations. Arecibo, 

Bayamón, Cabo Rojo, Dorado, Florida, Hatillo, 

Isabela, Lares, Maricao, Mayagüez, Rincón, 

Sabana Grande, San Germán, San Juan, San 

Sebastián, Toa Baja, Utuado, Vega Alta, and Vega 

Baja; St. Thomas. 


Arecibo: Tanamá, Reserva Mata de Platano, 

Ackerman 3414 (UPRRP); Río Abajo, Liogier et 

al. 30827 (NY). Río Tanamá, Stevens 7865 (NY). 

Bayamón: Liogier 10200 (US). Isabela: Bo. 

Planas, Guajataca Forest, Axelrod 7172 (UPRRP, 

US). Lares: Stevens & Hess 4946 (NY). Maricao: 

Maricao Forest, González-Más 1995 (MAPR). 

Mayagüez: Cerro Las Mesas, N.L. Britton & 

Brown 3878 (NY, US); Guanajibo, near 

Mayagüez, N.L. Britton et al. 4336 (NY, US). 

Rincón: Punta Higuero, González-Más & Seda 

2689 (MAPR).Sabana Grande: Susúa Forest 

Reserve, González-Más 2030 (MAPR). San Juan: 

Martín Peña, Stevenson 3726 (NY, US). San 

Sebastián: Cidial, González-Más 928 (MAPR, 

NY). Utuado: Stevens & Hess 4699 (NY). Vega 

Alta: Quebrada Arenas, González-Más 1114 

(MAPR, NY, US). Vega Baja: Rt. 644, ca. 1.5 mi. 

from Rt. 155, Taylor 8859 (UPRRP). 

3. Rhynchospora brachychaeta C. Wright in 

Sauvalle, Anales Acad. Ci. Méd. Habana 8: 

85. 1871. Type: Cuba; Pinar del Río. Wright 

3782, (holotype: GH; isotypes: NY!, US!). 

Rhynchospora blauneri Britton, Bull. Torrey Bot. 

Club 50: 56. 1923. Type: Puerto Rico; Sierra 

de Luquillo. Blauner 247 (holotype: NY!). 


rhizome short, creeping, the culms spaced at 

intervals along the rhizome. Culms ascending, 3- 

5 mm wide, filiform, obtusely trigonous to 

subterete, finely ribbed, firm, glabrous, green. 

Leaves 2-4 (per culm), primarily basal; sheaths 

elongate, finely ribbed, green, glabrous; ligule 

absent; blades linear-filiform, often capillary, 10- 

20 cm × 0.3-0.8 mm, involute proximally, 

crescentform-capillary to terete distally, antrorsely 

scabrous on margins distally, green, glabrous, 

attenuate to apex. Inflorescence a terminal and 1- 

3 small lateral partial obtriangular or narrowly 

elliptic panicles of fascicled spikelets from the 

upper sheathing bracts; panicles 6-10 × 4-10 mm; 

spikelets narrowly ovoid-lanceoloid, 3-3.2 × 

(0.6-) 0.7-1.2 mm, with acuminate apex, cuneate 

at base, 10-25 per partial panicle, with 3-4 scales; 

fertile scale ovate, 1, 2.1-3 × 1.9-2.2 mm, clawed 

at base, inrolled around flower and developing 

achene, reddish brown or brown, glabrous, with a 

single pale narrow midcosta prolonged beyond 

apex as a 0.2-0.3 mm long mucro, lateral veins 

indistinct, margins entire, inrolled in fertile scales, 

the apex acuminate. Stamens 3, or sometimes 1-2 

abortive, the anthers 1.5-2 mm long; style 2branched. 

Achene biconvex with bulging center, 

obovate, 1.2-1.4 × 1-1.1 mm, faintly cellularreticulate 

and brownish on sides, the turgid center 

smooth, light brown to yellowish brown; style base 

triangular to narrowly triangular, 0.5-0.7 mm long, 

0.5-0.7 mm wide at base, pale brown; bristles 1-4, 

short to rudimentary, antrorsely barbed. 


States (Alabama, Florida, and Mississippi), Belize, 

Nicaragua, Greater Antilles, and Guyana. 

Distribution in Puerto Rico: known only from 

a single collection made in the Sierra de Luquillo 

Mountains by Blauner (247) in 1852-53. 

4. Rhynchospora breviseta (Gale) Channell, 

Rhodora 58: 336. 1956. Rhynchospora 

oligantha var. breviseta Gale, Rhodora 46: 

129. 1944. Type: United States; Florida. 

Curtiss 5687 (holotype: GH; isotypes: NC, 

US!). 

Rhynchospora oligantha sensu Kükenthal, Repert. 

Spec. Nov. Regni Veg. 23: 207. 1926, non A. 

Gray, 1835. 


short, firm or ligneous, somewhat bulbous, crownlike; 

roots fine; sheathing base to 1 mm wide. 

Culms erect or weakly ascending, 0.2-0.5 (-0.7) 

mm wide, obtusely trigonous, capillary, delicate, 

finely ribbed, essentially smooth, pale green,


glabrous. Leaves numerous, ascending, basal and 

lower cauline; sheaths short, herbaceous, finely 

veined, pale green, inner band yellowish brown at 

orifice, glabrous; ligule absent or sometimes 

merely a narrow thickened band of tissue at adaxial 

junction of sheath and blade; blades filiformsetaceous, 

7-45 cm × 0.2-0.5 mm, shallowly 

canaliculate adaxially, long-attenuate with a 

constriction towards apex, subflattened above this 

with an acute to acuminate tip which is antrorsely 

scabrous on margins, finely veined, cellularreticulate 

adaxially, smooth, green, glabrous. 

Inflorescence a single spikelet or small 

corymbiform panicle of 2-3 spikelets subtended 

at base by an elongated involucral bract appearing 

as a continuation of the culm, rarely a remote 

lateral partial panicle present, the branching often 

simple; spikelet pedicels flattened to flattenedtrigonous, 

ciliate on margins; spikelets 

subflattened-fusiform, becoming turgid and 

torulose with developing achenes, 7-9.5 (-10) × 

(1.4-) 1.8-3 mm, acuminate at both apex and base, 

the rachis anfractuose-elongate, spirally zig-zag 

above sterile basal scales, with 7-10 scales; fertile 

scales 3-8, widely ovate-elliptic, 4.3-6 × 3-4 mm, 

curvate-keeled, dorsally obtuse to rounded, 

abruptly narrowed proximally to a claw-like base, 

thinly herbaceous, finely cellular-striate, dull to 

semi-glossy, cinnamon-brown, margins narrowly 

scarious, undulate, midcosta fine, pale green or 

yellowish green to brown distally, extended 

beyond narrowly acute apex as a short mucro. 

Stamens 3, the anthers typically 0.4-0.8 mm long, 

triangular-apiculate, truncate at base; style 2branched. 

Achene biconvex, obovate, 1.8-2.2 × 

1.3-1.7 mm, rounded at apex, cuneate at base, 

transversely rugulose with faint longitudinally 

striae, variable in color, light brown, dark brown, 

dark blue, bluish black, or charcoal black, often 

with a whitish or bluish powdery bloom; style base 

short-conic or depressed-conic, abruptly 

contracted at base, 0.3-0.5 mm long, 0.6-0.7 mm 

wide at base, subflattened above base, brown; 

bristles 6, plumose to apex or sometimes antrorsely 

barbed at tip, silvery brown, short, ½ length of 

achene or shorter. 


States and the Greater Antilles. 

Distribution in Puerto Rico: High elevation 

montane rock outcrops and wet sands of coastal 

plain savanna. Known only from Luquillo 

Mountains (El Yunque and East Peaks) and Laguna 

Tortuguero and vicinity. Ceiba, Dorado, Manatí, 

Río Grande, and Vega Baja. 


Dorado: Woodbury s.n. (UPR). Luquillo: El 

Yunque, East Peak Road, Woodbury s.n. (NY, 

UPR). East Peak area, Woodbury s.n. (UPR). 

Manatí: Tortuguero, Woodbury s.n. (NY, UPR). 

5. Rhynchospora ciliata (G. Mey.) Kük., Bot. 

Jahrb. Syst. 56 (Beibl. 125): 16. 1921; 

Dichromena ciliata Vahl, Enum. Pl. 2: 240. 

Oct.-Dec. 1805, nom. illeg., non Dichromena 

[as ‘Dichroma’] ciliatum Pers., Apr.-Jun. 

1805, nom. illeg.; Schoenus ciliatus G. Mey., 

Prim. Fl. Esseq. 23. 1818; Rhynchospora 

vahliana Griseb., Fl. Brit. W. I. 577. 1864, 

nom. illeg.; Rhynchospora nervosa var. ciliata 

(G. Mey.) Kük., Bot. Jahrb. Syst. 75: 295. 

1951; Rhynchospora nervosa subsp. ciliata 

(G. Mey.) T. Koyama, Madroño 20: 254. 1970, 

non Rhynchospora ciliaris [ as ‘ciliata’] 

(Michx.) Vahl, Enum. Pl. 2: 235. 1805. [Vahl’s 

incorrect publication of Rhynchospora ciliata, 

based on Schoenus ciliaris Michx., should be 

interpreted as an orthographic variant]. 

Lectotype: Puerto Rico. West s.n. (C-Vahl), 

here designated. 

Dichromena pura Nees in Martius, Fl. Bras. 2: 

112. 1842; Rhynchospora pura (Nees) 

Griseb., Fl. Brit. W. I. 577. 1864. Type: St. 

Vincent. Lindley s.n. (holotype: CGE; isotype: 

NY). 

Fig. 54. A-D; 65. H 


rhizomes short; roots medium-sized, to 1 mm 

thick. Culms ascending, (0.7-) 1-1.7 (-2) mm wide, 

obtusely trigonous or shallowly triquetrous, finely 

ribbed, often indistinctly so, green, glabrous 

distally, often hirsute proximally; sheathing base 

of culms and lower nodes 1-4 mm wide. Leaves 

5-15 per culm, ascending, basal and lower cauline; 

sheaths short, loose, finely veined, light green to 

straw-colored, whitened at base, hirsute to 

glabrous; ligule absent; blades linear, 8-65 cm × 

1.5-4.5 mm, flattened, often folded or margins 

subinvolute, acuminate at apex, finely and 

distinctly veined abaxially, essentially glabrous, 

green, finely cellular-reticulate roughened 

adaxially, glabrous or hirsute proximally, whitish

338 


Fig. 54. A-D. Rhynchospora ciliata. A. Habit, and detail of abaxial side of leaf blade. B. Inflorescence. C. Flower. D. Achene. E- 

J. Scleria lithosperma. E. Habit. F. Detail of culm showing summit of sheaths with contraligules. G. Inflorescence branch. H. 

Pistillate scales and pistil. I. Stamen and staminate scale. J. Achene. From Acevedo-Rdgz., P. 1996, Flora of St. John, Mem. New 

York Bot. Gard. 78.


or pale green, margins and abaxial midvein 

antrorsely scabrous, long-ciliate proximally. 

Inflorescence a glomerate, hemispherical head of 

5-16 spikelets at the summit of the culm, 10-17 

(-20) mm in diam.; involucral bracts 7-8, leaf-like, 

longer than the inflorescence, white on adaxial 

surface basally, green and glabrous to hirsute 

abaxially at base, the margins ciliate proximally; 

spikelets ovate to widely-ovate, 5-10 × 2-4 mm, 

acute to narrowly acute at apex, rounded at base, 

with 10-35 scales; scales all fertile except for basal 

scale of spikelet, ovate, 3.5-5 × 2-3 mm, broadly 

boat-shaped, curvate, slightly keeled distally, 

dorsally obtuse to rounded, glabrous, whitish, 

frequently red-lineolate, midcosta narrow, pale, 

green distally, extending beyond the acute, obtuse, 

rounded or emarginate apex as a short, straight or 

recurved mucro. Stamens 3, the anthers (1.7-) 2-3 

mm long, bluntly apiculate, truncate at base with 

minute crystalline lobes or papillae; style 2branched. 

Achene biconvex, turgid, obovate to 

depressed-obovate, 1.3-1.8 × 1.1-1.6 mm, rounded 

to broadly rounded at apex forming a very fine 

rim around base of style, short-cuneate at base, 

transversely rugulose, shiny, stramineous to dark 

yellowish brown at maturity; style base triangular 

with concave margins that are shortly decurrent 

on shoulders of achene, 0.5-0.8 mm long, about 

as wide as the achene, rounded or obtuse at tip, 

whitish to light brown or brown at maturity; 

bristles absent. 




Islands: In sandy or clayish soils; wet places, 

roadsides, grassy areas, lawns, fields, pastures, 

forest on serpentine, and dry coastal forests. 

Adjuntas, Aguas Buenas, Arecibo, Bayamón, Cabo 

Rojo, Caguas, Canóvanas, Carolina, Cayey, 

Comerío, Fajardo, Guayama, Guaynabo, 

Humacao, Isabela, Lares, Luquillo, Maricao, 

Mayagüez, Moca, Naguabo, Orocovis, Peñuelas, 

Río Grande, Sabana Grande, Salinas, San Germán, 

San Juan, San Lorenzo, San Sebastián, Trujillo 

Alto, Utuado, Vega Baja, Vieques, Yabucoa, and 

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

Common names: Puerto Rico: Botoncillo, 

Coquí blanco, Yerba de estrella. 

Note: In a revision of Rhynchospora section 

Dichromena by Thomas (1984), R. ciliata was 

treated as a subspecies of Rhynchospora nervosa 

(Vahl) Boeck. However, cytogenetic studies by 

Luceño et al. (Canad. J. Bot. 76: 440-449. 1998) 

indicate that R. ciliata should be treated as a 

distinct species from R. nervosa. 


Adjuntas: road 10, km 9, González-Más 865 (NY, 

US). Aguas Buenas: Bo. Sumidero, González-Más 

1176 (MAPR, NY, US). Arecibo: Río Abajo State 

Forest, Acevedo-Rdgz. 10580 (US). Bayamón: 

Johnston 1023 (US); Underwood & Griggs 897 

(NY, US). Cabo Rojo: Joyuda, Hess 15 (MAPR). 


Condesa Section, Axelrod et al. 1765 (UPRRP). 

Carolina: 204 St. Country Club, Ortega 120 

(UPRRP). Fajardo: S of Luquillo on Rt. 988, 

Taylor 7706, 7709 (UPRRP). Guayama: Sierra de 

Cayey, Bo. Carite, Ackerman 3468 (UPRRP). 

Guaynabo: Bo. Mamey, Axelrod & Sastre 6106 

(UPRRP). Isabela: Rt. 446, ca. 1-2 km N of 

boundary of Bosque Guajataca, Taylor & Taylor 

8926 (UPRRP). Luquillo: Road 988, Km 12.5, 

Bisbal 1 (UPRRP). Maricao: Road 120, González- 

Más 433 (MAPR); Maricao: Cordillera Central, S 

of Maricao on Rt. 120, Grimes et al. 3254 (NY). 

Mayagüez: Las Mesas, Holm 55 (US); Holm 108 

(MO). Moca: Road 110, km 9.2, González-Más 

1809 (MAPR). Naguabo: Bo. Florida, Road 191, 

km 27.5, González-Más 1387 (MAPR); Sierra de 

Naguabo, Río Prieto and adjacent hills, Shafer 

3672 (NY, US). Orocovis: Bo. Barros, Axelrod & 

Rogowitz 9989 (UPRRP). Peñuelas: Escuela 

Adolfo Grana, Fraticelli 18 (MAPR). Río Grande: 

Along Rt. 186, bordering Carribean National 

Forest, Kelloff et al. 364 (US). San Juan: Santurce, 

Heller & Heller 11 (NY, US); Río Piedras, Barrett 

211 (MAPR, US). San Germán: Sargent 488 (US). 

San Lorenzo-Yabucoa border, Taylor 7603 (NY, 

UPRRP). San Sebastián: Bo Cidral, Road 111, 

González-Más 910 (MAPR). Trujillo Alto: road 

172, González-Más 1562 (NY). Utuado: Road 

from Utuado to Lares, Underwood & Griggs 86 

(NY, US). Vieques: Isabel Segunda to Santa Maria, 

Shafer 2611 (NY, US). Yabucoa: Sierra de 

Yabucoa, N.L. Britton et al. 6248 (NY). Yauco: 

Bo. Susua Alta, Bosque Estatal de Susúa, García 

& Caminero 3310 (MAPR). ST. JOHN: Bordeaux, 

N.L. Britton & Shafer 562 (US); Lameshur, 

Woodbury 226/6693 (VINPS). ST. THOMAS: Crown, 

N.L. Britton & Marble 1346 (US); Raunkiaer s.n. 

(US); Eggers 369 (NA).

340 


6. Rhynchospora colorata (L.) H. Pfeiff., Repert. 

Spec. Nov. Regni Veg. 38: 89. 1935; Schoenus 

coloratus L., Sp. Pl. 43. 1753; Schoenus 

stellatus Lam., Encycl. 1: 741. 1784, nom. 

illeg.; Dichroma ciliatum Pers., Syn. Pl. 1: 

58. 1805, nom. illeg.; Rhynchospora stellata 

Griseb., Abh. Koenigl. Ges. Wiss. Göttingen 

8: 271. 1857; Dichromena colorata (L.) 

Hitchc., Rep. (Annual) Missouri Bot. Gard. 

4: 141. 1893. Lectotype: Jamaica. Sloane, Voy, 

Jamaica 1: t. 78, f. 1. 1707, designated by 

W.W. Thomas, Mem. New York Bot. Gard. 

37: 83. 1984. 

Dichromena leucocephala Michx., Fl. Bor.-Amer. 

1:37. 1803; Rhynchospora leucocephala 

(Michx.) Boeck., Vidensk. Meddel. Dansk 

Naturhist. Føren. Kjøbenhavn ser. 3, 1: 144. 

1869. Type: United States; Carolina and 

Georgia. Bosc s.n. (holotype: P; isotype: G). 


rhizomes producing slender, scaly stolons up to 

20 cm long, 1-2 mm thick; roots medium-sized, 

to 1 mm thick; sheathing base of culm 3-6 mm 

wide. Culms solitary, erect, obtusely trigonous 

proximally to obtusely triquetrous distally, 0.7-2 

mm wide, very firm but flexuous, finely ribbed, 

smooth, pale green, glabrous. Leaves 3-11, 

ascending, basal and lower cauline; sheaths short, 

herbaceous, finely veined, pale green, glabrous; 

ligule absent; blades linear, 3-56 (-60) cm × 

(0.5-) 1-4 mm, flattened to margins incurved, 

attenuate to triquetrous apex, finely veined 

abaxially, very finely cellular-reticulate and 

essentially smooth adaxially, veins not evident, 

margins and abaxial midvein essentially smooth 

proximally, antrorsely scabrous distally, green to 

yellowish green, glabrous. Inflorescence a 

congested hemispherical to globose head of (5-) 

10-50 spikelets at the summit of the culm; 

involucral bracts (3-) 4-7 (-8), linear, exceeding 

inflorescence head, green distally, white both 

adaxially and abaxially basally, the lowermost to 

20 cm long; head 7-16 (-18) mm in diam.; spikelets 

ovoid to ovoid-ellipsoid, (3-) 5-9 × 1.5-2.5 mm, 

acute to obtuse at apex, cuneate to sub-rounded at 

base, with 7-30 scales; scales all fertile except for 

basal scale of spikelet, ovate to oblong-ovate, 

(2.5-) 3-4.5 × 2-4 mm, boat-shaped, dorsally acute 

to more commonly obtuse or rounded, thinly 

herbaceous, dull, white to dingy white, reddish 

lineolate proximally, glabrous, margins entire or 

narrowly scarious, midcosta indistinct, evident 

only at tip, not prolonged beyond acute to obtuse 

apex. Stamens 3, the anthers 1-2 mm long, 

apiculate, truncate at base with crystalline papillae; 

style 2-branched. Achene biconvex, obovate to 

widely obovate, 0.9-1.2 (-1.5) × 0.9-1.4 mm, 

truncate to slightly arched at apex, rounded at base, 

transversely rugulose, often with an indistinct 

cellular-reticulate margin, glistening, stramineous 

to dark brown or dark brown; style base triangular 

to shallowly triangular, 0.3-0.7 mm long, 0.8-1.2 

mm wide at base, broadly and shortly attenuate to 

obtuse apex, subflattened, crustose, whitish to 

grayish brown or brown; bristles absent. 


States west to Texas, Mexico, Central America, 

West Indies, and South America. 

Distribution in Puerto Rico: Wet to moist 

sandy soils of open marshlands, swampy areas, 

moist swales, pastures, drainage canals, open 

savannas, sandy thickets, second-growth coastal 

forest, and roadside ditches. Aguada, Añasco, 

Arecibo, Cataño, Dorado, Guánica, Guayama, 

Humacao, Luquillo, Manatí, Ponce, Río Grande, 

Toa Baja, Utuado, and Vega Baja. 


Añasco: Sargent 639 (US). Arecibo: Río Abajo 


Cataño: Millspaugh 152 (NY). Guánica: Guánica 

Lake, Sargent 38 (US). Guayama: Algarrobo, 

Stevenson 2128 (US). Luquillo: Bo. Mata de 

Platano, Axelrod & Pérez 6569 (UPRRP). Manatí: 

Bo. Tierras Nuevas Saliente, Axelrod & Sastre 

6467 (UPRRP, US). Río Grande: Río Espíritu 

Santo, Coco Beach, Liogier et al. 33789 (NY); 

Punta Comején, Liogier et al. 29203 (NY). Toa 

Baja: Bo. Sabana Seca, Road 165, Km 16, 

Meléndez & Moya 83 (UPRRP). Vega Baja: at the 

E end of Laguna Tortuguero along Rd. 687, Taylor 

7647 (NY, UPRRP). 

7. Rhynchospora contracta (Nees) J. Raynal, 

Adansonia Ser. 2. 17: 277. 1978; 

Haloschoenus contractus Nees in Martius, Fl. 

Bras. 2(1): 123. 1842; Dichromena contracta 

(Nees) Steud., Syn. Pl. Glumac. 2: 136. 1855; 

Rhynchospora micrantha var. contracta 

(Nees) Kük., Bot. Jahrb. Syst. 75: 276. 1951. 

Type: Brazil; Piauí. Martius s.n. (holotype: 

M).


Juncus parviflorus Poir. in Lamarck, Encycl. 

Suppl. 3: 160. 1813, non Ehrhart, 1791. Type: 

Puerto Rico. Ledrú 11 (holotype: P; isotype: 

P-2). 

Scleria micrantha Poir. in Lamarck, Encycl. Suppl. 

5: 108. 1817. Type: Puerto Rico. Ledrú s.n. 


Rhynchospora micrantha sensu C. B. Clarke in 

Urb., Symb. Antill. 2: 117. 1900, non Vahl, 

1805. 

Caespitose annual, 4.5-45 cm tall. Culms 

ascending, 0.4-0.8 mm wide, triquetrous to 

compressed-trigonous, soft, finely ribbed, smooth, 

green, glabrous. Leaves numerous, basal and 

cauline, the upper cauline ones representing leaflike 

bracts subtending inflorescence corymbs; 

sheaths short, inflated, finely veined, green to pale 

green, glabrous; ligule absent; blades ascending, 

narrowly linear, flattened, (2-) 3-25 cm × (0.4-) 

0.5-2.2 mm, green, glabrous, finely veined, veins 

scarcely evident adaxially, distinct abaxially and 

pale green, margins antrorsely scabrous distally, 

smooth proximally, midvein antrorsely scabrous 

distally on abaxial side. Inflorescence a terminal 

and series of 2-5 lateral compound corymbs with 

ascending to divergent or reflexed branches from 

the upper sheathing bracts; terminal corymb 

widely ovoid to depressed-ovoid, 8-60 × 11-75 

mm, lateral corymbs in same size range as terminal 

except for the lowermost one which is often 

reduced; spikelets small, ovate-elliptic with acute 

apex, 1.5-2 × 0.5-0.7 mm, the scales 5-8, widely 

spreading with developing achenes; fertile scales 

4-5, 1-1.3 × 0.6-1.2 mm, ovate to widely ovate, 

dorsally obtuse to rounded on mature scales, acute 

on immature scales, submembranous, 

subtranslucent, brown with minute reddish 

lineations, acute to subacuminate at apex, midcosta 

fine, antrorsely scabrous distally, pale brown, 

extending as a short mucro below scale apex, not 

prolonged beyond apex. Stamens 2, the anthers 

small, oblong-elliptic, 0.3-0.5 mm long; style 2branched. 

Achene broadly biconvex to nearly 

rounded in cross section, 0.8-1 × 0.9-1 mm, 

depressed-obovate, broadly rounded at apex, subrounded 

at base with short stipitate cellular 

reticulate base, surface rugose, stramineous or 

light brown to blackish brown at maturity, margins 

costate; style base compressed proximally, as wide 

as achene body, forming 2 strap-like lobes that 

are decurrent on shoulders of achene body to about 

the middle, abruptly narrowing distally into a short 

triangular-lanceolate tip, 0.3-0.4 mm long; bristles 

absent. 


America, West Indies, tropical South America, and 


Distribution in Puerto Rico: On red clay or 

sandy soils in moist to wet or sometimes dry 

habitats, marshy areas, clearings, grassy areas, 

seepage bogs, gravelly areas, dry scrub forest, and 

along trails. Bayamón, Dorado, Humacao, Lares, 

Luquillo, Mayagüez, Moca, Maricao, Naguabo, 

San Juan, and Yabucoa. 


Bayamón: Sintenis 1211 (US). Dorado: Johnston 

887 (NY). Humacao: Bo. Candelero Abajo, 

vicinity of Palmas del Mar, Proctor & Rivera 

46705 (US). Lares: Sargent 3258 (US). Luquillo: 

Bo. Pitahaya, Otero 524 (MO). Maricao: Bo. 

Indiera Fría, mountain ridge E of Río Postrero, 

Proctor et al. 45213 (US). Mayagüez: Sintenis 107 

(US); Guanajibo, near Mayagüez, N.L. Britton & 

Cowell 4077 (MO, NY). Moca: Rt. 110, km 10.8, 

González-Más 1790 (MAPR, NY). Naguabo: 

Sierra de Naguabo, Loma Icaco, Shafer 3399 (NY, 

US). San Juan: Río Piedras, Stevenson 1218 (US). 

Yabucoa: in arenosis ad Punta del Naranjo, Sintenis 

5301 (MO, NY, US). 

8. Rhynchospora corymbosa (L.) Britton, Trans. 

New York Acad. Sci. 11: 84. 1892; Scirpus 

corymbosus L., Cent. Pl. 2: 7. 1756. 

Lectotype: India. Collector unknown. (LINN- 

71.48), designated by Gordon-Gray, Strelitzia 

2 : 150. 1995. 

Rhynchospora aurea Vahl, Enum. Pl. 2: 229. 1805. 

Type: India. Koenig s.n. (holotype: C-Vahl). 

Robust, coarse, rhizomatous perennial, 60- 

200 cm tall; rhizome short, stout, hardened. Culm 

ascending, 2-8 (-11) mm wide, trigonous with 

plane sides to triquetrous, hardened and stiff, finely 

ribbed, finely antrorsely scabrous, with 

subappressed barbs on marginal ribs, pale green 

to green, glabrous, the edges of the channel 

antrorsely scabrous. Leaves 5-15 per culm (not 

including upper leaf-like bracts subtending 

inflorescence corymbs), arched-ascending, 

spreading, basal and cauline; sheaths elongate, 

inflated distally, often spongy-thickened on basal

342 


leaves, herbaceous, substiffened, finely veined, 

antrorsely scabrous along margins on cauline 

sheaths, smooth on basal sheaths, pale green or 

sometimes tinged with brown, glabrous; ligule 

absent; blades linear, 40-190 cm × 8-23 mm, 1costate, 

herbaceous to substiffened, finely manyveined, 

essentially smooth, green, glabrous, 

margins and midvein beneath antrorsely scabrous, 

the apex long-acuminate, triquetrous. 

Inflorescence a terminal and a series of 2-5 

corymbose lateral partial panicles from the upper 

sheathing bracts; terminal corymb depressed 

obovoid, generally wider than long, 6-21 × 7.5- 

17.5 cm, generally the largest, the others gradually 

reduced below; spikelets many, in hemispherical 

fascicles at branch tips, ovoid-ellipsoid or 

ellipsoid, (4.5-) 5-7 (-7.5) × (1-) 1.2-2 mm, acute 

to acuminate at both base and apex, with 5-8 

scales; fertile scales 2-3, ovate-elliptic to widely 

ovate-elliptic or linear-elliptic (terminal scale), 4- 

5.6 × 1.8-3.8 mm, thin, submembranous, dorsally 

obtuse to rounded at maturity, upper sterile and 

lower fertile scales inrolled at anthesis, finely 

cellular-striate, uniformly reddish brown with very 

fine red lineations, glabrous, margins narrowly 

scarious, midcosta fine, pale brown to brown, 

extending beyond the acute to acuminate apex as 

a short mucro on proximal spikelet scales, 

antrorsely scabrous distally near tip; Stamens 3, 

the anthers 2.3-3.5 mm long; style entire to shortly 

bi-lobed at tip. Achene biconvex, obtrullate or 

obovate, 2.3-3.3 × 1.7-2 mm, transversely 

rugulose, papillate, brown to dark brown, dull, 

cuneate to base, subacute at apex, the margins 

thickened with constrictions and protuberances; 

style base slightly wider and thicker than achene 

body, triangular with acuminate apex and 2-lobed 

sagittate base, 3-4.3 mm long, 1.7-2.1 mm wide 

at base, finely antrorsely scabrous (at least distally) 

spongy-thickened, brown, semi-glossy, with 

longitudinal medial groove on both sides that is 

often whitish at bottom of declivity; bristles 6, fine, 

subulate, up to 5 mm long, typically exceeding 

achene body and 2/3 length of style base, but 

sometimes 1-2 shorter than achene body, reddish, 

finely antrorsely barbed with blackish barbs. 


Distribution in Puerto Rico: In wet swampy 

and marshy grounds or pastures at both low and 

high elevations. Arecibo, Bayamón, Cidra, 

Dorado, Mayagüez, Naguabo, San Juan, Toa Baja, 

and Utuado. 

Common name: Puerto Rico: Junco pico. 


Arecibo; Bosque de Río Abajo, Los Puercos, 

Acevedo-Rdgz. 155 (SJ); Río Abajo, Liogier et al. 

30823 (NY). Bayamón: Bo. Sabana Seca, Otero 

537 (MO); Sintenis 1241 (US); Bayamón to 

Cataño, N.L. Britton et al. 2901 (US). Cidra: along 

railroad near Pueblo Viejo, Heller 6398 (MO, NY, 

US); Dorado: Bo. San Antonio, González-Más 350 

(MAPR). Mayagüez: Stevens 713 (MAPR, NY). 

Naguabo: Bo. Río Blanco, Axelrod & Thomas 

11022 (UPRRP). Stevenson 2094 (NY). San Juan: 

Río Piedras, Stevenson 1290 (US). Toa Baja: road 

866, km 7.2 to Punta Salinas, González-Más 1145 

(MAPR, NY). Utuado: Sargent 3275 (US). 

9. Rhynchospora depressirostris M.T. Strong, 

Brittonia 52: 241. 2000. Type: Puerto Rico; 

Dorado. Stevenson 889 (holotype: US!; 

isotypes: NY!, UPR!). 

Rhynchospora tenuis sensu Britton & P. Wilson, 

Bot. Porto Rico 5: 102. 1923, non Willdenow 

ex Link, 1820. 

Fig. 55. A-E 


rhizome short, the culms and numerous basal 

leaves forming dense clumps. Culms ascending, 

0.4-0.9 mm wide, obtusely trigonous to 

compressed-trigonous or subterete, soft, flexuous, 

finely ribbed, pale green, glabrous. Leaves 

ascending, numerous, primarily basal, 1-3 cauline; 

sheaths quite long, particularly cauline ones, 

closely clasping culm, herbaceous, finely ribbed, 

pale brown to stramineous proximally, glabrous; 

ligule essentially absent, but sometimes a faint rim 

of tissue present at adaxial junction of sheath and 

blade; blades narrowly linear, 6-40 cm × 0.3-1.1 

mm (unfolded), V-shaped to subfolded or 

sometimes involute, firm, herbaceous, finely 

veined abaxially, smooth, glossy, and minutely 

cellular-reticulate adaxially, light green or pale 

brown abaxially, green adaxially, glabrous, 

margins smooth proximally, antrorsely scabrous 

distally, the apex long-acuminate, triquetrous near 

tip. Inflorescence 1-3 (4) corymbose partial 

panicles from the upper sheathing bracts, panicles 

somewhat strict, the terminal panicle 2-7 × 1-1.7 

cm, with 10-50 spikelets; sheathing bracts shorter 

and a little narrower than leaf blades, to 20 cm 

long; spikelets ovoid-ellipsoid, 2.5-3.2 × 0.6-1


mm, narrowly acute to acuminate at apex, 

becoming falcate with age, the scales 5-7, 

spreading with maturing achenes; fertile scales 3- 

4, ovate-elliptic or widely ovate-elliptic to ovatelanceolate, 

1.6-2.7 × 0.9-1.4 mm, dorsally obtuse 

to rounded, herbaceous, distal scales of spikelet 

thinly herbaceous to submembranous, minutely 

cellular-striate, semi-glossy, reddish brown with 

slightly darker brown lineations, glabrous, margins 

narrowly scarious, slightly crisped, midcosta very 

fine, indistinct except at apex, prolonged beyond 

narrowly acute to acuminate apex as a short mucro, 

lateral nerves indistinct. Stamens 3, the anthers 

0.8-1.2 mm long, minutely lobed at base, 

glandular-apiculate; style 2-branched. Achene 

biconvex, rounded-obovate, 0.7-0.9 × 0.8-1 mm, 

slightly narrowed to truncate or sub-rounded at 

short stipitate base, truncate at apex, transversely 

rugose, narrowly cellular-reticulate along margins 

and at base, shining, glossy, red to reddish brown 

or brown; style base extremely depressedtrigonous, 

oval-shaped from top view, not lobed, 

to 0.2 mm long, often 0.1 mm long or less, 0.5-0.7 

mm wide at base, brittle, dark brown or blackish; 



Distribution in Puerto Rico: White silica sand 

areas in the vicinity of Laguna Tortuguero and 

Dorado and at Punta Guayanes; Cataño, Dorado, 

Vega Baja, and Yabucoa. 

Note: Rhynchospora depressirostris is closely 

related to R. tenuis Link and has been treated as 

that species by Britton & P. Wilson (1923), 

González-Más (1964), and Liogier & Martorell 

(1982: 220; 2000: 245). However, it consistently 

differs from the latter species in the style base and 

achene morphology. The type of R. tenuis Link is 

a plant from South America and has a style base 

that is shallowly triangular, 0.2-0.6 mm long, and 

2-lobed at base, the lobes decurrent on shoulders 

of achene; and the achene is obovate and has a 

surface that is finely rugose or rugulose with 6-8 

rugae per face. In R. depressirostris, the style base 

is extremely depressed-trigonous, 0.1 (-0.2) mm 

long, with an oval-shaped (from top view), 

unlobed base that is not decurrent on shoulders of 

achene; and the achene is rounded-obovate to 

depressed-obovate and has a rugose surface (more 

coarsely wrinkled than R. tenuis) with only 3-4 

(-5) coarse rugae per face. 


Cataño: Bo. Palmas, Axelrod & Díaz 12728 

(UPRRP). Dorado: wet sand, 11 Jan 1964, 

Woodbury s.n. (UPR); Dorado, 1 Nov 1964, 

Woodbury s.n. (NY); Vega Baja: Tortuguero, 12 

Mar 1960, Woodbury s.n. (NY, UPR); Tortuguero, 

May 1960, Woodbury s.n. (NY, UPR); Tortuguero, 

Sep 1960, Woodbury s.n. (UPR); Tortuguero 

lagoon, Vega Baja, 3 Mar 1963, Woodbury s.n. 

(UPR); Tortuguero, Sep 1968, Woodbury s.n. (NY, 

UPR, US); Tortuguero area, in savanna at sea level, 

Liogier et al. 33611 (UPR); SE shore of Laguna 

Tortuguero off rte. 687, Bradley et al. 22111 

(GMUF); Tortuguero, Woodbury s.n. (NY, UPR, 

US). Yabocoa: Guayanes, May 1970, Woodbury 

s.n. (NY, UPR); 

10. Rhynchospora divaricata (Ham.) M. T. 

Strong, comb. nov. - Fimbristylis divaricata 

Ham., Prodr. 14. 1825. TYPE: Puerto Rico. 

Ledrú s.n. (holotype: P-Desv.!). 

Fimbristylis hirsuta Hochst. ex Steud., Syn. Pl. 

Glumac. 2: 116. 1855. TYPE: Surinam, 

Hostmann 1127 (holotype: P; isotypes: BM- 

2!, G-2!, MO!, P). 

Rhynchospora trichodes C. B. Clarke in Urban, 

Symb. Antill. 2: 116. 1900. Schoenus 

hispidulus Vahl, Enum. Pl. 2: 219. 1805. 

Dichromena hispidula (Vahl) Kunth, Enum. 

Pl. 2: 279. 1837. Haloschoenus hispidulus 

(Vahl) Nees in Martius, Fl. Bras. 2(1): 124. 

1842. Rhynchospora hispidula (Vahl) Boeck., 

Linnaea 37: 604. 1873, non Grisebach, 1866. 

TYPE: French Guiana, coll. ign. (holotype: B, 

destroyed, photo US ex B). 

Caespitose annual, 10-50 cm tall. Culms 

ascending,(0.5-) 0.6-3 mm wide, trigonous or 

subtriquetrous, often with obtuse angles, 

somewhat firm, flexible, longitudinally channeled 

along one side medially, finely ribbed, green, 

pilose. Leaves 3-6, ascending, basal and cauline, 

12-40 cm long; sheaths eligulate, subinflated, 

loosely surrounding culms, finely veined, coarsely 

pilose to subglabrate, inner band membranous on 

proximal sheaths, only the orifice membranous on 

distal sheaths, the orifice concave, long-pilose on 

margin; blades flattened or folded, (1.5-) 2-5 mm 

wide, finely veined, green, glabrate, margins and 

abaxial midvein pilose. Inflorescence a terminal 

and 1-2 lateral lax, compound, corymbose partial 

panicles from the upper leaf-like bracts which may 

elongate to 23 cm; prophyllar bracts often bearing 

a sterile axillary spikelet; partial panicles very

344 


variable in size, particularly in width, 3.5-20 × 

(3-) 5-20 cm; lower primary corymb branches and 

spikelet pedicels often strongly reflexed or 

recurved at maturity, pilose to glabrous, the lowest 

primary branches often greatly elongating up to 

15 cm; spikelets borne singly at tips of branches 

or secondary and tertiary branchlets, often 

nodding, broadly ovoid to globose, 3.2-4.5 × 2.3- 

4 mm, obtuse or broadly acute at apex, the scales 

spreading with developing achenes; scales 11-48 

per spikelet, submembranous, dorsally obtuse to 

rounded, glabrous, brown, reddish brown lineolate, 

margins broadly scarious, midcosta very slender, 

brown, inconspicuous, the tip pilose or glabrate, 

shorter than to equaling obtuse to rounded apex 

of scale, lateral nerves indistinct, distal fertile 

scales membranous, translucent; fertile scales 9- 

45, broadly ovate, circular, oblate, subrhombic, or 

ovate to oblong-ovate (distal scales), 2.3-3 × 1.2- 

3 mm; sterile scales 2-3, ovate, 1.5-2 × 1.2-2 mm. 

Stamens 3, the anthers 0.8-1.2 mm long; style 2branched. 

Achene biconvex, oblate-obovate to 

transversely elliptic-obovate, wider than long, 1- 

1.4 × 1.2-1.5 mm, gradually rounded at apex, 

truncate at base with 2 lateral rounded swellings 

on either side of the short 0.2-0.4 mm long strapshaped 

stipitate base, transversely rugose, shiny, 

pale brown to dark brown or blackish at maturity; 

style base depressed-triangular or lunate, 0.3-0.5 

mm long and as wide as achene, 2-lobed, the lobes 

extending down along margins of achene to about 

the middle; bristles absent. 

General distribution: Central America, Puerto 

Rico, Trinidad, and northern South America. 

Distribution in Puerto Rico: Known in Puerto 

Rico only from the type collection. 

Note: An earlier described name discovered 

for this species, Fimbristylis divaricata Ham., has 

priority over Rhynchospora trichodes C. B. Clarke, 

a new name that was coined for the illegitimate 

homonym, R. hispidula (Vahl) Boeck. A transfer 

from Fimbristylis to Rhynchospora is thus 

necessary. Because the type of Rhynchospora 

hispidula is from French Guiana and no other 

record of this species exists from Puerto Rico or 

anywhere else in the West Indies, the locality of 

the type specimen of Rhynchospora divaricata is 

questionable. However, this species is known from 

Trinidad and the collector, Ledru, was known to 

have collected on that island. This suggests that 

the specimen may perhaps be mislabeled. Other 

occurrences of Rhynchospora divaricata in Puerto 

Rico are based on specimens in the Delesert 

herbarium (G) and Willdenow herbarium (B). A 

hand sketched rendering and inflorescence 

fragment of the G specimen at NY shows that it is 

that species. The sketch of the specimen label reads 

“Porto rico” only, and below that on the sheet the 

poor scribbling indicates that Salzmann is the 

collector. However, Salzmann has not been 

recorded as collecting in Puerto Rico. His 

collections in the New World were exclusively 

from South America, mainly Bahia, Brazil 

between 1827-1830. The name Rhynchospora 

trichodes was based on Schoenus hispidulus Vahl. 

In his protolog, Vahl (1805) was uncertain of the 

locality of the specimen he based his new species 

on citing it as “Habitat in Carolina “(?)” ex herb. 

Lamarckii.” Kunth (1837: 279) indicated that 

Vahl’s new species was from French Guiana, not 

Carolina, “Guiana (nec Carolina)” and cites a 

specimen from the Willdenow herbarium (B- 

Willd. 1116) which strangely enough is labeled 

by Willdenow as occurring in “Portorico”. A 

photograph of a specimen in B ex US, now 

probably destroyed, is labeled in Vahl’s hand as 

Schoenus hispidulus Vahl, fide Vahl, ex Willd. 

Herb., and as occurring in “Cajenna” (Cayenne, 

French Guiana). A label attached to this sheet, in 

Vahl’s hand, is inscribed “Schoenus hispidulus 

Willd. herb. no. 1116, fol 1,” and bears a lengthy 

description of the plant. The B sheet appears to be 

type material and the specimen in the Willdenow 

herbarium (B-Willd. 1116) has probably been 

mislabeled and is from French Guiana. 

Fimbristylis divaricata was treated by Steudel 

(1855) as conspecific with F. hirtella Ham., Prodr. 

14. 1825, nom. illeg., non Vahl, 1805 (F. hamiltonii 

(Ham.) Steud., as a new name), also cited from 

Puerto Rico by Hamilton. A search by Odile Poncy, 

pers. comm. of the Desvaux herbarium at P for 

the type collection of F. hirtella Ham. was 

unsuccessful. 

11. Rhynchospora domingensis Urb., Symb. 

Antill. 7: 170. 1912. Rhynchospora longiflora 

var. domingensis (Urb.) Kük., Bot. Jahrb. 

Syst. 74: 396. 1949. Type: Dominican 

Republic; vicinity of Constanza. Türckheim 


Rhynchsopora longiflora of authors, non C. Presl, 

1828.



rhizome short, stout; roots medium to coarse, 0.7- 

1.5 mm thick; sheathing base of culm 3-10 mm 

wide. Culms sub-erect to ascending, 0.8-3 mm 

wide, trigonous or triquetrous proximally, obtusely 

trigonous to subterete distally, stiff and hardened, 

finely ribbed, scabridulous, often smooth at nodes 

and at very base, pale brown proximally, green 

distally, chestnut brown at nodes proximally. 

Leaves 6-20, ascending, basal and cauline; sheaths 

somewhat loose, short on basal leaves, longer on 

cauline leaves, to 6 cm long, herbaceous, finely 

veined, light brown to brown proximally, golden 

brown adaxially on basal leaves, glabrous; ligule 

absent; blades linear-lanceolate, 9-60 cm × 1.8-6 

mm, V-shaped proximally to subflattened or 

subplicate distally, acuminate or long-attenuate to 

triquetrous apex, finely veined abaxially, finely 

cellular-reticulate and essentially smooth 

adaxially, scabridulous abaxially (at least distally), 

margins and abaxial midvein scabridulous, (at least 

distally), green. Inflorescence a terminal and series 

of 2-3 lateral, lax (at maturity), compound, partial 

corymbiform panicles from the upper sheathing 

bracts; terminal partial panicle 2-8 × 2-10 cm, the 

lateral smaller, on slender peduncles to 10 cm long; 

sheathing bracts equaling to slightly overtopping 

subtending partial panicles, to 8 cm long; spikelets 

lanceoloid to fusiform, 8-12 × 1-1.5 (-1.7) mm, 

long-acuminate at apex, cuneate at base, with 6-8 

(9) scales, pedicellate, solitary at branch tips or 

subfasciculate on secondary branches; fertile 

scales 3-5, ovate-elliptic (sterile basal scales of 

spikelet), ovate-lanceolate to oblong-lanceolate or 

lanceolate (terminal scale of spikelet), 4-7.5 (-8) 

× 1.6-2.8 mm, slightly curvate keeled, dorsally 

obtuse to subrounded, herbaceous to thinly 

herbaceous or submembranous, finely cellularstriate, 

semi-glossy, light brown, reddish brown 

medially along midcosta, essentially glabrous, 

margins narrowly scarious, midcosta fine, 

extended beyond acute apex as a 0.5-1 mm long 

awn. Stamens 3, the anthers 2-2.8 mm long, 

glandular-apiculate, truncate with crystalline 

trichomes at base; style entire or shortly bi-fid or 

2-branched at tip. Achene biconvex, obovate or 

broadly oblong-obovate, 1.5-2 × 1-1.4 mm, 

broadly rounded or obtuse at apex, shortly 

attenuate at base, often margined but indistinctly 

so, semi-glossy, glossy at maturity, finely and often 

indistinctly cancellate with 13-20 longitudinal 

rows of very shallow horizontal pits on each face, 

pale brown to yellowish brown, brown at maturity; 

style base lance-attenuate with concave margins, 


flattened, attenuate to apex, shallowly articulated 

at base, pale brown with reddish lineations or dots; 

bristles 3-6, subulate, finely antrorsely barbed, pale 

or reddish, exceeding the achene. 


Puerto Rico. 

Distribution in Puerto Rico: Forested habitats 

on slopes, thickets, and along streams. Known to 

occur only in west-central Puerto Rico in Adjuntas 

and Jayuya. 

Note: Rhynchospora domingensis is closely 

related to Rhynchospora longiflora C. Presl, but 

characteristically has narrower leaf blades (1.8- 

6 vs. 8-12 mm wide) and conspicuously red-, 

purple-, and black-dotted or -lineolate upper 

culms, sheathing bracts, inflorescence branches, 

and scales (vs. not conspicuously dotted or 

lineolate). The culms and abaxial distal portion of 

leaf blades and sheathing bracts are scabridulous 

or short appressed-strigose (vs. smooth or 

essentially so). Spikelet scales are light brown and 

red-lineolate (vs. dark reddish or rusty brown). 

Lastly the achenes have more rounded shoulders 

at apex and are glossy-brown at maturity (vs. 

abruptly obtuse or subtruncate at apex and yellowbrown). 


Adjuntas: Guilarte Peak way, Woodbury s.n. (NY, 

UPR). Jayuya: Cerro de Punta, Liogier et al. 30576 

(NY, UPR). 

12. Rhynchospora fascicularis (Michx.) Vahl, 

Enum. Pl. 2: 234. 1805; Schoenus fascicularis 

Michx., Fl. Bor.-Amer. 1: 37. 1803. Type: 

United States; “Carolina”. Bosc. s.n. 

(holotype: P, photo at GH; isotype fragment: 

NY). 

Caespitose, rhizomatous perennial, 30-95 cm 

tall; rhizome short; roots medium-sized. Culms 

erect to ascending, 0.6-1.7 (-2) mm wide, obtusely 

3-angled proximally to terete distally, 

subherbaceous, firm, not easily compressed, finely 

ribbed, green, glabrous. Leaves 4-11, ascending, 

primarily basal, 3-5 cauline; sheaths short, 

herbaceous, finely veined, pale brown to brown, 

glabrous; ligule absent; blades narrowly linear, 6- 

30 cm × 1-3 mm, V-shaped or involute, 

herbaceous, finely veined, green to pale, glabrous,

346 


margins essentially smooth, but sometimes 

remotely and bluntly scabrous, the apex filiform, 

long-acuminate to antrorsely scabrous apex. 

Inflorescence a simple fascicle of spikelets or 

corymbosely compound group of 3-5 fascicles, 

sometimes with 1-2 lateral fascicles from the upper 

sheathing bracts; fascicles hemispherical, 5-10 × 

5-15 mm; spikelets 3-4 × 1-1.8 mm, ovoidellipsoid 

or ellipsoid, acute to short-acuminate at 

apex, short-cuneate at base, the spikelet scales 5- 

6, spreading with maturing achenes; fertile scales 

3-4, ovate, 2-3 × 1.3-2.8 mm, often inrolled around 

flowers, herbaceous to submembranous, dorsally 

rounded, reddish brown, glabrous, margins 

narrowly to broadly scarious, erose, acute at apex, 

midcosta distinct, very narrow, brown to pale 

brown, lateral nerves indistinct, sides often with 

fine blackish striations running longitudinally and 

curving out to the margins. Stamens 3, sometimes 

1-2 abortive, the anthers 1-1.5 mm long; style 2branched. 

Achene biconvex, obovate to broadly 

so, suborbicular, or elliptic-obovate, 1.3-2 × 1.1- 

1.5 mm, indistinctly transversely rugulose to 

smooth, umbonate or gradually biconvex, dark 

brown to blackish with a yellowish brown to deep 

red circular patch medially, acute to abruptly 

acuminate to base, truncate at apex; style base 

shallowly triangular to triangular or triangularsubulate, 

0.5-1 mm long, 0.7-1 mm wide at base, 

brown, the margins smooth to sometimes scabrous; 

bristles 6, filiform, short or rudimentary to 

exceeding the achene body, reddish, antrorsely 

barbed, often with trichomes at base. 


States west to Texas, Mexico, Central America, 

Greater Antilles, and northern South America. 

Note: This species is represented in Puerto 

Rico by two infraspecific taxa. Rhynchospora 

fascicularis subsp. fascicularis var. distans 

(Michx.) Chapm. is known from Laguna 

Tortuguero only, while R. fascicularis subsp. 

fascicularis var. fascicularis is more widespread. 

Below a key is given to distinguish the two. 

Key to the varieties of Rhynchospora 

fascicularis subsp. fascicularis in Puerto Rico 

1. Achenes obovate to broadly so or suborbicular, 

1.1-1.5 mm wide, umbonate; style base 

triangular; bristles short or rudimentary, rarely 

exceeding achene body.............. R. fascicularis 

subsp. fascicularis var. fascicularis 12a. 

1. Achenes elliptic-obovate, 1-1.3 mm wide, not 

umbonate; style base triangular-subulate; 

bristles exceeding achene body....................... 

.................................................. R. fascicularis 

subsp. fascicularis var. distans 12b. 

12a. Rhynchospora fascicularis subsp. 

fascicularis var. fascicularis. 

(See above for type and general distribution). 

Distribution in Puerto Rico: On wet to moist 

sandy or mossy habitats; swampy areas, savannas, 

and coastal marshes at low elevation. Cataño, 

Dorado, Manatí, Mayagüez, Salinas, San Juan, 

Vega Alta, and Vega Baja. 


Bayamón: Sabana Llana, N.L. Britton & E.G. 

Britton 8700 (NY). Cataño: Heller & Heller 1380 

(NY, US). Dorado: Woodbury s.n. (NY); Johnston 

883 (NY). Manatí: western end of Laguna 

Tortuguero, N.L. Britton & E.G. Britton 7872 (NY, 

US). San Juan: Martín Peña, Stevenson 501 (US). 

Vega Alta: Bo. Sabana, NE of Regadera, Proctor 

45807 (US). Vega Baja, Heller & Heller 1313 (NY, 

US). 

12b. Rhynchospora fascicularis subsp. 

fascicularis var. distans (Michx.) Chapm., Fl. 

South U. S. 527. 1860; Schoenus distans 

Michx., Fl. Bor.-Am. 1: 36. 1803; 

Rhynchospora distans (Michx.) Vahl, Enum. 

Pl. 2: 235. 1805. Type: United States. 

Carolina, Michaux s.n. (holotype: G; holotype 

fragment: NY). 


States, Bermuda, and Puerto Rico. 


from Laguna Tortuguero. 


Vega Baja: Laguna Tortuguero, Liogier et al. 

33592 (UPR). Historically cited for Puerto Rico 

by C. B. Clarke in Urban, Symb. Antill. 2: 125. 

1900, based on Sintenis 1232 (not located). 

13. Rhynchospora filiformis Vahl, Enum. Pl. 2: 

232. 1805; Dichromena filiformis (Vahl) 

Kunth, Enum. Pl. 2: 281. 1837; Spermodon 

filiformis (Vahl) Nees in Martius, Fl. Bras. 

2(1): 118. 1842. Type: Puerto Rico. Ledrú s.n. 

(holotype: C-Vahl; probable isotype: P).


Rhynchospora podosperma C. Wright in Sauvalle, 

Anales Acad. Ci. Méd. Habana 8: 87. 1871. 

Type: Cuba. Wright 3791 (holotype: GH; 

isotype: NY!, US!). 

Rhynchospora longispicata Boeck., Linnaea 37: 

600. 1873. Type: French Guiana. Sagot 1389 

(cited as Sagot 389) (lectotype: BM!; 

isolectotype: K!), here designated. 

Rhizomatous perennial, 13-50 cm tall, culms 

sprouting in close clusters at nodes of rhizome; 

rhizome short, knotty; roots fine to medium, to 1 

mm thick. Culms ascending, bulbous and 

horizontal at very base, abruptly arched-ascending, 

trigonous to obtusely trigonous, firm but flexuous, 

essentially smooth, green, glabrous. Leaves 3-7, 

ascending, primarily basal, 1-2 lower cauline or 

sometimes one to about the middle; sheaths soft, 

finely veined, green, glabrous; ligule a very narrow 

thickened band of tissue at the adaxial junction of 

sheath and blade; blades linear-filiform, thickened, 

3-40 cm × 0.3-3 mm (unfolded), V-shaped to 

somewhat crescentform-capillary or trigonous 

proximally, triangular-channeled distally, often 

folded proximally, firm but flexuous, finely veined 

abaxially, smooth, shining, and cellular-reticulate 

adaxially, green, glabrous, margins generally 

smooth proximally, antrorsely scabrous on margins 

distally, the apex attenuate to tip. Inflorescence a 

terminal and 1-4 lateral corymbose partial panicles 

from the upper sheathing bracts, the lowermost 

long-peduncled; terminal panicle 1.5-5 × 0.6-4.5 

cm, with (1-) 2-20 (-30) spikelets, rarely reduced 

to a single spikelet; spikelets linear, narrowly 

ellipsoid-lanceoloid, (8-) 9-13 (-14) × 1-1.8 mm, 

attenuate at apex, narrowly cuneate at base, with 

6-13 scales; fertile scales 3-9, ovate-lanceolate to 

linear-lanceolate, 5.3-11.5 × 0.8-2.8 mm, variable 

in cross section, dorsally rounded and inrolled 

around developing pistil, lower fertile and sterile 

scales subcoriaceous, smooth and semi-glossy, 

glabrous, brown medially, light brown on sides 

with brown lineations, margins scarious, midcosta 

pale brown to greenish brown, indistinct, very 

narrow, prolonged beyond the acute to acuminate 

apex as an antrorsely scabrous awn, upper fertile 

scales hidden, thin and membranous, linearlanceolate. 

Stamens 3, the anthers 2.5-4 mm long, 

apiculate, truncate at base with crystalline papillae; 

style deeply 2-branched. Achene biconvex, 

obovate to oblong-obovate, 1.5-2.2 (including 

stipitate base) × 1.1-1.3 mm, truncate to concave 

at apex, narrowed below to a tounge-like, finely 

cellular-reticulate stipitate base, relatively smooth 

or distinctly to faintly cellular-reticulate medially, 

light brown to brown with a longitudinal gray 

stripe medially on each side, margins often 

prolonged at apex on either side as a short tooth, 

cellular-reticulate-papillose, the epidermal cells 

bulging at maturity forming a pebbly surface; style 

base triangular, 0.3-0.5 mm long, 0.3-0.7 mm wide 

at base, rimmed by the truncate to concave apex 

of achene, brittle, brownish black; bristles absent. 


America, Greater Antilles, and tropical South 

America. 


Islands: Historically known only from a Ledrú 

collection from Puerto Rico that is the type of the 

species. However, Britton and P. Wilson (1923) 

cite a Ledrú collection from St. Thomas. Not 

collected nor seen since. 

14. Rhynchospora gigantea Link in Sprengel, 

Jahrb. Gewächsk. 1(3): 76. 1820. Type: 

Brazil. Hoffmannsegg s.n. (holotype: B-Willd. 

1129, microfiche US ex B). 

Coarse, robust perennial, 0.6-1.5 m tall; 

rhizome short, thick, knotty, hardened; roots 

coarse, 2-4 mm thick, golden brown, lustrous, 

spongy, easily compressed with internal chambers. 

Culms solitary, ascending, (2-) 3-7.5 mm wide, 

up to 10 mm wide at swollen nodes, finely ribbed, 

smooth, glabrous, trigonous, somewhat soft and 

often hollow proximally just below leaf nodes of 

cauline leaves, firm distally. Leaves 9-20, archedascending, 

spreading, basal and cauline; sheaths 

spongy-thickened (at least proximally), glabrous, 

to 20 cm long, smooth proximally, septate 

nodulose distally; ligule absent; blades narrowly 

linear, flattened to plicate proximally, those borne 

from basal sheaths often spongy-thickened 

proximally, (3.5-) 4-15 dm × (5-) 9-23 (-25) mm, 

glabrous, abaxial side bright green, distinctly 

septate-nodulose, adaxial side dark green, lustrous, 

glossy, smooth, margins and midvein beneath 

antrorsely scabrous, long-acuminate to triquetrous 

apex. Inflorescence of 2-5 remote to subcontiguous 

large compound partial corymbose panicles 

(including the terminal one), (10-) 12-50 (-60) × 

5-15 cm, each subtended at base by a leaf-like

348 


involucral bract, reduced distally, the spikelets 

congested at ray tips in globose or hemispherical 

clusters of 20-40, 12-16 mm in diam; spikelets 

ovoid-ellipsoid, (4.5-) 5-6 (-6.5) × (1.2-) 1.5-2.2 

mm, short-acuminate at apex, obtuse at base, with 

7-11 scales; fertile scales 4-7, ovate-elliptic, the 

distal ones ovate-lanceolate to lanceolate, (3-) 4- 

6 (-6.5) × (2-) 2.2-3.1 (-3.3) mm, thin, uniformly 

brown, glabrous, finely indistinctly veined, the 

apex acute to short-acuminate. Stamens 3, the 

anthers linear, 2-3 mm long; style entire or shortly 

bi-lobed at apex. Achene elliptic-obovate, 2-2.5 

mm × 1-1.2 mm, papillate, transversely rugulose 

medially on sides, the thickened margins with 

constrictions and protuberances; style base 

narrowly triangular, 2-2.8 × 1-1.2 mm, as wide 

and thick as achene at base, spongy-thickened with 

a narrow vertical channel medially on each side; 

bristles 6, exceeding the achene, shorter than to 

equaling tip of style base, antrorsely barbed. 

General distribution: Mexico, Nicaragua, 

Greater Antilles, and tropical South America. 

Distribution in Puerto Rico: In wet sandy or 

muddy soils, marshy and swampy areas, pastures, 

ditches, swales, and savannas at low elevation. 

Dorado, Loíza, Manatí, Mayagüez, San Juan, Toa 

Baja, Vega Alta, and Vega Baja. 


Dorado: Bo. San Antonio, González-Más 350 

(US); vicinity of Dorado, N.L. Britton et al. 6649 

(NY). Loíza: Punta Cangrejos, Johnston & 

Stevenson 1675 (NY). Manatí: Laguna Tortuguero, 

Liogier et al. 30438 (NY); Sintenis 6672 (US). 

San Juan: Santurce, Heller & Heller 980 (NY, US); 

Martín Peña, Stevenson & Johnston 1675 (US). 

Toa Baja: Rd. 866, km 7.2, Punta Salinas, 

González-Más 1145 (US). Vega Baja: Puerto 

Nuevo, Rd. 686, km 4, González-Más 1091 (NY, 

US); vicinity of Vega Baja, N.L. Britton et al. 5786 

(NY). 

15. Rhynchospora holoschoenoides (Rich.) 

Herter, Revista Sudamer. Bot. 9: 157. 1953; 

Schoenus holoschoenoides Rich., Actes Soc. 

Hist. Nat. Paris 1:106. 1792. Type: French 

Guiana. LeBlonde s.n. (holotype: P). 

Schoenus cyperoides Sw., Prodr. 19. 1788, non 

Retzius, 1786; Rhynchospora cyperoides 

Mart., Denkschr. Königl. Acad. Wiss. 

München 6: 149. 1817 (“1816”). Type: 

Jamaica. Swartz s.n. (holotype: S-Sw. R- 

5609). 


rhizome short, thickened; roots coarse. Culm 

ascending, 1.3-3 mm wide, trigonous, firm, finely 

ribbed, green, smooth, glabrous, the angles smooth 

proximally to bluntly scabrous distally. Leaves 7- 

12, ascending, primarily basal, 2-4 lower cauline; 

sheaths herbaceous, often spongy-thickened, 

finely veined, septate-nodulose, green to pale 

brown proximally, glabrous; ligule a very narrow, 

thickened band or flap of tissue at adaxial junction 

of sheath and blade, inconspicuous on upper 

sheaths; blades narrowly linear, 13-75 (-82) cm × 

2.2-6 (-6.5) mm (unfolded), often folded and 

spongy-thickened proximally, flattened distally, 

herbaceous, finely veined, septate-nodulose, 

sometimes indistinctly so, green, glabrous, 

margins and midvein beneath antrorsely scabrous, 

the apex attenuate to triquetrous apex. 

Inflorescence a single and often 1-2 lateral simple 

to compound open corymbs; terminal corymb 4- 

13 × 3.5-10 cm, the heads of spikelets 2-20, 7-13 

mm in diam; spikelets numerous, tightly 

compacted in globose capitula, ovate-elliptic, 

(2.5-) 3-5 × 1.1-1.8 mm, often longitudinally 

falcate, acuminate at apex, obtuse at base, with 5- 

8 scales; fertile scales 3-5, ovate to ovate-elliptic, 

2.4-4.2 (-4.5) × 1.4-2.5 mm, thin, membranous, 

curvate-keeled, often somewhat longitudinally 

folded, acute to acuminate at apex, reddish brown 

or golden brown, lustrous, glabrous, margins erose, 

midcosta indistinct except at apex, dark brown, 

extending as smooth to bluntly scabrous short 

mucro, lateral nerves. Stamens 3, 2 in terminal 

scale of spikelet, the anthers 1.1-1.8 mm long, 

apiculate; style entire or sometimes shortly bilobed 

at apex. Achene biconvex, 2-sided, widely 

obovate to narrowly obovate, 1.4-2 × (0.9-) 1-1.5 

(-1.6) mm, rounded at apex, cuneate to base, finely 

transversely rugulose, often indistinctly so and 

appearing smooth, shiny, lustrous, brown to dark 

brown or grayish brown, the margins above middle 

antrorsely scabrous; style base linear-lanceolate, 

4-angled, 1.8-3.1 mm long, 0.2-0.4 (-0.6) mm wide 

at base, antrorsely scabrous on margins, pale 

brown to reddish brown; bristles 6, filiform, terete 

in cross section, antrorsely barbed, reddish, to 3 

mm long, surpassing achene, shorter than to 

equaling tip of style base. 


America, West Indies, tropical South America, 

Africa, and Madagascar.


Distribution in Puerto Rico: Occurring in a 

diversity of wet habitats including, swampy 

grounds, savanna, meadows, open fields, grassy 

areas, marshy areas, pastures, wet mountain 

forests, dry scrub forests, clearings, hollows, peaty 

hillsides, roadside ditches, and exposed and 

disturbed sites generally at low and middle 

elevations, but occurring at high elevation in the 

Luquillo Mountains. Arecibo, Camuy, Cayey, 

Ceiba, Ciales, Cidra, Dorado, Fajardo, Guayama, 

Humacao, Loíza, Luquillo, Manatí, Maricao, 

Maunabo, Mayagüez, Moca, Naguabo, Patillas, 

Río Grande, San Juan, San Sebastián, Vega Alta, 

Vega Baja, and Yabucoa. 



10568 (MAPR, UPRRP, US). Cayey: Bo. 

Guavate, edge of Carite Commonwealth Forest, 

Breckon & Lewis 4033 (MAPR, MO); Cerro La 

Santa, Guavate, Liogier et al. 33506 (MO, NY). 

Ceiba: Caribbean National Forest, near Pico del 

Este, Axelrod 8352A (UPRRP). Ciales: Sargent 

3052 (US). Cidra: Pueblo Viejo, Hioram 313 (US). 

Dorado: Johnston 895 (US). Fajardo: Sintenis 

1236 (US). Guajataca: Boy Scout Reservation, 

hwy. 119, km 18, south from Camuy, Wadsworth 

175 (NY). Guayama: Carite Forest Reserve, 

Axelrod et al. 608 (NY, UPRRP). Humacao: Pozal, 

Playa de Humacao, Eggers 588 (US). Luquillo: 

top of El Yunque Mt., González-Más 1536 

(MAPR, NY, US); Santa Catalina, Stevens & Hess 

2745 (MAPR, NY). Manatí: Laguna Tortuguero, 

N.L. Britton et al. 3846 (NY). Maricao: González- 

Más 457 (NY, US). Maunabo: Cuchilla de ls 

Panduras, Axelrod et al. 2055 (UPRRP). 

Mayagüez: Guanajibo, Fredholm 4586 (MAPR). 

Moca to Mayagüez, González-Más 4011 (MAPR). 

Naguabo: Sierra de Luquillo, Caribbean National 

Forest, Proctor & Thomas 43204 (US). Patillas: 

on the property of Tropic Ventures, along Rt. 184, 

just S of Carite, Miller et al. 6586 (MO, UPRRP). 

Río Grande: Luquillo Mountains, Acevedo-Rdgz. 

10515 (MAPR, UPRRP, US). San Juan: Río 

Piedras to Trujillo Alto, Chase 6368.5 (US); 

Laguna San José, Hioram s.n. (NY). San 

Sebastián: Sargent 244 (US). Vega Baja: 

Tortuguero Lagoon, González-Más 1060 (NY, 

US). Yabucoa: Punta Guayanes, Axelrod et al. 

2749 (MO, NY, UPRRP). 

16. Rhynchospora jamaicensis Britton, Bull. 

Torrey Bot. Club 41: 1. 1914. Type: Jamaica; 

Hardwar Gap. Britton 3322 (holotype: NY!). 

Rhizomatous perennial, 50-175 cm in length; 

rhizome short, knotty; roots medium to coarse, to 

1 mm thick; sheathing base of culm about same 

width of culm distally, with few nodes. Culms 

often clumped together on rhizome, ascending to 

scandent or trailing, 1.8-4 mm wide, trigonous 

proximally, triquetrous to compressed trigonous 

distally, hardened, finely ribbed, light brown with 

brown nodes proximally, green distally, short 

pilose to glabrate, the nodes few and remote 

proximally, many and approximate medially. 

Leaves 8-30, ascending, spreading, primarily 

cauline; sheaths elongate proximally, short 

medially, herbaceous, finely veined, green, short 

pilose abaxially to wooly on inner band, the inner 

band herbaceous; ligule absent; blades linear, 6- 

36 cm × 2-7 mm, plicate or subflattened 

proximally with dorsally obtuse to rounded pleats, 

the margins often involute, attenuate to triquetrous 

apex, herbaceous, finely veined abaxially with dull 

surface, glossy and somewhat smooth adaxially 

with two lateral veins evident, margins and abaxial 

midvein antrorsely scabrous or setose-scabrous (at 

least distally), green, short pilose abaxially (at least 

proximally), glabrous adaxially or short pilosescabrous 

on sides, particularly distally, the adaxial 

midvein closely strigose. Inflorescence a terminal 

and series of 3-6 lateral, remote to subcontiguous, 

compound, pyramidal partial panicles from the 

upper sheathing bracts; sheathing bracts reduced 

distally, overtopping proximal partial panicles, 

shorter than to slightly exceeding terminal panicle; 

terminal panicle 4-7 × 3-6 cm, the lateral smaller, 

gradually reducing in size proximally, on short, 

erect, stiff peduncles; panicle branches pilose to 

glabrescent; spikelets 20-160 per partial panicle, 

ovoid-lanceoloid, 4-7 × 1-1.7 mm, acuminate at 

apex, short-cuneate at base, with 8-12 scales; 

fertile scales 5-9, ovate-elliptic to ovate-lanceolate, 

(2.6-) 3-5.5 × 1.6-2.7 (-3) mm, boat-shaped, 

slightly curvate, dorsally obtuse to rounded, thinly 

herbaceous, finely cellular-striate, dull (flat), 

brown with lighter brown or reddish brown 

margins, glabrous, margins narrowly scarious, 

entire, midcosta fine, inconspicuous, pale, 

extended beyond the acute to acuminate apex as a 

mucro. Stamens 2, the anthers 1.5-2.2 mm long,

350 


subulate-apiculate, truncate at base with minute 

crystalline papillae or lobes; style shortly 2branched 

at apex. Achene biconvex, sometimes 

unequally so, broadly obovate to rounded-obovate, 

0.8-1.1 × 0.7-1 mm, obtuse to rounded or 

subtruncate at apex, widely cuneate to rounded at 

base, cellular-cancellate with 11-16 longitudinal 

rows or cells, glossy, brown, the cell walls light 

brown; style base triangular-lanceolate, 0.9-1.5 

mm long, 0.5-0.7 mm wide at base, flattened, 

shallowly 2-lobed at base, brittle, crustose, light 

brown to brown, often whitened; bristles 

rudimentary or 1-2, subulate, antrorsely barbed, 

reddish, shorter than to rarely equaling achene. 


Distribution in Puerto Rico: Occurs in moist 

montane forests from 800-1300 m; openings on 

wet slopes, thickets on road banks, forest trails, 

and roadside ditches. Adjuntas, Cayey, Ceiba, 

Jayuya, Juana Díaz, Luquillo, Orocovis, Ponce, 

Río Grande, San Lorenzo, and Yauco. 

Note. This species was placed in synonymy 

with Rhynchospora polyphylla (Vahl) Vahl by 

Kükenthal (1949). However, it differs from R. 

polyphylla in having pubescence abaxially on 

sheaths and leaf blades (vs. essentially glabrous); 

spikelets 4-7 × 1-1.7 mm (vs. 3-5 × 1 mm); spikelet 

scales that are brown with lighter brown or reddish 

brown margins (vs. uniformly stramineous to light 

reddish brown); achenes 0.7-1 mm broad (vs. 0.6- 

0.8 mm broad); achene surfaces with 11-16 

longitudinal rows of cells on each face (vs. 5-10 

longitudinal rows); and a triangular-lanceolate 

style base 0.9-1.5 mm long (vs. narrowly triangular 

and 0.3-0.7 mm long). 


Adjuntas: Alto de la Bandera, Stevens 8254 (NY); 

Monte Guilarte, Liogier et al. 30313 (NY). Ceiba- 

Naguabo: Caribbean National Forest, along road 

near Pico del Este, Axelrod 8365 (UPRRP). 

Jayuya: Near Cerro de Punta, Liogier 10832 (NY, 

US). Juana Díaz: Sargent 3196 (US). Luquillo: 

road toward top of El Yunque, González-Más 1545 

(MAPR, NY, US); Near Pico de Este, Luquillo 

Mts., Liogier et al. 33302 (NY). Río Grande: 

Caribbean National Forest, along Mt. Britton trail, 

Axelrod 6216 (US). San Lorenzo: Sierra de Cayey, 

Carite Forest Reserve, Proctor et al. 47103 (US). 

17. Rhynchospora lindeniana Griseb., Cat. Pl. 

Cub. 244. 1866. Type: Cuba. Linden 1945 

(holotype: GOET; isotype: GH!). 

Rhynchospora bahamaensis Britton, Torreya 13: 

217. 1913; Rhynchospora lindeniana var. 

bahamaensis (Britton) Gale, Rhodora 46: 223. 

1944. Type: Bahamas. Britton & Brace 588 


Caespitose perennial forming dense tussocks, 

10-75 cm long; rhizomes short, inconspicuous; 

roots medium, to 1 mm thick. Culms ascending, 

0.3-0.8 mm wide, obtusely trigonous to subterete, 

slender to filiform, firm but flexuous, very finely 

ribbed, smooth, light green, glabrous. Leaves 

numerous, erect, spreading, primarily basal, 1-3 

lower cauline; sheaths finely veined abaxially 

often indistinctly red-dotted, pale green to pale 

brown, glabrous; ligule essentially absent except 

for a thickened area of tissue at adaxial junction 

of sheath and blade; blades setaceous to linearsetaceous, 

4-50 cm × 0.2-1 (1.2) mm, flattenedcrescentform 

to widely v-shaped, or subplicate, 

long-attenuate to apex, abaxial surface smooth 

with a single prominent pale midvein, often 

scabrous distally, lateral veins indistinct, adaxial 

surface semi-glossy, finely transversely rugulose, 

sometimes indistinctly so, margins very finely 

serrate-scabrous, light green, glabrous. 

Inflorescence a terminal and series of 3-5 lateral 

corymbose partial panicles from the upper 

sheathing bracts; corymbiform partial panicles 

small, congested, 6-20 (-30) × 4-10 mm, the 

proximal on long, flattened-setaceous peduncles, 

antrorsely scabrous on margins, 0.2-0.3 mm wide; 

spikelets ovoid-ellipsoid, 2-3 (-3.2) × 0.6-1 mm, 

acute to acuminate at apex, narrowly cuneate at 

base, with 5-7 scales; fertile scales 1-3, widely 

ovate to rounded-ovate, 1-1.7 × 1-1.5 mm, curvatekeeled, 

dorsally obtuse to rounded, deeply boatshaped, 

thinly herbaceous to thickly membranous, 

finely cellular striate or roughened, semi-glossy, 

dark brown to ferruginous, dark brown on sides, 

glabrous, margins broadly whitish scarious, 

midcosta fine, indistinct, pale green, prolonged as 

a short mucro beyond the acute to obtuse apex. 

Stamen 1, the anther 0.6-1 mm long, apiculate, 

truncate at base; style 2-branched. Achene planoconvex 

or unequally biconvex, ovate to oblongovate, 

1.1-1.5 × 0.7-0.9 mm, narrowed to a 

truncate apex, acute to cuneate at base, indistinctly 

transversely rugulose, semi-glossy to glossy, light 

reddish brown to dark brown or red medially; style 

base triangular-acuminate or sometimes prolonged 

and subulate, compressed, 0.6-1 mm long, 0.5 mm


wide at base, whitish; bristles 6, subulate, fragile, 

weakly ascending, finely and closely antrorsely 

barbed, reddish or reddish brown, exceeding 

achene, shorter than to equaling the tip of style 

base, clustered on abaxial side of achene. 

General distribution: Mexico (Quintana Roo), 

Guatemala, and the Greater Antilles. 

Distribution in Puerto Rico: On serpentine 

soils in open forests, along trails, rocky slopes and 

ravines, gravelly areas, and thickets, primarily in 

uplands, 600-900 m. Maricao, Mayagüez, Sabana 

Grande, San Germán, and Yauco. 

Note: Gale (1944) treated Rhynchospora 

bahamaensis Britton as a variety of R. lindeniana. 

She separated typical R. lindeniana from var. 

bahamaensis by its longer spikelets (3-3.2 mm 

long vs. 2-2.4 mm long respectively); spikelets 

with a solitary achene vs. having 2 achenes; and 

longer achenes (1.2-1.4 mm long vs. 1.1-1.2 mm 

long respectively). However, spikelets of the 

isotype are 2-2.6 mm long and have 2 achenes, as 

in var. bahamaensis, and the difference in the 

length of the achenes is not significant. Achenes 

from the type of R. lindeniana are plano-convex 

with a nearly flat or slightly concave side, 

characteristic of those of R. bahamaensis in which 

the developing achenes are nearly parallel to each 

other on the rachilla and tightly pressed together 

at maturity, the nearly flat or slightly concave side 

being where the two achenes press against each 

other. We were not able to find a single, consistent 

character to separate these two taxa and they are 

here treated as conspecific. 


Maricao: Indiera Fría, N.L. Britton et al. 4540 

(NY); mountain ridge E of Río Pastrero, Proctor 

et al. 45212 (US); Río Maricao, N.L. Britton et al. 

2427 (NY, US); Maricao to Monte Alegrillo, N.L. 

Britton et al. 2552 (NY, US); Monte Alegrillo, 

Hess 598 (MAPR, NY); Monte de Estado, 

Caminero & García 147 (MAPR). Mayagüez: Las 

Mesas, Liogier 10522 (NY, US). Sabana Grande: 

Bo. Susúa Alta, Susúa Forest Reserve, Axelrod et 

al. 10189 (UPRRP, US). Yauco: Susúa, Woodbury 

s.n. (NY). 

18. Rhynchospora marisculus Nees in Martius, 

Fl. Bras. 2(1): 142. 1842. Lectotype: Brazil; 

Rio de Janeiro. Martius 3193 (M), designated 

by T. Koyama, Mem. New York Bot. Gard. 

23: 56. 1972. 

Rhynchospora borinquensis Britton, Bull. Torrey 


Sierra de Naguabo. Shafer 3515 (holotype: 



rhizome short, thick, woody. Culms crowded along 

rhizome, ascending, 1.5-4.5 mm wide, trigonous 

to obtusely trigonous, stiff, hardened and brittle, 

asperous with minute whitish blisters or prickles, 

finely ribbed, pale green, glabrous. Leaves 8-15, 

ascending, basal and cauline, the lowermost 

bladeless; sheaths short, somewhat stiffened, 

finely veined, green to pale green, often tinged 

with brown, glabrous; ligule absent; blades 

narrowly linear, 7-90 cm × (1.2-) 1.5-7.5 (-10) mm, 

plicate to subflattened, substiffened, green, semiglossy 

adaxially, asperous with minute whitish 

blisters or prickles abaxially, smooth and finely 

veined (indistinctly so) adaxially, glabrous, smooth 

proximally, finely antrorsely serrate-scabrous 

distally, long-acuminate to triquetrous tip. 


compound partial corymbs with erect to strongly 

ascending branches from the upper sheathing 

bracts; corymbs strict, the terminal corymb 5-17 

× 1.5-4 cm, often curving or nodding at maturity, 

with typically well over 100 spikelets, the spikelets 

in fascicles at branch tips; spikelets narrowly 

ovoid-ellipsoid to ovoid-lanceoloid, often slightly 

curvate, 4.5-6 (-6.5) × 0.8-1.5 mm, acuminate at 

apex, narrowly cuneate at base, with 5-10 scales; 

fertile scales 2-6, ovate-elliptic to ovate-lanceolate 

or linear ovate-lanceolate, 3.4-5 × 1.2-2.4 mm; 

upper fertile scales of spikelet reduced, thin, 

subtranslucent or membranous, yellowish brown, 

but with perfect flowers; lower fertile scales of 

spikelet dorsally obtuse to rounded, thin to 

submembranous, finely striate, brown or tinged 

with reddish brown, often with dark brown 

lineations, glabrous, midcosta fine, pale reddish 

brown, extending beyond apex as a mucro on distal 

scales of spikelet, as a short awn on proximal 

scales of spikelet, lateral nerves indistinct, margins 

entire, narrowly scarious, the apex acute to 

acuminate. Stamens 3, the anthers 1.5-2.3 mm 

long, lobed with crystalline projections at base, 

subulate and awl-like at tip, bearing minute 

crystalline projections; style 2-branched. Achene 

biconvex, sometimes with unequal sides, the 

abaxial side usually more tumid than the adaxial

352 


side, obpyriform, 1.5-2 × 0.8-1.2 mm, subattenuate 

to base, rounded to subtruncate at apex, 

transversely rugulose, semi-glossy, yellowish 

brown, margins and apex of achene body forming 

a narrow, constricted edge, pale at junction with 

style base; style base triangular-lanceolate, 

flattened, 1.2-1.6 mm long, 0.5-0.7 mm wide at 

base, brittle, pale brown to brown or sometimes 

whitish; bristles 6, subulate to compressedsubulate, 

3-6 mm long, often 3 times as long as 

achene body, overtopping achene and style base, 

finely antrorsely barbed, reddish. 

General distribution: Mexico (Chiapas, 

Veracruz), Central America, West Indies, and 

South America south to Argentina. 


from wet openings in forested areas in the Luquillo 

and Naguabo Mountains, in the municipalities of 

Ceiba, Naguabo, and Río Grande. 


Ceiba-Naguabo: Caribbean National Forest, near 

Pico del Este, Axelrod et al. 10171 (UPRRP). 

Luquillo: Near Pico del Este, Luquillo Mts., 

Liogier et al. 33291b (NY). Naguabo: road 191, 

km 13.5, Sierra de Luquillo, González-Más 1418 

(MAPR, NY). Río Grande: El Yunque, Sargent 

526 (US). 

19. Rhynchospora microcarpa Baldwin ex A. 

Gray, Ann. Lyceum Nat. Hist. New York 3: 

202. 1835. Lectotype: United States; North 

Carolina. Curtis s.n. (NY), designated by W. 

W. Thomas, Brittonia 44: 33. 1992. 


tall; rhizome short, inconspicuous. Culms 

ascending, 1-3.2 (-4) mm wide, obtusely trigonous, 

hardened but flexuous, finely and coarsely ribbed, 

smooth, green, glabrous. Leaves numerous, 

ascending, primarily basal, 2-4 cauline; sheaths 

elongate, thickly herbaceous, finely veined, green 

distally, pale brown proximally, glabrous; ligule 

absent; blades linear, 7-52 cm × (0.7) 1-3.5 (-4) 

mm (flattened), thickly herbaceous, flattened, Vshaped, 

or subfolded, often spirally twisted and 

contorted distally with age, long-attenuate to 

triquetrous apex, finely veined abaxially, very 

finely cellular-reticulate and subsmooth adaxially, 

margins and abaxial midvein antrorsely scabrous 

distally, smooth proximally, green, glabrous. 

Inflorescence a terminal and series of 1-3 (-5) 

lateral corymbose partial panicles from the upper 

sheathing bracts; partial panicles, (1-) 1.5-8 × 1-6 

cm, the smaller lateral panicles on relatively short 

peduncles 1-5 cm long, spikelets in small fascicles 

at branch tips; spikelets ovoid to ovoid-globose, 

2.5-3.5 (-4) × 1.5-2 mm, acute to narrowly acute 

at apex, acute to sub-rounded at base, with 6-9 

scales; fertile scales 3-6, widely ovate to rounded 

or very broadly obovate, 1.5-2.7 (-3) × 1.3-2.6 mm, 

deeply boat-shaped or cuplike, curvate-keeled, 

dorsally obtuse to subrounded, thinly herbaceous, 

finely cellular-striate, semi-glossy, dark brown to 

brownish black, glabrous, margins scarious to 

narrowly so, erose, midcosta very fine, extended 

beyond the obtuse to broadly obtuse or emarginate 

apex as a short mucro Stamens 3, the anthers 

(0.5-) 1-2 mm long, minutely apiculate, truncate 

with crystalline papillae at base; style 2-branched. 

Achene thinly biconvex, obovate to widely 

depressed-obovate, 1.1-1.5 × 0.8-1.3 (-1.4) mm, 

obtuse to subtruncate at apex, cuneate at base, 

deeply alveolate, slightly transversely rugose, 

glistening, pale brown to brown or red-brown; 

style base short, deltate, flattened (2-sided), 0.2- 

0.5 mm long, 0.5-1 mm wide at base, brown; 

bristles 6, stiffly erect, antrorsely barbed, reddish, 

varying in length, reaching ½ length of achene 

body to equaling or shortly exceeding the tip of 

style base. 


States west to Texas, Belize, and the Greater 

Antilles. 

Distribution in Puerto Rico: On wet sand of 

savanna and thickets, Laguna Tortuguero and 

vicinity in the municipalities of Manatí and Vega 

Baja. 


Manatí: Laguna Tortuguero, N.L. Britton & E.G. 

Britton 8128 (NY, US). Vega Baja: Tortuguero 

Lake, González-Más & Seda 2171 (MAPR); 

González-Más 1082 (NY), 2173 (US). 

20. Rhynchospora nitens (Vahl) A. Gray, Manual, 

ed. 5. 568. 1867; Scirpus nitens Vahl, Enum. 

Pl. 2: 272. 1805; Psilocarya nitens (Vahl) A. 

W. Wood., Amer. Bot. Fl. 364. 1870. Type: 

United States; “Carolina,” Bosc s.n. 

(holotype: probably at C-Vahl). 

Psilocarya portoricensis Britton, Bull. Torrey Bot. 


Laguna Tortuguero. Britton et al. 3850 

(holotype: NY!).


Caespitose annual, extremely variable 

vegetatively in size, 4.5-85 cm tall; roots fine to 

medium. Culms erect to ascending, 0.5-4.5 mm 

wide, compressed-trigonous, obtusely 4-angled 

(rectangular), or subterete, soft, surface appearing 

roughened under high magnification due to very 

fine cellular reticulation, finely and coarsely 

longitudinally ribbed, deeply longitudinally 

channeled along one side, remotely scabrous along 

edges of channel, pale green to green, glabrous. 

Leaves numerous, ascending, basal and cauline, 

the upper ones (sheathing bracts) subtending 

inflorescence corymbs; sheaths short, herbaceous, 

inflated, finely veined, green, glabrous; ligule 

absent; blades narrowly linear, 4-58 cm × 1-7 (-8) 

mm, flattened proximally, often folded or 

subinvolute distally, finely veined, midvein 

indistinct on adaxial side of blade, surface 

appearing roughened due to very fine cellular 

reticulation, remotely scabrous to smooth on 

margins and midvein abaxially, regularly scabrous 

on margins and midvein distally near apex, green, 

glabrous. Inflorescence a series of (1-) 2-4 simple 

to compound corymbs of ascending rays from the 

upper sheathing bracts, the terminal corymb 0.7- 

19 × 0.8-15 cm wide; sheathing bracts reduced 

distally, generally narrower and shorter than leaf 

blades; spikelets ovoid to oblong-ovoid with acute 

apex, (3.8-) 4-8 (-9) × (1.8-) 2-2.8 (-3) mm, acute 

at apex, broadly obtuse to sub-rounded at base, 

with only one sterile basal scale above prophyllar 

bract; fertile scales numerous, ovate to ovatedeltate, 

2.1-3 × (1.5-) 1.7-2.2 mm, thin, somewhat 

translucent, widely boat-shaped, dorsally acute to 

obtuse, minutely longitudinally striate, medium 

brown to reddish brown with very fine dark brown 

or blackish lineations, glabrous, margins entire, 

the narrow midcosta distally pale brown and 

proximally dark brown, often extending beyond 

the acute apex as a short mucro on proximal scales, 

lateral nerves indistinct. Stamens 2, the anthers 

0.7-1 mm long; style 2-branched. Achene widely 

biconvex, obovate to broadly obovate or 

subrounded, broadly rounded to subtruncate at 

apex, obtuse to sub-rounded at base, 1-1.1 × 1 mm, 

transversely rugose, cellular-reticulate at base, 

lustrous, yellowish brown to golden brown, 

maturing to dark reddish brown or brown-black, 

the rugae often lighter in color; style base 

depressed-triangular, mustache-shaped, 2-lobed at 

base, 0.2-0.6 mm long, about as wide as achene 

body at its base, crustose, pale brown to 

subwhitened; bristles absent 

General distribution: Eastern and 

southeastern United States west to Texas, Belize, 

Nicaragua, and the Greater Antilles. 

Distribution in Puerto Rico: In wet muddy 

areas over white silica sand, Laguna Tortuguero 

and vicinity, in the municipalities of Manatí, Vega 



Manatí: W side of Rt. 687, SE side of Laguna 

Tortuguero, Strong et al. 415 (GMUF). Vega Alta: 

Bo. Sabana, near drainage canal, ca. 1.8 km due 

NE of jct. Rds. 690 and 691 (Regadera), Proctor 

et al. 47743 (US); Proctor & Thomas 44143 

(US).Vega Baja: Bo. Algarrobo, just S of Laguna 

Tortuguero, Proctor et al. 45032 (US). 

21. Rhynchospora odorata C. Wright ex Griseb., 

Cat. Pl. Cub. 242. 1866. Type: Cuba. Wright 

3787 (lectotype: US!; isolectotypes: GH, 

NY!), here designated. 

Rhynchospora marisculus sensu Britton & P. 


Nees von Esenbeck, 1842. 

Perennial, 0.7-1.5 (-1.8) m tall; rhizome stout, 

emitting short, scaly stolons to 4 cm long; roots 

medium to coarse, to 1.5 mm thick. Culms 

ascending, stout, 1.5-5.5 (-6) mm wide, trigonous, 

hardened, finely ribbed, smooth, green, glabrous. 

Leaves arched-ascending, spreading, basal and 

cauline, the cauline reduced distally on culm; 

sheaths short on basal leaves, herbaceous, finely 

veined, pale brown proximally, light green distally, 

glabrous; ligule absent; blades linear, 7-67 cm × 

(3-) 4-8 (-10) mm, flattened, attenuate to 

triquetrous tip, finely veined abaxially, very finely 

cellular-reticulate adaxially, margins and abaxial 

midvein antrorsely scabrous (at least distally and 

at apex), essentially smooth, green, glabrous. 

Inflorescence a terminal and 1-4 (5) lateral, remote 

(at least proximally), compound, corymbiformfasciculate 

partial panicles from the upper 

sheathing bracts; terminal partial panicle 3-10 

(-15) × 2-5 cm; spikelets ovoid, 5.6-8 × 1.5-2.6 

mm, acuminate at apex, short-attenuate to abruptly 

narrowed at base, with 7-10 scales; fertile scales 

3-5, widely ovate-elliptic, 3.5-6.5 × 2.2-3.4 mm, 

deeply boat-shaped, curvate-keeled, dorsally 

obtuse, thinly herbaceous, finely cellular striate,

354 


semi-glossy, ferrugineous, glabrous, margins 

escarious or at most very narrowly so, midcosta 

fine, pale brown to brown, extended beyond 

acuminate to attenuate slightly recurved apex as a 

0.3-1 mm long awn. Stamens 3, sometimes 

marescent, the anthers of lower fertile flowers 1.7- 

3.3 mm long, those of terminal scales often twice 

as small, apiculate, truncate and minutely lobed 

at base; style deeply 2-branched. Achene biconvex, 

tumid, rounded-obovate, 1.6-2 (-2.1) (including 

stipe) × (1.3-)1.4-1.6 mm, rounded to a truncate 

apex, shortly attenuate at base to a 0.5 mm long 

glossy reddish brown stipe, transversely rugulose, 

finely cellular oblong-cancellate between the 

rugae, dull, brown or dark brown; style base 

shallowly deltate with an ellipsoid rim at base (as 

seen from above), 0.6-1.1 mm long, 0.9-1.1 mm 

wide at base, subflattened, with setose margins, 

brown, often with a crusty whitened surface; 

bristles 6, ascending to spreading, subulate, finely 

antrorsely barbed, setose at base, reddish, usually 

well exceeding the tip of style base, to twice length 

of achene. 


States and the Greater Antilles. 

Distribution in Puerto Rico: On wet white 

silica sand in open areas or among thickets, Punta 

Cangrejos and Laguna Tortuguero and vicinity. 

Carolina, Manatí, and Vega Baja. 

Note: Wright in Sauvalle, Fl. Cub. (179: 1868) 

cited R. odorata with his collection number 3787. 

However, the original protologue for this taxon 

appeared earlier in Grisebach’s Cat. Pl. Cub. (242: 

1866) where no collection number is given for the 

cited material: “Cuba occ. pr. Hanabana, in 

humidis, (Wr. a. 1865)”. The locality of this 

material is unknown, so the specimen cited later 

by Wright (Wright 3787) is formally chosen here 

as lectotype for R. odorata. 


Carolina: Punta Cangrejos, Stevenson 1706 (US). 

Manatí: Tortuguero, Woodbury s.n. (NY, US). 

22. Rhynchospora plumosa Elliott, Sketch Bot. 

S. Carolina 1: 58. 1816. Type: United States; 

Florida. Baldwin s.n. (holotype: PH; isotype: 

NY). 


rhizome short, thickened, knotty; roots medium, 

to 1 mm thick. Culms erect, 0.4-1.5 mm wide, 

slender, terete proximally to compressed-trigonous 

with rounded angles distally, finely ribbed, smooth 

proximally to sparsely scabrous on margins 

distally at apex, pale brown to brown proximally, 

green distally, glabrous. Leaves numerous, 

ascending or sometimes arching, primarily basal, 

1-3 cauline; sheaths short, herbaceous, finely 

veined to coarsely so on basal sheaths, pale brown 

to brown or sometimes dark brown on basal leaves, 

glabrous; ligule absent; blades filiform, 2-40 cm 

× 0.2-1.5 (-1.7) mm, crescentform proximally to 

crescentform-capillary distally, articulated towards 

apex and becoming flattened-trigonous to 

flattened, attenuate to apex, margins smooth 

proximally, antrorsely scabrous distally above 

articulation, finely veined, smooth, green, 

glabrous. Inflorescence a single terminal small 

contracted corymbiform panicle at apex of culm, 

rarely a second lateral one below, 0.6-4 × 0.4-1 

(-1.5) cm, subtended at base by a single sheathing 

bract, overtopping panicle, to 12 cm long; spikelets 

ovoid to ovoid-ellipsoid, (2.5-) 2.8-5 × 1-1.5 mm, 

short-attenuate to sharp tip at apex, cuneate to 

short-attenuate at base, in congested fascicles at 

branch tips, with 5-6 scales; fertile scales 1 (2), 

rounded-ovate or rounded-obovate, 1.7-3.7 (-4.2) 

× 1.4-2.5 (-2.7) mm, curvate-keeled, dorsally 

widely obtuse to rounded, finely cellular-striate, 

semi-glossy, dark brown, often dark brown on 

sides, glabrous, margins escarious or at most 

narrowly scarious, midcosta fine, indistinct 

proximally, pale distally, mucronulate at the obtuse 

to acute or sharply acute apex. Stamens 2-3, the 

anthers small, 0.3-0.7 mm long, apiculate, truncate 

at base; style 2-branched. Achene broadly 

biconvex (nearly rounded in cross section), 

obovoid, 1.4-1.8 × 1-1.3 mm, rounded at apex, 

cuneate at base, transversely rugulose to ridged, 

uniformly rusty brown; style base depressed, 

conical-apiculate or discoid, 0.2-0.5 mm long, 0.5- 

0.7 mm wide at base, articulated at base with 

achene, often with tufts of crystalline fimbrillae 

along each margin at base, light brown; bristles 6, 

silvery plumose to near apex, subulate and 

antrorsely barbed at apex, closely investing 

achene, slightly shorter than achene to exceeding 

the tip of the style base, incurved at apex, the tips 

of the longer ones often meeting just above apex 

of style base. 


States west to Texas, Central America, and the 

Greater Antilles.


Distribution in Puerto Rico: On wet, white 

silica sand in open areas, known only from northcentral 

Puerto Rico, in the vicinity of Laguna 

Tortuguero and Dorado. Dorado, Vega Alta, and 

Vega Baja. 


Vega Baja: Tortuguero, 12 Mar 1960, Woodbury 

s.n. (US); 1970, Woodbury s.n. (US); Jan 1975, 

Woodbury s.n. (NY, US). 

23. Rhynchospora pusilla Chapm. ex M. A. 

Curtis, Amer. J. Sci. Arts 57: 409. 1849. Type: 

United States; Florida. Chapman 4473 


Rhynchospora intermixta C. Wright in Sauvalle, 

Anales Acad. Ci. Méd. Habana 8: 88. 1871. 

Type: Cuba; Pinar del Río. Wright 3794 

(holotype: GH; probable isotype: NY). 

Rhynchospora bruneri Britton in Britton & P. 


Puerto Rico; Luquillo Mountains. Britton & 

Bruner 7630 (holotype: NY!; isotypes: UPR!, 

US!). 


inconspicuous; roots fine, capillary. Culms 

ascending, 0.3-0.7 mm wide, filiform, trigonous 

with blunt angles proximally to compressed 

trigonous or subterete distally, delicate, flexuous, 

finely ribbed, smooth, green, glabrous. Leaves 

numerous, ascending, basal and 1-2 lower cauline; 

sheaths short, herbaceous, finely veined, pale 

brown, glabrous; ligule absent; blades short, 2-25 

cm × 0.2-0.7 mm, setaceous, involute proximally, 

becoming filiform and canaliculate, flexuous, 

often curving or wavy, attenuate to triquetrous or 

subflattened tip, finely veined abaxially, finely 

cellular reticulate adaxially, antrorsely scabrous 

on margins distally, green, glabrous. Inflorescence 

a terminal and series of 1-2 lateral, small, open or 

subdense, compound, partial corymbiform 

panicles from the upper sheathing bracts; terminal 

corymb 1-2.5 × 0.4-1 cm, the lateral corymbs 

smaller, on short peduncles; sheathing bracts 

equaling to shortly exceeding corymbs, to 3 cm 

long; spikelets ovoid, terete, 1.8-2.6 (-3) × 0.5-1 

mm, acute to narrowly acute at apex, cuneate at 

base, fasciculate at tips of short primary branches, 

rarely solitary, short-pedicellate, with 4-8 scales; 

fertile scales 2-5, ovate to broadly ovate or 

rounded-ovate, 1.4-1.8 × 1.2-1.6 mm, deeply boat- 

shaped, curvate-keeled, dorsally obtuse to 

subrounded, submembranous, finely cellularstriate, 

semi-glossy, brown, margins scarious, 

often undulate, midcosta slender, pale, extended 

beyond the obtuse to sub-rounded apex as a short 

mucro. Stamens 1-3, the anthers 0.4-1.2 mm long, 

bluntly apiculate, truncate at base; style 2branched. 

Achene biconvex, oblong-obovate, 0.7- 

0.9 × 0.5-0.6 mm, rounded at apex, attenuate to a 

short stipe at base, transversely rugulose, 

stramineous to brown; style base short bulbiform, 

0.1-0.2 mm long, 0.3-0.4 mm wide at base, reddish 

brown; bristles absent. 


States west to Texas, Mexico (Chiapas, Tabasco), 


Distribution in Puerto Rico: White silica 

sands of Laguna Tortuguero and vicinity and in 

exposed rock crevices on and near summit of El 

Yunque in the Luquillo Mountains. Río Grande, 


Note: The original description of this taxon 

by Curtis (1849) was barely validly published, only 

2 explicit characters were given for it. 


Vega Baja: Laguna Tortuguero, Sep 1960, 

Woodbury s.n. (NY, US); Liogier & Martorell 

34636 (NY); González-Más & Woodbury 3686 

(MAPR); Liogier & Martorell 34559 (UPR); 

Breckon et al. 5804 (US). Río Grande: Sierra de 

Luquillo, Caribbean National Forest, Roca El 

Yunque, Proctor 40814 (US); Gleason & Cook M- 

143 (NY); N.L. Britton & Bruner 7630 (NY). 

24. Rhynchospora racemosa C. Wright in 

Sauvalle, Anales Acad. Ci. Méd. Habana 8: 

86. 1871. Type: Cuba. Wright 727 (holotype: 

probably at GOET; isotypes: GH!, S, US!). 

Rhynchospora polyphylla sensu Grisebach, Cat. 

Pl. Cub. 246. 1866, non (Vahl) Vahl, 1805. 


rhizome short, hardened, nodose, fibrillose; roots 

medium to somewhat coarse, to 1 mm thick. Culm 

erect to ascending, borne loosely or tightly 

crowded together along rhizome, slender, 0.8-2.3 

mm wide, often narrowing to 0.5 mm distally, 

obtusely trigonous proximally, trigonous to 

triquetrous distally, densely nodose, covered in leaf 

sheaths much of its length, hardened proximally, 

firm and flexuous distally, coarsely to finely

356 


ribbed, finely roughened or scabridulous at least 

distally, smooth proximally, golden brown at nodes 

proximally, green distally. Leaves 13-40, 

ascending, spreading, primarily cauline, those at 

base reduced or represented by sheaths only; 

sheaths short, herbaceous, finely veined, light 

brown proximally on basal sheaths, grayish green 

on cauline sheaths, scaberulous and often 

appressed to spreading short-pubescent distally 

(particularly on inner band) to glabrescent, the 

inner band reddish or yellowish brown with minute 

shiny dark red or purplish red lineations, 

muriculate at orifice; ligule essentially absent, but 

often a thickened band of tissue at adaxial junction 

of sheath and blade; blades linear, 9-60 cm × 1.3- 

4 mm, flattened proximally to plicate distally, longacuminate 

to triquetrous tip, finely veined 

abaxially, somewhat glossy and smooth adaxially 

with indistinct veins, margins and abaxial midvein 

finely antrorsely serrate-scabrous, grayish green, 

essentially glabrous, short pilose at base abaxially 

at junction with sheath. Inflorescence a terminal 

and series of 3-7 remote, strict, racemose partial 

panicles from the upper sheathing bracts; sheathing 

bracts leaf-like proximally to linear setaceous 

distally on culm, exceeding subtending partial 

panicles proximally, shorter than to equaling 

terminal partial panicles; terminal partial panicle 

largest, 3-7 × 0.6-1 cm, on filiform peduncles 2-6 

cm long; spikelets narrowly ellipsoid-lanceoloid, 

5.5-8.5 (-9) × 1-1.5 mm at anthesis and early stages 

of maturity, at later stages of maturity, rachilla 

elongates up to 20 mm long and is distinctly 

anfractuose, acuminate at apex, cuneate at base, 

with 6-7 (-8) scales, subsessile, solitary or in 

fascicles of 2-3 at branch tips; fertile scales 2-3, 

ovate-lanceolate to lanceolate, (3-) 3.5-6 (-6.2) × 

(1.8) 2-2.6 (-2.8) mm, slightly curvate-keeled, 

dorsally obtuse to subrounded, thinly herbaceous, 

semi-glossy, stramineous or stramineousferrugineous, 

often sparsely reddish lineolate, 

glabrous, margins narrowly scarious, often 

undulate, midcosta fine, inconspicuous, extended 

beyond acute apex as a 0.4-0.6 mm long antrorsely 

scabrous mucro. Stamens 2-3, the anthers 2-3 mm 

long, with triangular-papillate appendage at apex, 

truncate at base with minute papillae or lobes; style 

entire or shortly 2-branched at tip. Achene 

biconvex, subtumid, oblong-obovate, (1.4) 1.5-1.9 

× 1-1.3 mm, obtuse or subtruncate at apex, cuneate 

at base, cellular-cancellate with 15-18 (-20) 

longitudinal rows of cells on each face, pale brown 

to brown or brownish black, shiny at maturity; 

style base oblong-lanceolate, mitriform, (0.8-) 1- 

1.5 mm long, 0.6-0.8 mm wide at base, compressed 

or subflattened, with truncate to sub-rounded or 

acute tip, brittle, pale grayish green; bristles 

wanting or 2-4, often rudimentary, 1-2 sometimes 

reaching 2/3 length of achene, subulate, antrorsely 

barbed, translucent, yellowish or reddish. 


Distribution in Puerto Rico: On serpentine 

soils; steep, often rocky, wooded slopes and 

roadside banks. Maricao, Sabana Grande, and 

Yauco; known only from the southwestern region. 


Maricao: Bo. Maricao Afuera, González-Más 3363 

(MAPR); Rosario to Maricao road, González-Más 

2424 (MAPR); Sabana Grande: Maricao State 

Forest, Proctor & McKenzie 43827 (US); Susúa 

Forest Reserve, Breckon & Weaver 6471 (MAPR). 

Yauco: Bo. Susua Alta, García & Caminero 3145 

(MAPR). 

25. Rhynchospora radicans subsp. 

microcephala (Bertero ex Spreng.) W.W. 

Thomas, Mem. New York Bot. Gard. 37: 60. 

1984; Dichromena microcephala Bertero ex 

Spreng., Syst. Veg. 1: 202. 1824; 

Rhynchospora microcephala (Bertero ex 

Spreng.) Kük., Bot. Jahrb. Syst. 75: 311. 1951, 

non Britton ex Small, 1903. Type: Jamaica. 

Bertero s.n. (holotype: TO; photo at MICH; 

isotype: MO). 

Dichromena radicans sensu Urban, Symb. Antill. 

4: 121. 1903; and sensu Britton & P. Wilson, 

Bot. Porto Rico 5: 99. 1923, non 

Schlechtendal & Chamisso, 1831. 

Caespitose annual or short lived perennial, 10- 

63 cm tall; roots fine. Culms arching to erect, 0.6- 

1.7 (-2) mm wide, firm and flexuous, triquetrous 

to obtusely so, compressed distally, coarsely and 

finely ribbed, smooth, green, glabrous, often 

elongating and lopping over to the ground at 

maturity, the inflorescence heads proliferating. 

Leaves (2-) 3-4 (-5), ascending, primarily cauline 

or from sterile shoots, the basal leaves, when 

present, often with reduced blades or represented 

by bladeless sheaths; sheaths loose, short to 

medium in length, finely veined with pale veins, 

green distally, pale green proximally, glabrous; 

ligule absent; blades linear, herbaceous, 5-30 cm 

× 1.4-5 mm, flattened to slightly involute, finely


nerved, distinctly so abaxially, with whitened to 

pale green nerves, often ciliate marginally at base, 

green to dark green, essentially smooth, glabrous. 

Inflorescence a conical to hemispherical head of 

3-6 (-7) spikelets at the summit of the culm, 0.8- 

1.5 cm in diam., the central spikelet largest; 

involucral bracts (3-) 4-5 (-6), leaf-like, spreading 

to reflexed, exceeding the inflorescence, ciliate 

basally along margins, the basal bract longest, 4- 

15 cm long, 1.5-4.5 mm wide; spikelets 3-6 (-7), 

ovoid, 7-12 × 2-4.3 mm, acute to acuminate at 

apex, rounded at base, with ca. 20-30 scales; scales 

all fertile except for basal scale of spikelet, ovate 

to ovate-lanceolate, 4.2-5.2 × 1.8-3.6 mm, boatshaped, 

dorsally obtuse to rounded, 

submembranous, margins broadly scarious, 

whitish or stramineous to pale brown, reddish 

lineolate, glabrous, the midcosta fine, 

inconspicuous, pale green, extended beyond the 

narrowly acute to acuminate apex as a very short 


apiculate, truncate at base with minute crystalline 

papillae or glands; style 2-branched. Achene 

biconvex, very widely obovate, 0.9-1.2 × 0.8-1.4 

mm, broadly rounded at apex, cuneate to widely 

cuneate or sub-rounded at base, transversely 

rugulose, stramineous to brownish orange or bony 

white; style base very shallowly triangular to 

shallowly lunate, 0.1-0.4 mm long, 0.6-1.2 mm 

wide at base, brownish to stramineous or greenish; 



America, West Indies, western South America, and 

sporadically to the Guianas and Amapá, Brazil. 

Recently recorded from Hawaii. 


Islands: On clay soils and limestone substrate, in 

open areas and clearings in wet mountain forest, 

swampy areas, lawns, roadsides and disturbed 

habitats. Adjuntas, Añasco, Arecibo, Bayamón, 

Cabo Rojo, Caguas, Canóvanas, Cayey, Ceiba, 

Fajardo, Guayama, Hatillo, Jayuya, Juana Díaz, 

Las Marias, Luquillo, Maricao, Maunabo, 

Mayagüez, Naguabo, Orocovis, Ponce, Río 

Grande, San Germán, San Lorenzo, Utuado, Vega 

Baja, Villalba, and Yabucoa; St. Thomas and 

Tortola. 



Britton et al. 5393 (NY, US). Añasco: new road 

#2, González-Más 1457 (MAPR). Arecibo: Bo. 

Río Arriba, Axelrod 7985 (HAC, NY, UPRRP, US). 

Bayamón: Johnston & Stevenson 1140 (NY, US). 

Caguas: Beatriz de Caguas, Goll 420 (US). 


Condesa Section, Axelrod & Schulz 5637 

(UPRRP). Cayey: Carite Forest Reserve, Axelrod 

& Axelrod 4347 (NY, UPRRP, US). Ceiba: 

Luquillo Mountains, Acevedo-Rdgz. et al. 10762 

(US). Fajardo: Río Arriba, N.L. Britton & Shafer 

1676 (NY, US). Guayama: Carite Forest Reserve, 

Axelrod et al. 1009 (UPRRP). Juana Díaz: Bo. 

Collores, dirt road from end of Rt. 512, Axelrod & 

Sastre 5257 (UPRRP, US). Las Marías: La Juanita, 

near Las Marías, N.L. Britton et al. 3896 (NY, US). 

Luquillo: El Verde Field Station, Taylor 10130 

(NY, UPRRP). Maricao: Fish Hatchery, González- 

Más 402 (MAPR, NY). Maunabo: La Pica, road 

3, km 106.2, González-Más 1337 (MAPR, NY). 

Mayagüez: Las Mesas, near Mayagüez, Holm 126 

(US); Near Las Vegas, Fredholm 4421 (MAPR). 

Naguabo: Sierra de Naguabo, Bo. de Maizales, 

N.L. Britton & Cowell 2181 (NY, US); Orocovis: 

Bo. Barros, Axelrod & Rogowitz 9974 (UPRRP). 

Pico del Este, Luquillo Mts, Liogier et al. 33266 

(NY). Ponce: On the Adjuntas Road, 8 mi. from 

Ponce, Heller 6301 (MO, NY, US). Río Grande: 

El Verde Research Station, Rt. 186 at the Río 

Sonadora, Taylor 11684 (MO), 11798 (MO, 

UPRRP); El Yunque, Hioram 363 (NY, US). San 

Germán: Rosario, Stevens & Hess 3809 

(MAPR).Utuado: Vicinity of Utuado, N.L. Britton 

& Cowell 982 (NY). Vega Baja: Stevens & Hess 

2503 (MAPR). Villalba: Toro NegroReserve 

Forest, González-Más 2005 (MAPR). Yabucoa: 

Cuchilla de Panduras, Axelrod et al. 1367 

(UPRRP). ST. THOMAS: Signal gorges, Eggers s.n. 

(MO). TORTOLA: Road Town to High Bush, N.L. 

Britton & Shafer 769 (US); Sage Mountain, 

D’Arcy 758 (MO). 

26. Rhynchospora rariflora (Michx.) Elliott, 

Sketch Bot. S. Carolina 1: 58. 1816; Schoenus 

rariflorus Michx., Fl. Bor.-Amer. 1: 35. 1803; 

Rhynchospora micrantha Vahl, Enum Pl. 2: 

231. 1805, nom. illeg. Type: United States; 

Georgia. Michaux s.n. (holotype: P, photo at 

GH; isotype fragment: NY). 

Rhynchospora setacea Vahl, Enum. Pl. 2: 233. 

1805. Type: South America. Collector 

unknown (holotype: C-Vahl). 

Caespitose perennial, (15-) 20-72 cm tall; 

rhizomes slender and knotty, stiff, often forming

358 


a dense mass or sometimes short horizontally 

creeping; roots medium, to 0.7 mm thick. Culms 

ascending or reclining, 0.3-0.8 (-1) mm wide, 

trigonous with blunt angles or subterete, wiry, 

flexible, delicate, finely ribbed, smooth except for 

antrorsely scabrous margins of channel distally, 

green, glabrous. Leaves numerous, ascending, 

basal; sheaths short, the lowermost bladeless or 

nearly so, herbaceous, finely veined, stramineous 

proximally, pale brown or green distally, glabrous, 

the inner band brown with minute blackish spots 

at orifice; ligule absent; blades filiform, 4-40 cm 

× 0.2-1 mm (flattened), often curving or wavy with 

age, grading from involute to subterete and 

canaliculate, long-attenuate to triquetrous 

antrorsely scabrous apex, finely veined abaxially, 

finely cellular-reticulate adaxially, smooth, 

margins scabrous (at least distally), green, 

glabrous. Inflorescence a terminal and 1-2 lateral, 

remote, small, open, corymbiform panicles of 2- 

13 (-23) spikelets from the upper sheathing bracts, 

1-5 × 1-2 (-3) mm, the capillary branchlets 

elongating, ascending to spreading, with solitary 

spikelets terminating branch tips; spikelets ovoid 

to broadly ovoid or ovoid-ellipsoid, 3.5-4.5 (-5) × 

1.5-1.8 mm, acute at apex, shortly attenuate at 

base, with 5-8 scales; fertile scales 3-5, ovate to 

rounded-ovate, 2-3.7 (-4) × 1.9-3.2 (-3.5) mm, 

curvate-keeled, dorsally obtuse, thinly herbaceous, 

finely cellular-striate, semi-glossy, light to dark 

brown, often blackish brown-lineolate on sides, 

margins narrowly scarious, often undulate, 

midcosta very fine, indistinct, extending beyond 

acute to obtuse apex as a short mucro. Stamens 3, 

the anthers 1.5-2.5 mm long, apiculate, truncate 

at base with minute papillae or lobes; style 2branched. 

Achene biconvex, obovate to oblongobovate, 

(1.3-) 1.4-1.7 × (1.0-) 1.1-1.4 mm, 

rounded to truncate apex, short attenuate at base, 

transversely rugulose, cellular oblong-cancellate 

between the rugae, two small whitish ascending 

tongues of spongy tissue prominent on either side 

at the base, light or dark brown, dark reddish 

brown at base; style base deltate, 0.4-0.7 mm long, 

0.7-1 mm wide at base, flattened, brittle, brown, 

often with a crusty whitened surface; bristles 5-6, 

subulate, antrorsely barbed, setose at base, reddish 

brown, unequal in length, the longest equaling or 

only slightly exceeding middle of achene. 

General distribution: Eastern and 

southeastern United States west to Texas, Central 

America, and the Greater Antilles. 

Distribution in Puerto Rico: On wet to moist 

white silica sand in open areas, Dorado and Laguna 

Tortuguero. Dorado and Vega Baja. 


Dorado: 3 Dec 1960, Woodbury s.n. (NY). Vega 

Baja: Tortuguero, 17 May 1960, Woodbury s.n. 

(NY, US). 

27. Rhynchospora recognita (Gale) Kral, Novon 

9: 205. 1999; Rhynchospora globularis var. 

recognita Gale, Rhodora 46: 245. 1944. Type: 

United States; Virginia. Fernald & Long 6070 

(holotype: GH; isotypes: P, PH, US!). 

Schoenus cymosus Muhl., Descr. Gram. 8. 1817, 

non Willdenow, 1797. 

Rhynchospora cymosa sensu Torrey, Fl. N. Middle 

United States 1: 56. 1823; sensu Urban, Symb. 

Antill. 4: 124. 1903, and many authors 

thereafter, non Elliott, 1816. 

Caespitose perennial, in small clumps, 10-102 

cm tall; rhizome short, inconspicuous. Culms 

stiffly erect, firm to subhardened, slender, 

trigonous, obtusely trigonous, or subterete, 

smooth, finely ribbed, green, glabrous, minute 

whitish punctations often evident on culms and 

leaves. Leaves numerous, ascending, primarily 

basal, 3-5 cauline, the blades of basal leaves often 

curling; sheaths short, herbaceous to thickly 

herbaceous, finely veined, green to pale green or 

stramineous, glabrous; ligule absent; blades 

narrowly linear, 3-36 cm × 1-4.5 mm, flattened to 

V-shaped or subplicate, finely veined abaxially, 

smooth, semi-glossy and finely cellular-reticulate 

adaxially, attenuate to triquetrous apex, margins 

and abaxial midvein essentially smooth 

proximally, antrorsely scabrous distally, green, 


1-3 remote lateral corymbose panicles of spikelets 

from the upper sheathing bracts; terminal panicle 

1-6 × 1-3 cm, the smaller lateral panicles remote, 

the lowermost on long slender peduncles, the 

spikelets in fascicles at branch tips; spikelets 

broadly ovoid, 3-4 × 1.5-2 mm, acute at apex, acute 

to obtuse at base, with 5-6 scales; fertile scales 2- 

3, ovate to subrounded, deeply boat-shaped, 1.8- 

3 × 2-3.2 mm (flattened), curvate-keeled, dorsally 

obtuse to subrounded, thinly herbaceous, finely 

cellular-striate, dull, brown, often tinged with dark 

brown or brown-black on sides, margins scarious, 

erose, midcosta slender, pale brown, extending


beyond obtuse to subemarginate apex of scale as 

a short mucro. Stamens 3, the anthers 0.5-2 mm 

long, apiculate, truncate with minute glands or 

papillae at base; style 2-branched. Achene thickly 

biconvex, broadly obovoid to subglobose, 1.3-1.8 

× 1.2-1.5 mm, rounded at apex, acute to subobtuse 

and very short-stipitate at base, transversely 

rugulose, yellow-brown to dark brown; style base 

depressed-conical to triangular, 0.3-0.8 mm long, 

0.6-0.9 mm wide at base, subflattened, abruptly 

flaring to an elliptic base as seen from above, 

brittle, light brown to brown; bristles 5-6, subulate, 

antrorsely barbed, reddish, extending ½ to 2 /3 

length of achene body. 

General distribution: North central and 

southeastern United States west to Texas and 

California, Mexico (Chiapas), Central America, 

and the West Indies. 

Distribution in Puerto Rico: Occurs on red 

laterite soils or silica sand deposits; savannas, 

grassy hillsides, slopes, fields, and seepage areas 

at lower and middle elevations. Barranquitas, 

Maricao, Mayagüez, Patillas, Río Grande, San 

Juan, Vega Baja, and Yabucoa. 


Barranquitas: Monte Torrecilla, N.L. Britton et al. 

5646 (NY, US). Maricao: Bosque de Maricao, 

González-Más 3864 (MAPR). Mayagüez : Cerro 

Las Mesas, N.L. Britton & Hess 2717 (NY, US); 

Cerro de Las Mesas, Hess & Stevens 3852 (NY); 

Las Mesas, Holm 82a (US), 317 (NY); Liogier et 

al. 30529 (NY). Río Grande: Rd. to East Peak, 

Woodbury s.n. (UPR). San Juan: Santurce, Heller 

& Heller 13 (NY, US). Vega Baja: Tortuguero 

(east), 15 May 1977, Woodbury s.n. (NY, UPR). 

Yabucoa: Guayanes, Woodbury s.n. (NY, UPR). 

28. Rhynchospora rugosa (Vahl) Gale, Rhodora 

46: 275. 1944; Schoenus rugosus Vahl, Eclog. 

Amer. 2: 5. 1798; Rhynchospora glauca Vahl, 

Enum. Pl. 2: 233. 1805, nom. illeg. Type: 

South America; Boca Chica no.27. Rohr s.n. 

(holotype: C-Vahl, photo at F, US). 


rhizome short, knotty; roots coarse. Culms erect, 

(0.8-) 1.2-2.5 (-3) mm wide, trigonous to obtusely 

trigonous, very firm, finely ribbed and channeled, 

smooth, green, glabrous. Leaves 5-15, ascending, 

primarily basal, 2-5 cauline; sheaths short, finely 

ribbed abaxially, smooth, green, glabrous; ligule 

absent; blades , 9-55 cm × (1.3-) 1.7-4.5 (-5) mm, 

flattened to subplicate or folded, smooth, green, 

glabrous, margins and abaxial midvein smooth 

proximally to minutely antrorsely scabrous 

distally, long-acuminate to triquetrous apex. 


remote cymose-paniculate partial panicles from 

the upper sheathing bracts; panicles narrowly 

obtriangular, 1-8 (-11) × 0.8-2.7 (-3.5) cm, the 

spikelets in fascicles at branch tips; spikelets 

ovoid-ellipsoid, 3.5-4.7 (-5) × 1.1-1.7 mm, acute 

at apex, short-cuneate at base, with 6-9 scales; 

fertile scales 3-5, spreading with developing 

achenes, ovate to oblong-ovate, (2.3-) 2.5-3.5 × 

1.8-2.8 mm, thinly herbaceous, dorsally obtuse to 

subrounded, finely cellular-striate, brown to dark 

brown or brownish black, glabrous, finely and 

indistinctly veined, margins broadly scarious, 

often erose, midcosta narrow, pale brown, 

extending beyond the acuminate apex as a 0.1-0.5 

mm long mucro. Stamens 3, the anthers 0.6-1.7 

mm long, apiculate; style 2-branched. Achene 

biconvex, obovate to elliptic-obovate, 1.6-1.9 × 

1-1.3 mm, cuneate at base, truncate at apex, 

transversely rugulose, light brown or yellowish 

brown; style base triangular, 0.6-1 mm long, 0.6- 

0.8 (-1) mm wide at base, ca. 2/3 as wide as achene, 

minutely antrorsely scabrous along margin, pale 

brown to dark brown; bristles 6, antrorsely barbed, 

equaling to exceeding apex of style base, often 

setose at base. 


America, West Indies, and tropical South America. 

Distribution in Puerto Rico: Occurs in wet to 

moist, rich laterite soils, red clays, or silica sand 

deposits, in savannas, thickets, pastures, open 

places, mountain slopes, along streams, and steep 

exposed roadside banks. Aguada, Bayamón, 

Cataño, Ceiba, Cidra, Lares, Las Piedras, Luquillo, 

Manatí, Mayagüez, Moca, Naguabo, Río Grande, 

San Juan, San Sebastián, and Vega Baja. 


Aguada: Sintenis 5742 (US). Cataño: back of 

Cataño, Heller 6410 (MO, NY, US). Cidra: Perkins 

s.n. (NY). Lares: Sintenis 5901 (MO, NY, US). 

Luquillo: Pico del Este, Mts., Liogier et al. 33284 

(NY). Mayagüez: Las Mesas, Holm s.n. (US); 

Liogier et al. 30539, 30691 (NY). Moca: road 110, 

km 9.2, González-Más 1808 (MAPR, NY, US). 

Naguabo: Sierra de Naguabo, Río Icaco and 

adjacent hills, Shafer 2534 (NY, US). Naguabo/

360 


Río Grande: Caribbean National Forest, along 1 

km stretch of Tradewinds Trail, Axelrod 6913 

(UPRRP, US), 6915 (NY, UPRRP). San Juan: Road 

from Río Piedras to Trujillo Alto, Hioram 806 (NY, 

US). San Sebastián: Sargent 213 (US). Vega Baja: 

Tortuguero, González-Más 4016 (MAPR). 

29. Rhynchospora tenerrima Nees ex Spreng., 

Syst. Veg. 4 (Curae posteriores): 26. 1827. 

Type: West Indies. “Nov. Holl.” [an error for 

West Indies], Kohaut s.n., distributed as F. 

Sieber agrost. no. 116 (holotype: AWH). 

Schoenus setaceus P. J. Bergius, Acta Helv. Phys.- 

Math. 7: 130. 1772; Rhynchospora setacea 

(P. J. Bergius) Boeck., Vidensk. Meddel. 

Dansk Naturhist. Foren. Kjøbenhavn 1869: 

159. 1869, non Vahl, 1805. Type: Surinam. 

Rolander s.n. (C-Rottb). 

Scleria setacea Poir. in Lamarck, Encycl. 7: 4. 

1806. Type: Puerto Rico. Ledrú s.n. (holotype: 

probably at P). 

Caespitose annual, 7-55 cm long; roots fine 

to medium. Culms ascending, 0.3-0.8 mm wide, 

obtusely trigonous, soft, flexuous, finely ribbed, 

minutely punctate distally, green, glabrous. Leaves 

numerous, ascending, primarily basal, 1-3 cauline; 

sheaths short, scarcely inflated, herbaceous, finely 

veined, pale brown, glabrous; ligule absent; blades 

filiform, 3-36 cm × 0.3-1.3 (-1.4) mm, 

crescentform-involute proximally, triangularchanneled 

distally, herbaceous, finely veined, 

minutely punctate, (at least distally), pale green, 

glabrous, margins essentially smooth proximally, 

antrorsely scabrous at apex, long-attenuate to 

triquetrous or subflattened apex. Inflorescence a 

terminal and 1-3 lateral, small, contracted, 

corymbose partial panicles from the upper 

sheathing bracts; terminal panicle 7-15 × 4-14 mm, 

the spikelets (1-) 2-10, rather congested; spikelets 

narrowly ovoid-ellipsoid to ellipsoid-lanceoloid, 

5-6 × 0.9-1.8 mm, acuminate at apex, cuneate at 

base, with 5-11 scales; fertile scales 3-6, ovateelliptic 

to ovate-lanceolate, (2.7-) 3-4.8 (-5) × 1- 

2.8 mm, obtusely angled dorsally, becoming subrounded 

with developing achenes, sterile and 

lower fertile scales thinly coriaceous, glabrous, 

glossy, somewhat uniformly light brown to brown, 

with darker reddish brown lineations, upper fertile 

scales submembranous, dull, margins broadly 

scarious with reddish lineations, midcosta very 

fine, pale, extending beyond acute to acuminate 

apex of scale as a 0.5-1 mm long antrorsely 

scabrous awn, lateral nerves indistinct. Stamens 

2, the anthers 1-1.6 mm long, apiculate, truncate 

with minute crystalline protuberances at base; style 

2-branched. Achene biconvex, obovate, 1.5-1.8 × 

1.1-1.5 mm, margins at apex flaring, prolonged 

into an ear-like projection on each side that is 

confluent with margins of style base, abruptly 

narrowing to a narrow, whitish translucent, 

cellular-reticulate stipe at base, transversely rugose 

medially, widely cellular-reticulate-papillate on 

margins, pale brown to brown, often with a gray 

or reddish longitudinal stripe medially; style base 

tounge-like, dialated at the tip as seen from dorsal 

or front view, pyramidal as seen from lateral view, 

the edges decurrent along edges of apical ears of 

achene, 0.4-0.6 mm long, 0.2-0.3 mm wide at base, 

soft, brittle, brown to brownish black; bristles 

absent. 

General distribution: Mexico (Chiapas, 

Tabasco), Central America, West Indies, and 

tropical South America. 

Distribution in Puerto Rico: In wet to moist 

sandy soils of open fields, pastures, open savanna, 

marshy or swampy areas, hillside seepage bogs, 

coastal swamps, and roadside banks and ditches 

at lower and middle elevations. Cabo Rojo, 

Cataño, Cayey, Dorado, Fajardo, Guayama, 

Humacao, Lares, Las Piedras, Manatí, Patillas, Río 

Grande, San Lorenzo, San Sebastián, Manatí, 

Mayagüez, Río Grande, San Juan, Vega Alta, Vega 

Baja, and Yabucoa. 


Cabo Rojo: Punta Arena, González-Más 2146 

(MAPR, NY, US). Cataño: Vicinity of Cataño, N.L. 

Britton et al. 6971 (NY). Cayey: Road 184, 

Guavate-Carite, km 2, González-Más 1570 

(MAPR, NY, US); Cerro La Santa, Guavate, 

Liogier et al. 29813 (NY). Dorado: Dorado to 

Cerro Gordo, Rt. 693, km 13, González-Más 307 

(MAPR, NY, US). Guayama-Patillas: Carite Forest 

Reserve, Axelrod & Alemán 9893 (UPRRP). 

Humacao: Punta Santiago, Liogier & Oquendo 254 

(UPR). Lares: Stevens & Hess 4942 (NY). Manatí: 

Bo. Tierras Nuevas Saliente, Proctor & 

Concepción 42201 (US). Mayagüez : Las Mesas, 

Holm 73 (MO); Guanajibo, near Mayagüez, N.L. 

Britton & Cowell 4075 (NY, US). Patillas: edge 

or Carite Reserve, Taylor 8230 (UPRRP). Río 

Grande: Río Espíritu Santo, Coco Beach, Liogier


et al. 33785 (NY); El Yunque, Fosberg 44199 

(US). San Juan: Santurce. Heller & Heller 599 

(NY, US). San Lorenzo: Guayabota, from San 

Lorenzo to Patilla, Liogier & Martorell 34329 

(MO, NY). San Lorenzo-Yabucoa border: Taylor 

7601 (NY). San Sebastián: Bo. Eneas, González- 

Más 924 (MAPR, NY, US). Vega Alta: Bo. Sabana, 

Proctor & Thomas 44147 (US). Vega Baja: 

Tortuguero Lagoon, González-Más 1051 (NY, 

US); Liogier & Martorell 34613, 34624 (NY). 

Yabucoa: Sintenis 5168 (US); Cerro de Pandura, 

Santa Elena, Liogier et al. 30991 (NY). 

30. Rhynchospora uniflora Boeck., Flora 63: 

439. 1880. Type: Brazil; Rio de Janiero. 

Glaziou 9336 (holotype: B, destroyed). 

Rhynchospora elongata Boeck., Beitr. Cyper. 1: 

26. 1888. Type: Puerto Rico. Sintenis 1360 

(isotype: S). 

Rhizomatous perennial forming dense 

clumps, (20-) 30-110 cm tall; rhizome short, 

nodose, woody; roots medium to coarse, 0.5-1.5 

mm thick. Culm erect, 0.7-1.5 (-2) mm wide, 

obtusely trigonous proximally to trigonous 

distally, stiff, internally soft and pithy, often hollow 

proximally, finely ribbed, essentially smooth, often 

antrorsely scabrous on angles distally, tan or light 

yellowish brown proximally and often with 

patches of brown at nodes, green distally, very 

finely and sparsely antrorsely setose-scabrous, 

glabrescent. Leaves numerous, ascending, dense, 

primarily cauline, the lower cauline blades often 

reduced, the mid to upper cauline elongate; sheaths 

elongate, herbaceous, finely veined, pale green or 

brown, very finely and sparsely antrorsely setosescabrous 

near orifice, glabrous proximally; ligule 

a very fine thickened band of tissue at adaxial 

junction of sheath and blade; blades narrowly 

linear, 10-40 (-50) cm × (0.8-) 1-2.5 (-3) mm, 

flattened to plicate, long-attenuate to triquetrous 

apex, acute to acuminate at apex on reduced lower 

cauline blades, finely veined abaxially with a pale 

distinct midcosta and two lateral costa, finely 

veined adaxially, margins and abaxial midcostae 

finely and closely antrorsely appressed setosescabrous 

(at least distally), bluish or grayish green, 

minutely pale green or whitish punctate, very 

finely and sparsely antrorsely setose-scabrous or 


1-5 lateral, often remote, small, simple to 

compound, fasciculate or loosely corymbosefasciculate 

partial panicles from the upper 

sheathing bracts; terminal partial panicle a single 

fascicle or compound corymb of 2-5 fascicles of 

spikelets, 1-5 × 1-2 cm, with 10-60 spikelets, the 

lateral smaller, on very slender peduncles to 5 cm 

long or sometimes very short peduncled and 

appearing sessile; spikelets lanceoloid to oblonglanceoloid, 

4-6 × 0.7-1.3 mm, acuminate at apex, 

acute to obtuse at base, with 6-8 scales, widely 

spreading with developing achenes; fertile scales 

4-6, 1-3 bisexual, 1-2 terminal scales of spikelet 

staminate only, ovate-lanceolate to lanceolate, 3- 

5 × 0.8-1.5 mm, shallowly boat-shaped, slightly 

curvate, dorsally obtuse to rounded, thinly 

herbaceous, glossy, pale reddish brown or 

brownish yellow, finely ferrugineous or brownish 

lineolate, margins entire, escarious, midcosta fine, 

pale green, extended beyond acute to acuminate 

apex as a short mucro. Stamens 3, the anthers 2-3 

mm long, glandular-apiculate, truncate at base with 

minute papillae of glands; style entire or shortly 

2-branched at apex. Achene biconvex, obovate, 

elliptic obovate, or rounded-obovate, 1.5-2.2 × 

0.9-1.6 mm, rounded at apex, obtusely narrowed 

to short-stipitate base, finely transversely rugulose 

to faintly so and appearing smooth, light brown to 

chestnut colored, dull to subglossy; style base 

conic-subulate, 1-1.5 mm long, 0.4-0.6 mm wide 

at base, brittle, crustose, grayish green, often 

purple-dotted; bristles absent. 


Venezuela, and southern Brazil. 


from mountain summits in the Naguabo and 

Luquillo Mountains. Naguabo and Río Grande. 

Note: Kükenthal (1949) treated R. uniflora 

and R. elongata as separate taxa based on length 

of culms (20-70 cm tall vs. 45-100 cm tall 

respectfully); width of the leaves (0.5-1.5 mm wide 

vs. 1.5-2 mm wide respectfully); number of 

flowers in the spikelets (1-2 vs. 3 respectfully); 

width of the style base in relation to that of the 

achene (½ vs. 2 /3 respectfully); and achene surface 

(indistinctly transversely rugulose vs. distinctly 

transversely rugulose respectfully). We have not 

found these characters to be consistent in any set 

of specimens studied over the entire range of this 

plant, nor have we found any other characters to 

support separation of the two taxa as circumscribed 

by Kükenthal.

362 



Naguabo: Sierra de Naguabo, El Yunque, NW, 

Shafer 3661 (NY, US). Río Grande: Luquillo 

Forest, summit of El Yunque, González-Más & 

Brown 2364 (MAPR); González-Más & Woodbury 

3454 (MAPR); Sierra de Luquillo, Sintenis 1356 

(US). 

31. Rhynchospora velutina (Kunth) Boeck., 

Vidensk. Meddel. Dansk Naturhist. Foren. 

Kjøbenhavn 1869: 149. 1869; Dichromena 

velutina Kunth, Enum. Pl. 2: 282. 1837; 

Psilocarya velutina (Kunth) Nees in Martius 

Fl. Bras. 2(1): 115. 1842. Type: Brazil. Sello 


Psilocarya rufa Nees in Martius, Fl. Bras. 2(1): 

117. 1842; Rhynchospora rufa (Nees) Boeck., 

Vidensk. Meddel. Dansk Naturhist. Foren. 

Kjøbenhavn 1869: 149. 1869. Type: Guyana. 

Schomburgk 667 (lectotype: US!, 

isolectotype: BM, G, K), here designated. 

Rhizomatous perennial, 66-174 cm long; 

rhizome short, stout and thickened, hardened; roots 

coarse, 1-2 mm thick. Culm erect to ascending, 1- 

4.6 mm wide, widest at base, often rooting at 1-2 

nodes just above base, trigonous with blunt angles 

to obtusely trigonous, firm, not hardened, finely 

ribbed, essentially smooth, green, often with pale 

green ribs, glabrous or with a few trichomes 

proximally. Leaves 6-11, ascending, basal and 

cauline, those at very base often with very short 

blades or blades absent; sheaths medium length, 

soft, herbaceous, closely and finely veined with 

pale green veins, light green or pale brown 

proximally, green distally, hirsute to glabrescent; 

ligule absent; blades narrowly linear, 10-90 cm × 

1.5-5 (-6) mm, flattened to plicate, herbaceous, 

finely veined, green and pilose to glabrescent 

abaxially, sometimes pilose proximally, the adaxial 

surface rugose or warty to somewhat smooth or 

roughened, glossy, green to whitish, finely 

cellular-reticulate with white crystals evident in 

some cells, glabrous, margins smooth to scabrous 

distally, long-attenuate to triquetrous apex. 

Inflorescence a terminal and series or 1-3 (-4) 

lateral corymbose partial panicles from the upper 

sheathing bracts, reduced in size proximally; 

terminal panicle 2.5-15 × 11 cm, with 17-190 

spikelets, central axis triquetrous, long-ciliate on 

angles, sheathing bracts and panicle bractlets often 

long-ciliate on margins; spikelets ovoid-ellipsoid, 

5.5-7 (-9) × (1.3-) 1.5-2.5 mm, acute to acuminate 

at apex, acute to obtuse at base, with 12-23 scales; 

fertile scales 9-18, widely ovate or widely ovatedeltate 

to widely ovate-lanceolate, ovatelanceolate, 

or lanceolate (apical scales), 3.2-6.2 × 

1.4-3.5 mm, obtuse to sub-rounded dorsally, thinly 

herbaceous to submembranous, finely cellularstriate, 

uniformly reddish or yellowish brown, 

often with dark brown lineations medially on sides, 

glabrous, margins widely scarious, entire, slightly 

undulate, midcosta very fine, inconspicuous, often 

setose at apex, extending beyond apex as a short 

slightly recurved awn, lateral nerves indistinct, 

acute at apex, the apex of distal spikelet scales 

often recurved. Stamens 3, the anthers 2-3.5 mm 

long, with an elongate papillate appendage at apex, 

truncate at base with minute lobes or papillae; style 

2-branched. Achene biconvex, at maturity often 

falcate in longitudinal section, obovate or obovatedeltate, 

1.5-1.7 × 1.3-1.6 mm, truncate at apex, 

cuneate at base, transversely rugulose-papillate, 

with cellular-reticulate-papillate margins, brown, 

yellowish brown, or dark brown; style base 

triangular to triangular-lanceolate, 0.8-1.2 mm 

long, 1.2-1.4 mm wide at base (nearly as wide as 

achene at base), 2-lobed at base, the lobes 

overlapping apex of achene, brittle, crustose, pale 

brown to light brown; bristles absent. 


America, Greater Antilles, and tropical South 

America. 

Distribution in Puerto Rico: Historically, 

known only from Martín Penã marsh and Río 

Piedras in San Juan. 


San Juan: Martín Peña, Johnston 842 (NY); Río 

Piedras, Johnston & Stevenson 1883 (NY, US). 

32. Rhynchospora wrightiana Boeck., Flora 64: 

78. 1881; Rhynchospora gracillima C. Wright 

in Sauvalle, Anales Acad. Ci. Méd. Habana, 

8: 85. 1871, non Thwaites 1864; 

Rhynchospora distans var. λ. gracillima Kük., 

Repert. Spec. Nov. Regni Veg. 23: 208. 1926. 

Type: Cuba. Wright 3781 (holotype: GH; 

isotypes: NY!, US!). 

Rhynchospora distans var. tenuis Britton, Trans. 

New York Acad. Sci. 11: 90. 1892. Type: 

United States; Georgia. Cutler s.n. (holotype: 

probably at PH).


Rhynchospora pallida sensu C. B. Clarke in 

Urban, Symb. Antill. 2: 126. 1900, pro parte, 

non M.A. Curtis, 1849. 

Rhynchospora brachychaeta sensu Small, Fl. s.e. 

U.S. 196. 1903, non C. Wright, 1871. 

Densely caespitose perennial, (8-) 14-56 (-65) 

cm tall; rhizome short, inconspicuous; roots fine 

to medium, less than 1 mm thick. Culms erect to 

ascending, 0.3-1.7 mm wide, wiry, trigonous to 

obtusely trigonous or subterete, firm but flexible, 

finely ribbed, smooth, green, glabrous. Leaves 

numerous, ascending to loosely spreading, basal 

and cauline, reduced distally on culm; sheaths 

short, herbaceous, finely veined, pale brown to 

light reddish brown, glabrous; ligule absent; blades 

V-shaped, folded, or involute proximally, 

triangular-channeled distally, 3-30 (-35) cm × 0.2- 

1.3 (-2) mm, often curving or wavy at maturity, 

attenuate to triquetrous apex, finely veined 

abaxially, finely cellular-reticulate adaxially, 

essentially smooth, antrorsely scabrous distally on 

margins, bluish green adaxially, otherwise green, 

glabrous. Inflorescence a terminal and series of 1 

or rarely 2 lateral fascicles of spikelets from the 

upper sheathing bracts; fascicles 4-10 × 3-10 mm, 

the smaller lateral fascicles on short, slender erect 

to ascending peduncles; spikelets ovoid, (2.6-) 3- 

3.5 (-4) × 0.8-1.5 mm, acute at apex, cuneate at 

base, with 5-7 scales; fertile scales 2-3, roundedovate 

or rounded-obovate, 2-2.8 × 2-2.8 mm, 

curvate-keeled, deeply boat-shaped, dorsally 

obtuse to rounded or broadly rounded, thinly 

herbaceous, finely cellular-striate, semi-glossy, 

dark brown, margins essentially escarious or 

narrowly so, inrolled around flower at anthesis, 

midcosta fine, pale brown, extended beyond 

slightly recurved obtuse or acute to acuminate apex 

as a mucro or short awn. Stamen 1, the anthers 

0.7-1.3 mm long, apiculate, truncate at base with 

minute papillae or glands; style 2-branched. 

Achene biconvex, elliptic, 1.5-1.8 × 1.1-1.4 mm, 

truncate at apex, short-cuneate at base, faintly 

transversely rugulose or nearly smooth, dark 

brown to brownish black with a distinct or 

indistinct somewhat paler brown or reddish area 

medially; style base triangular, subcompressed, 


flattened with a broad bill-like extension at tip and 

narrowly elliptic swollen rim at base (as seen from 

above), light brown, often with crusty whitish 

surface; bristles 6, subulate, antrorsely barbed, 

reddish, variable in length, rarely exceeding the 

achene. 


States west to Texas and the Greater Antilles. 

Distribution in Puerto Rico: Historically 

known only from mountain summits in the Sierra 

de Luquillo Mountains, collected in 1960 by R. 

O. Woodbury at Laguna Tortuguero. Known from 

the municipalities of Ceiba, Dorado, Río Grande, 



Vega Baja: Tortuguero, 12 March 1960 Woodbury 

s.n. (UPR); Sep 1969, Woodbury s.n. (NY). Río 

Grande: Luquillo Mountains, Wilson 97 (NY, US); 

Sierra de Luquillo, in monte Jimenes, Sintenis 

1381 (NY, US). 


Rhynchospora divergens Chapm. ex M.A. Curtis, 

Amer. J. Sci. Arts, ser. 2, 7: 409. 1849. Cited 

by Urban (1903) based on the collection 

Blauner 247, the type of Rhynchospora 

blauneri Britton which is a synonym of 

Rhynchospora brachychaeta C. Wright. Also 

cited for Laguna Tortuguero by Liogier & 

Martorell (1982: 218; 2000: 243). However, 

no specimens have been found to substantiate 

this record. 

Rhynchospora longiflora C. Presl in Oken, Isis 21: 

269. 1828. Cited for Puerto Rico by Liogier 

& Martorell (1982: 218; 2000: 244), but no 

specimens have been found to confirm this 

record nor has it been recently collected. 

Rhynchospora macra (C. B. Clarke ex Britton) 

Small, Man. S. E. Fl., 180. 1933. Erroneously 

cited by Thomas (Brittonia 44: 27. 1992) as 

occurring in Puerto Rico (Thomas, pers. 

comm. 1999). 

Rhynchospora miliacea (Lam.) A. Gray, Ann. 

Lyceum Nat. Hist. New York 3: 198. 1835. 

Reported from Puerto Rico by Schultes, but 

no specimens were found to substantiate this 

record nor has it been seen or collected since 

(see Britton & P. Wilson, 1923). 

Rhynchospora nervosa (Vahl) Boeck., Vidensk. 

Meddel. Dansk Naturhist. Foren. Kjøbenhavn 

ser. 3, 1: 143. 1868. Erroneously cited by

364 


Thomas (Brittonia 44: 22. 1992) for Puerto 

Rico based on the type he listed for 

Dichromena persooniana Nees. The type 

listing of this name by Nees is a combination 

of localities for synonyms he cited under this 

taxon which included Dichromena ciliata 

Pers. (Florida, Caribbean islands) and 

Schoenus ciliatus G. Mey. (Essequebo, 

Guyana). 

Rhynchospora oligantha A. Gray, Ann. Lyceum 

Nat. Hist. New York 3: 212. 1835. Cited for 

Puerto Rico by Liogier & Martorell (1982: 

219; 2000: 244), but no specimens were found 

to confirm this record nor has it been recently 

collected. Specimens of Rhynchospora 

breviseta (Gale) Channell at the UPR 

herbarium have been misidentified as this 

species. 

Rhynchospora pedersenii Guagl., Darwiniana 39: 

321. 2001. Based on Rhynchospora gigantea 

var. latifolia H. Pfeiff., Repert. Spec. Nov. 

Regni Veg. 17: 236. 1921. Guaglianone 

(2001) cites a specimen from Puerto Rico 

(Otero 537, MO!) as this taxon. However, this 

specimen does not show the characteristics 

described for this taxon. It is glabrous 

throughout, the peduncles of the lateral 

inflorescence panicles are exserted above the 

sheath of their subtending bracts, and is 

otherwise characteristic of typical R. 

corymbosa. Rhynchospora gigantea var. 

latifolia was described as a somewhat hairy 

plant having puberulent culms and tomentose 

or pilose sheaths and proximal leaf blades. 

Perhaps the mold hyphae evident on the Otero 

specimen were mistaken for tomentose hairs 

by the author. 

Rhynchospora perplexa Britton in Small, Fl. s.e. 

U.S. 197. 1328. 1903. Reported for Puerto 

Rico by Kartesz (1998) in “A Synonymized 

Checklist of the Vascular Flora of the United 

States, Puerto Rico, and the Virgin Islands” 

(http://www.csdl.tamu.edu/FLORA/b98/ 

check98.htm). However, no specimen record 

exists (Kartesz, pers. comm., 2001). 

Rhynchospora pinetorum Britton & Small in 

Small, Fl. s.e. U.S. 183, 1503. 1903. 

(Rhynchospora globularis var. pinetorum 

(Britton & Small) Gale). Cited for Puerto Rico 

by Liogier & Martorell (1982: 218; 2000: 

243), but no specimens were found to confirm 

this record nor has it been recently collected. 

Specimens of Rhynchospora recognita (Gale) 

Kral (Rhynchospora globularis var. recognita 

Gale) in the UPR herbarium have been 

misidentified as this species. 

Rhynchospora sola Gale, Rhodora 46: 162. 1944. 

Reported for Puerto Rico by Kartesz (1998) 

in “A Synonymized Checklist of the Vascular 

Flora of the United States, Puerto Rico, and 

the Virgin Islands” (http://www.csdl. 

tamu.edu/FLORA/b98/check98.htm). 

However, no specimen record exists (Kartesz, 

pers. comm., 2001). 

Rhynchospora tenuis Willd. ex Link, Jahrb. 

Gewächsk. 1(3): 76. 1820. Specimens of 

Rhynchospora depressirostris have 

previously been treated as this. 

15. SCHOENOPLECTUS 

Schoenoplectus (Rchb.) Palla, Verh. K. K. Zool.-Bot. Ges. Wien 38 (Sitzungsber.): 49. 1888, nom. conserv. 

Emergent aquatics; perennials or annuals, with elongate, horizontal rhizomes or tufted. Culms 

trigonous or terete, glabrous, air cavities often present. Leaves well-developed or reduced to short 

protrusions from the bladeless sheaths; ligule present or absent. Inflorescences congested and head-like 

or anthelate with elongate rays, terminal, often appearing lateral (pseudolateral); involucral bracts 1-2 (- 

3), the lowest appearing as a continuation of the culm, or (1-) 2-6, flattened and leaf-like; rays with 

tubular prophylls at base; spikelets ovoid, ellipsoid or cylindrical. Flowers bisexual; bristles 0-6, when 

present strap- or needle-like, retrorsely barbed with straight or recurved barbs; stamens 2-3, the anthers 

with prickly or barbed appendage at apex, sagittate at base; style 2- or 3-branched, sub-flattened to strap 

like. Achene plano-convex, trigonous or lenticular, smooth or transversely rugulose, brown or blackish


at maturity. Widely distributed, primarily in North America, Asia, and Africa with approximately 80 

species. 

TYPE: Schoenoplectus lacustris (L.) Palla (≡ Scirpus lacustris L.). 

References: Strong, M.T. 1994. Taxonomy of Scirpus, Trichophorum, and Schoenoplectus 

(Cyperaceae) in Virginia. Bartonia 58: 29-68. Strong, M.T. 1998. Schoenoplectus. Pp. 639-640. In: 

D.M. Kearns et al., Cyperaceae, Flora of the Venezuelan Guayana. Vol. 4, Caesalpiniaceae-Ericaceae, 

eds. P.E. Berry, B.K. Holst, and K. Yatskievych, Missouri Botanical Garden Press, St. Louis, MO. 

Key to the species of Schoenoplectus 

1. Inflorescence simple; spikelets sessile in a single glomerate cluster ...................... 1. S. americanus 

1. Inflorescence compound, with elongate branches; spikelets solitary or in small glomerules of 2-4 at 

branch tips ..................................................................................................................... 2. S. validus 

1. Schoenoplectus americanus (Pers.) Volkart ex 

Schinz & R. Keller, Fl. Schweiz. ed. 2, 1: 75. 

1905; Scirpus americanus Pers., Syn. Pl. 1: 

68. 1805. Type: United States; South Carolina. 

Michaux. s.n. (holotype: P). 

Scirpus olneyi A. Gray ex Engelm. & A. Gray, 

Boston J. Nat. Hist. 5: 238. 1845; 

Schoenoplectus olneyi (A. Gray ex Engelm. 

& A. Gray) Palla, Bot. Jahrb. Syst. 10: 299. 

1888. Type: United States; Rhode Island. 

Olney s.n. (holotype: GH; isotypes: NY, S). 

Emergent, rhizomatous perennial, 30-200 cm 

tall; rhizomes horizontally creeping, elongate, 3.5- 

5 mm thick, reddish. Culms erect, sharply 

trigonous, wing-angled, with deeply concave 

sides, red to red-tinged near base, 3-10 mm wide. 

Leaves 1-2, blade-bearing, or the lower bladeless, 

confined to basal portion of culm; sheaths crossveined, 

the inner band firm with a U- or V-shaped 

notch at orifice; ligule short, ca. 1 mm long at 

adaxial junction of sheath and blade; blades crossveined, 

1-15 × 1-4 mm. Inflorescence 

pseudolateral with a single glomerate cluster of 

(3-) 5-12 spikelets; involucral bract 1, lancetriangular, 

obtuse to subacute, 1-2 (-3.5) cm long, 

appearing as a continuation of the culm, glabrous; 

spikelets ovoid, narrowly ovoid or ellipsoid, obtuse 

to subacute at apex, 5-10 (-20) × 4-7 mm; scales 

broadly ovate to orbicular, 2.5-3.2 × 2-2.5 mm, 

light brown to reddish brown, scarious-margined, 

the green to yellowish midcosta prolonged beyond 

the obtuse apex as a short awn, shorter than to 

equaling the shallowly emarginate apex. Stamens 

3, the anthers narrowly oblong, 2-2.5 mm long, 

with a very short, blunt, minutely bearded 

appendage at tip; style straplike, 2- or sometimes 

3-branched, the branches glabrous, minutely scaly. 

Achene plano-convex, suborbicular to broadly 

ovate, 2-2.5 × 1.6-2.2 mm, smooth, brownish to 

grayish brown; bristles 1-6, reddish, with 

downcurved-retrorse barbs, shorter than to 

equaling the achene. 




Islands: In brackish or saline waters of marshes 

or marshy grounds and roadside ditches. Arecibo, 

Barceloneta, Guánica, Guayama, Ponce, Río 

Grande, and San Juan; St. Croix, St. Thomas, and 

Tortola. 


Arecibo: Caño Tiburones, near Arecibo, Gleason 

& Cook F-7 (NY). Barceloneta: Bo. Palmas Altas, 

Caño Tiburones, Axelrod et al. 10521 (UPRRP, 

US). Guánica: Sargent 199 (US). Guayama: Bo. 

Mosquito, Rd 3, km 154, González-Más 1301 (NY, 

US). Río Grande: Bo. Zarzal, vicinity of Punta 

Picúa, Proctor 43543 (US). San Juan: Park of 

Santurce, Hioram 905 (US). 

2. Schoenoplectus validus (Vahl) Á. Löve & D. 

Löve, Bull. Torrey Bot. Club 81: 33, 1954; 

Scirpus validus Vahl, Enum. Pl. 2: 268. 1805; 

Scirpus lacustris subsp. validus (Vahl) T. 

Koyama, Canad. J. Bot. 40:927, 1962. Type: 

West Indies. Banks s.n. (holotype: C-Vahl). 

Scirpus lacustris sensu C.B. Clarke in Urban, 

Symb. Antill. 2: 93. 1900; and Urb., Symb. 

Antill. 4: 120. 1903, non Linnaeus, 1753. 

Fig. 55. F-K

366 


Emergent, rhizomatous perennial, 50-300 cm 

tall; rhizomes stout, scaly, horizontally creeping, 

7-15 mm thick, reddish; roots pale brown to brown, 

to 1 mm thick. Culms erect, terete, easily 

compressed, 3-20 mm wide at base, forming in a 

row along the rhizome. Leaves 3-5 bladeless 

sheaths, short mucronate or the uppermost with 

short blades, confined to basal portion of culm; 

sheaths glabrous, with scarious margins, open at 

acute to rounded summit, the ventral portion 

gradually narrowing towards base to V-shaped 

orifice; ligule short, firm, ca. 1.5 mm long, borne 

at adaxial base of blade; blades 1-2 on upper 

sheaths, 2-20 cm long, dorsally flattened, 

thickened towards summit with cartilaginous tip. 

Inflorescence pseudolateral, anthelate, compound 

to partially decompound, pendulous, the numerous 

spikelets solitary or in small glomerules of 2-4 at 

ray tips; involucral bracts 2-3, the lowest one erect, 

looking like a continuation of the culm, the others 

membranaceous, scale-like; rays flat, to somewhat 

obtusely angled, glabrous, scabrous on margins, 

pendulous; spikelets ovoid to ovoid-ellipsoid, 

acute, 5-7 (-9) × 3-4 mm; scales ovate to elliptic, 

2.5-3.2 × 1.8-2.2 mm, reddish brown, erose-ciliate, 

with scarious margins, the green midcosta 

prolonged as a short, scabrous awn at the 

emarginate apex. Stamens 3, marescent; anthers 

narrowly oblong, 1-2 mm long, tipped by a 

triangular-ovate appendage; style straplike, 2- or 

3-branched, the branches glabrous, with minute 

scales. Achene plano-convex, broadly ovoid, 1.7- 

16. SCLERIA 

Scleria P. J. Bergius, Kongl. Vetensk. Acad. Handl. 26: 142. 1765. 

2.3 × 1.3-1.5 mm, smooth, grayish brown, reddish 

or black; bristles 2-6, retrorsely barbed, reddish, 

shorter than to equaling or slightly exceeding the 

achene. 




Islands: In wet areas, often in standing water of 

marshes, open swamps or swampy grounds, 

pastures, roadside ditches, and disturbed areas. 

Arecibo, Guánica, Guayama, Mayagüez, Ponce, 

Salinas, San Germán, Toa Baja, Utuado, and Vega 

Baja; St. Croix, St. Thomas, and Tortola. 

Common name: Puerto Rico: Junco. 

Note: This species is sometimes included 

within a broad circumscription of European 

Schoenoplectus tabernaemontani (C. C. Gmel.) 

Palla. 


Arecibo: Bo. Río Abajo, Acevedo-Rdgz. & Cedeño 

10190 (UPRRP, US). Guánica: Sintenis 3846 (NY, 

US); Sargent 200 (US); N.L. Britton & Shafer 1859 

(NY, US); near Guánica, Heller 6290 (NY, US). 

Mayagüez: Vicinity of Mayagüez, Chase 6320 

(US). Ponce: Stevens 2404 (NY). Salinas: Rd. 3, 

km 156, González-Más 1294 (NY, US). San 

Germán: Road 2, km 200.9, González-Más 1202 

(NY). Toa Baja: Bo. Sabana Seca, Axelrod et al. 

9747 (UPRRP). Utuado: Bosque de Río Abajo, 

Saltillo, Acevedo-Rdgz. 339 (SJ). Vega Baja: Bo. 

Cabo Caribe, E side of Rt 686, near coast, Axelrod 

& Thomas 10208 (UPRRP, US). 

Perennials or sometimes annuals; rhizomes when present horizontal or short and nodose, hardened 

and knotty, sometimes tuberous. Culms erect, elongating and sprawling, or climbing, trigonous or 

triquetrous, harshly scabrous to smooth, glabrous or pubescent, green. Leaves well-developed at middle 

and upper nodes, the basal ones essentially bladeless; sheaths 3-angled, closed at summit, distinctly 

veined, the apex of the inner band with a rounded, obtuse, or triangular contraligule with distinct, straight 

or anastomosing veins, the margin thickened or cartilaginous, sometimes with a short to elongate scarious 

appendage; ligule absent or sometimes present; blades linear-elongate or sometimes lanceolate, flattenedplicate 

to plicate or somewhat inrolled along margins, 3-costate, herbaceous, weakly to harshly scabrous 

on margins and costae, glabrous or sometimes pubescent. Inflorescence paniculate or spike-like, terminal, 

or terminal and a series of axillary partial panicles from the upper leaf-like bracts; panicle branches 3angled, 

sometimes narrowly winged, scabrous or smooth, pubescent or glabrous; panicle bractlets linearlanceolate 

or setaceous, often ciliate or scabrous on margins; spikelets unisexual, or bisexual and 

androgynous, sessile or on pedicels to 1 cm long; staminate spikelets lanceolate or narrowly oblongovate, 

cylindrical or subcompressed, many-flowered; staminate scales numerous, the basal 2-3 often 2ranked 

and sterile, like the sterile pistillate spikelet scales, the upper fertile ones spirally imbricate,


Fig. 55. A-E. Rhynchospora depressirostris. A. Habit. B. Terminal inflorescence unit. C. Spikelet scale. D. Flower. E. Achene. 

F-K. Schoenoplectus validus. F. Habit. G. Inflorescence. H. Spikelet. I. Spikelet scale. J. Flower. K. Achene. (A-D, from Stevenson 

889; E, from Johnston 884; F-G and J, from Proctor 42230; H, I, and K, from Proctor 45573).

368 


narrowly ovate to lanceolate, membranous; pistillate spikelets ovoid to ovoid-lanceoloid or ellipsoid, 

subcompressed, often becoming obovoid after expanding to the width of the mature achene, cylindrical 

to subcompressed, with a single terminal flower which is usually subtended by a much reduced, lanceolate 

fertile scale, hidden by the uppermost well-developed sterile scale; pistillate spikelet scales 2-ranked, 

the uppermost two broadly boat-shaped or cupuliform and spreading widely with the developing achene, 

carina 1- to 3-nerved, lateral nerves indistinct. Flowers unisexual; hypogynium (when present) borne at 

base of achene, sessile or stipitate, smooth or crustaceous, entire or 3-lobed, sometimes 3- to 9-tuberculate 

near base, the lobes entire or dissected, sometimes reflexed; cupula supporting the hypogynium and 

ovary shallow and dish-like, 3-lobed or obtusely trigonous with smooth margins, or deep and cup-like, 

enveloping the hypogynium and often ciliate on margin; stamens 1-3, the anthers often prickly-appendaged 

at apex; styles capillary, 3-branched, the unbranched portion glabrous. Achene body globose or ovoid to 

ellipsoid, rounded or obtusely trigonous, rarely trigonous or triquetrous, sometimes subconic, with straight 

to recurved apex, apiculate, bony or crustaceous, white or sometimes variegated with purple, glabrous 

or pilose, the surface smooth, rugose, reticulate, trabeculate, papillate, verrucose, or warty. A genus of 

primarily warm-temperate and tropical regions worldwide, with approximately 200-225 species. 

LECTOTYPE: Scleria flagellum-nigrorum P. J. Bergius, nom. conserv. 

References: Core, E. L. 1936. The American species of Scleria. Brittonia 2: 1-105. Camelbeke, K. 

and P. Goetghebeur. 1998. Scleria (Cyperaceae). Pp. 641-659. In: D.M. Kearns et al., Cyperaceae, Flora 

of the Venezuelan Guayana. Vol. 4, Caesalpiniaceae-Ericaceae, eds. P.E. Berry, B.K. Holst, and K. 

Yatskievych, Missouri Botanical Garden Press, St. Louis, MO. Camelbeke, K., K. Spruyt and P. 

Goetghebeur. 2003. The genus Scleria (Cyperaceae) in Bolivia. Revista Soc. Boliv. Bot. 4(1): 139-170. 

Key to the species of Scleria 

1. Slender perennials or annuals, erect or ascending, never vine-like; hypogynium wanting or 

inconspicuous, when present, trigonous, not bearing a pearl-colored pebbly crust nor supporting 

tubercles; achene body often porose proximally............................................................................. 2 

2. Inflorescence a solitary fascicle of spikelets at the summit of the culm; achene body longitudinally 

ridged with a pair of iridescent pits on each side proximally..... 6. S. georgiana 

2. Inflorescence of 2 or more fascicles or panicles; achene bodies smooth, verrucose, or tuberculate, 

lacking iridescent pits proximally....................................................................... 3 

3. Inflorescence composed of a terminal and series of 2-3 remote, axillary, narrow partial 

panicles; spikelet scales finely appressed-pubescent .............................. 9. S. lithosperma 

3. Inflorescence interruptedly fasciculate-spicate; spikelet scales glabrous or coarsely 

setose............................................................................................................................... 4 

4. Spikelet scales glabrous ....................................................................16. S. verticillata 

4. Spikelet scales coarsely setose.................................................................................. 5 

5. Perennial with horizontally creeping rhizome; fascicles of spikelets nodding....... 

............................................................................................................ 4. S. distans 

5. Caespitose annual; fascicles or single spikelets ascending................ 8. S. hirtella 

1. Medium-sized to coarse perennials, erect or sometimes vine-like; hypogynium present, generally 

conspicuous, 3-lobed, trigonous, or discoid, sometimes bearing a pearl-colored pebbly crust, or 

supporting tubercles between it and achene body, or hidden by the well-developed cupula which is 

entire, ciliate, or lacerate on its margin; achene body rarely porose............................................... 6 

6. Hypogynium hidden by the well-developed cupula which is ciliate on its margin (sparsely and 

inconspicuously so in Scleria mucronata), not supporting tubercles. ................................. 7 

7. Edges of leaf blade abruptly narrowed at base, not decurrent or only very narrowly so along 

the lateral margins of the sheath; cupula sparsely and inconspicuously ciliate on margins; 

style base smooth or essentially so...................................... 12. S. mucronata 

7. Edges of leaf blade decurrent and forming broad wings along the lateral margins of the 

sheath; cupula densely ciliate on margins; style base finely antrorsely scabrid (at least 

proximally). ............................................................................................................. 8


8. Culms (5-) 6-12 mm wide; leaf blades (10-) 13-27 mm wide; achene body 2.5-3 mm 

wide; style base 0.8-1.2 mm long .................................................... 5. S. eggersiana 

8. Culms 3-6 mm wide; leaf blades 4-13 (-15) mm wide; achene body 1.2-1.9 mm wide; 

style base 0.3-0.5 mm long ................................................. 11. S. microcarpa 

6. Hypogynium not hidden by the cupula, smooth on margins, with or without tubercles between 

it and achene body.................................................................................................... 9 

9. Achene body reticulate or verrucose............................................................................... 10 

10. Hypogynium trigonous, with a pearl-colored pebbly crust on sides between the 

decurrent costae of the achene body; achene body shallowly reticulate to nearly 

smooth............................................................................................... 7. S. havanensis 

10. Hypogynium 3-lobed, without a pearl-colored pebbly crust on its sides; achene body 

distinctly reticulate or verrucose................................................................... 11 

11. Hypogynium deeply 3-lobed, the tips of the lobes appressed to the base of the 

achene body, lacking tubercles; achene body reticulate...... 12. S. muehlenbergii 

11. Hypogynium shallowly 3-lobed, the lobes short, not reaching achene body, 

supporting three entire or 2-lobed tubercles; achene body verrucose............. 12 

12. Plant glabrous except for the strigillose sheaths; achene body 1-2 mm 

diam.......................................................................................... 1. S. brittonii 

12. Plant pubescent, densely ciliate on angles of sheaths and culms, margins and 

abaxial midveins of leaf blades and inflorescence bracts, and carina of spikelet 

scales; achene body 2-3 mm diam ............................... 3. S. ciliata 

9. Achene body smooth....................................................................................................... 13 

13. Hypogynium trigonous, with a pearl-colored pebbly crust on sides between the 

decurrent costae of the achene body; achene body shallowly reticulate to nearly smooth 

............................................................................................. 7. S. havanensis 

13. Hypogynium 3-lobed or obscurely so; achene body smooth................................. 14 

14. Plants erect, not scrambling or climbing .................................. 10. S. melaleuca 

14. Plants vine-like or scandent, scrambling or climbing, often forming impenetrable 

masses........................................................................................ 15 

15. Contraligule typically with a scarious appendage at its apex; hypogynium 

obscurely 3-lobed, discoid, irregularly wrinkled ..................... 15. S. secans 

15. Contraligule lacking an appendage at apex with a cartilaginous, often ciliate 

or pubescent margin; hypogynium distinctly 3-lobed, the lobes 

reflexed..................................................................................................... 16 

16. Inflorescence panicles open; leaf blade margins with coarse barbs, 0.2- 

0.5 mm long; spikelet scales dark brown or purplish brown; achene body 

ovoid-globose to ovoid, 2.6-3.3 mm long; hypogynium lobes strongly 

revolute, obtuse, typically appressed to base of achene body 

.............................................................................. 2. S. canescens 

16. Inflorescence panicles contracted; leaf blade margins with fine barbs, 

less than 0.2 mm long; spikelet scales greenish brown; achene body 

globose, 2.3-2.6 mm long; hypogynium lobes weakly revolute, rounded, 

often spreading away from base of achene body................... 14. S. scindens 

1. Scleria brittonii Core in Small, Man. S. E. Fl. 

190. 1933, as a new name for Scleria glabra 

(Chapm.) Britton in Small, Fl. s.e. U.S. 200. 

1903, non Boeckeler, 1888; Scleria pauciflora 

var. glabra Chapm., Fl. South. U.S. 532. 1860; 

Scleria ciliata var. glabra (Chapm.) Fairey, 

Castanea 32: 48. 1967. Type: United States; 

Florida. Chapman s.n. (lectotype: NY, 

isolectotypes: NY!, US!), designated by 

Fairey, Castanea 32: 51. 1967. 

Slender, rhizomatous perennial, 20-70 (-90) 

cm tall; rhizome horizontal, nodose, 1.5-5 mm 

thick, often densely branched and interwoven with

370 


those of previous seasons. Culms erect, 

triquetrous, glabrous, smooth on angles, 0.7-1.7 

(-2) mm wide. Leaves 3-5 below inflorescence; 

sheaths unwinged, retrorsely scabrid on angles, 

densely strigillose, brownish, stained with purple 

distally; contraligule convex to rounded, strigillose 

with a scarious margin; ligule absent; blades 

plicate, the lowermost short, often bifid, 3-25 

(-40) cm × 1-2.5 mm, glabrous both abaxially and 

adaxially, margins, abaxial midvein, and adaxial 

lateral veins remotely antrorsely scabrous (at least 

distally), long-attenuate to triquetrous apex. 

Inflorescence composed of a terminal and series 

of 1-2 remote, turbinate, long-peduncled partial 

panicles from the upper leaf-like bracts, the 

terminal one largest 0.6-1.7 cm diam.; bractlets 

lanceolate-subulate, ciliate-scabrous; spikelets 5- 

7 mm long, the lowermost scales spreading to 4 

mm with mature achene; staminate scales yellowbrown; 

sterile pistillate spikelet scales 3, ovatelanceolate 

to lanceolate, 3-5 × 1.5-2.6 mm, 

glabrous, stramineous to light brown, reddish 

lineolate, glabrous on margins, carina 3-nerved, 

prolonged beyond the acuminate apex as a 

scabrous mucro. Staminate flower with 3 stamens, 

the anthers 2-3 mm long, with a linear-subulate 

apiculum beset with antrorse crystalline prickles. 

Achene body globose, 1.5-2 × 1.7-2 mm, 

transversely verrucose-reticulate, or sometimes 

nearly smooth, apiculate, 2-porose on each side, 

estipitate, white or sometimes with grayish 

patches; style base deciduous; hypogynium 

narrow, obtusely trigonous, 0.5-0.7 × 0.8-1 mm, 

supporting 3 entire or 2-lobed tubercles. 


States, Cuba, and Puerto Rico. 

Distribution in Puerto Rico: Sand savannas. 

Vega Baja, known to occur only in the vicinity of 

Laguna Tortuguero. 

Note: The glabrous culm, leaf blades, and 

inflorescence; densely strigillose sheaths; and 

consistently smaller achenes separate this species 

from the closely related Scleria ciliata. 


Manatí: Laguna Tortuguero, May 1960, Woodbury 

s.n. (US); Tortuguero area, Liogier 33563 (UPR). 

2. Scleria canescens Boeck., Beitr. Cyper. 1: 37. 

1888. Type: Puerto Rico. Guayama. Kuntze 

s.n. (holotype B, destroyed; isotype: NY!). 

Fig. 56. A, B 

Robust perennial, 0.5-3 m tall, growing in 

large colonies, often forming dense tangles; 

rhizome stout, nodose. Culms ascending, vine-like 

and clambering, solitary from nodes of the 

rhizome, sharply trigonous, 3-10 mm wide, 

antrorsely and retrorsely scabrous or scabrid on 

angles, sparsely strigose and scabrid on sides 

towards the apex, glabrous at base. Leaves 

numerous, the uppermost overtopping the 

inflorescence; sheaths stiff, loose, glabrescent, 

with a deltate or sub-rounded contraligule, with 

straight veining, the margin cartilaginous, 

canescent to glabrescent; ligule absent; blades 

plicate or subflattened, 25-60 cm × 4-10 mm, stiff, 

somewhat coriaceous, sharply retrorsely scabrous 

on margins and abaxial midvein, the apex 

acuminate to blunt, flattened tip. Inflorescence 

composed of a terminal, open, pyramidal panicle, 

(5-) 6-14 × (3.5-) 4-11 (-13) cm and 1 to 3 

additional smaller lateral panicles in the upper leaflike 

bracts; bractlets setaceous, ciliate on margins 

proximally; panicle branches minutely pubescent 

to glabrescent; staminate spikelets about the same 

length as the pistillate; pistillate spikelets ovateelliptic, 

4-6 × 2-3 mm; sterile pistillate spikelet 

scales 4-5, ovate-orbicular, the uppermost one 3.5- 

5 × 2-3.7 mm, the lower successively smaller, 

canescent, brown or reddish brown with purpleblack 

veins and margins, finely ciliate on margins, 

carina 1-nerved, shortly prolonged beyond the 

acute apex. Staminate flowers with 3 stamens, the 

anthers 2-3 mm long, with a subulate-conic 

apiculum. Achene body ovoid-globose to ovoid, 

2.6-3.3 × 2.4-2.8 mm, minutely apiculate, shiny 

white to purple-tinged; hypogynium 1-1.8 × 2-2.7 

mm, 3-lobed with broad, strongly revolute, obtuse, 

entire, ascending lobes which are appressed to base 

of achene body. 

General distribution: Apparently endemic to 

Puerto Rico, although reported by León (1946) 

from Cuba. 

Distribution in Puerto Rico: Wet montane, 

secondary, or palm forests, in openings, along 

trails, and forest edges. Adjuntas, Caguas, Cayey, 

Ciales, Guayama, Jayuya, Maricao, Naguabo, 

Orocovis, Patillas, and Río Grande. 

Common name: Puerto Rico: Cortadora. 


Adjuntas: Monte Guilarte State Forest, Proctor & 

Alemany 45833 (SJ). Barranquitas: Monte 

Torrecilla, N.L. Britton et al. 5544 (NY). Caguas:


Sierra de Cayey, Carite Forest Reserve, Proctor 

47890 (SJ, US). Cayey: Carite Forest Reserve, 

Axelrod & Axelrod 4375 (NY, UPRRP). Ciales: 

Cordillera Central, Toro Negro State Forest, 

Proctor & Haneke 44588 (SJ). Ceiba: Caribbean 

National Forest, Fs Rd 27, Axelrod et al. 481 

(UPRRP). Jayuya: Cordillera Central, Bo. Coabey, 

Proctor & Haneke 42037 (SJ). Las Piedras: 

Luquillo Forest, peak of El Toro, Hill 46 (NY). 

Luquillo: Sierra de Luquillo, along Pico del Este 

road, Taylor & Walker 11854 (NY, UPRRP). 

Maricao: Río Maricao, N.L. Britton et al. 2432 

(NY, US); Bo. Maricao Afuera, Río Maricao 

margins, Cedeño & de la Cruz 639 (MAPR). 

Miramontes, N.L. Britton & E.G. Britton 9537 

(NY). Naguabo: Sierra de Luquillo, summit of La 

Mina, Shafer 3317 (NY, US); Sierra de Luquillo, 

Proctor 47981 (US). Orocovis: Doña Juana, Toro 

Negro, Liogier & Martorell 35107 (NY). Patillas: 

Carite Forest Reserve, Axelrod et al. 1951 

(UPRRP). Ponce: Toro Negro Forest Reserve, 

Axelrod & Chávez 4285 (NY, UPRRP). Río 

Grande: El Yunque, Fosberg 44190 (US). 

3. Scleria ciliata Michx., Fl. Bor.-Amer. 2: 167. 

1803. Lectotype: United States; “Carolina”. 

M. Michaux (P), designated by Fairey, 

Castanea 32: 50. 1967. 


rhizome horizontal, nodose, 3-6 mm thick, often 

densely branched and interwoven with those of 

previous seasons. Culms ascending to erect, 

triquetrous to trigonous, sparsely pubescent on 

sides, densely ciliate on angles, 0.8-3 (-4) mm 

wide. Leaves 3-7; sheaths unwinged, densely 

ciliate on angles, short-pilose on sides, light brown 

to reddish brown, the lowermost often forming 

short, 2-fid blades; ligule absent or sometimes a 

faint line; contraligule obtuse to rounded, 

pubescent, the margin often densely ciliate; blades 

plicate to subflattened, 10-53 cm × 1-4 mm, 

pubescent to glabrous both abaxially and adaxially, 

margins and abaxial midvein densely ciliate, acute 

to acuminate to a triquetrous or 2-fid tip. 

Inflorescence composed of a single terminal 

panicle or often with a remote, smaller second one 

below from the upper leaf-like bracts, the terminal 

one 1-2.5 × 0.5-1 cm, subtended by an erect leaflike 

bract appearing as a continuation of the culm; 

bractlets linear-lanceolate, densely ciliate; 

spikelets unisexual (female) or androgynous, the 

staminate portion (spikelet) borne at the base of 

the cupula; staminate spikelets 4-7 × 0.8-1.6 mm; 

staminate scales brown; pistillate spikelets ovoid 

to ovoid-ellipsoid, slightly compressed, 3.5-6 × 

1-2 mm; sterile pistillate spikelet scales 3-4, ovate 

to lanceolate, 3-5 × 1.5-2.5 mm, pubescent to 

sparsely so, finely reddish lineolate, light brown, 

tinged with dark brown or purple on distal sides 

and margins, carina 1- to 3-nerved, prolonged 

beyond the acute to acuminate apex as an excurved 

awn. Staminate flower with 3 stamens, the anthers 

2.5-3 mm long, with a prickly apiculum. Achene 

body globose, 2-3 mm diam., transversely 

verrucose-reticulate, often papillate distally, or 

sometimes nearly smooth, apiculate, eporose, 

estipitate, white or sometimes with grayish patches 

or green-mottled; style base deciduous; 

hypogynium narrow, obtusely trigonous, 0.5-0.8 

× 1-1.5 mm, light yellow or stramineous on sides, 

supporting 3 entire or 2-lobed tubercles at base. 


Central America, and the West Indies. 

Distribution in Puerto Rico: Dry grassy slopes 

and damp ravines, on lateritic soil, 350-400 m. 

Maricao (Maricao Forest Reserve) and Mayagüez 

(Cerro Las Mesas and vicinity). 


Maricao: Bo. Maricao Afuera, Maricao Forest 

Reserve, Axelrod & Stenzel 11107 (UPRRP). 

Mayagüez: Las Mesas near Mayagüez, 29 Dec 

1914, Holm s.n. (US); 23 Nov 1914, Holm 45 (US). 

4. Scleria distans Poir. in Lamarck, Encycl. 7: 4. 

1806. Type: Puerto Rico. Ledrú 110 

(lectotype: P; isolectotype: P), designated by 

J. Raynal, Adansonia, ser. 2. 16: 216. 1976. 

Scleria nutans Willd. ex Kunth, Enum. Pl. 2: 351. 

1837. Type: Venezuela. Humboldt s.n. 

(holotype: B-Willd. 17336; isotypes: HAL, P). 

Slender perennial, 15-60 cm tall; rhizome 

scaly, horizontally creeping, 2-4 mm thick. Culm 

erect, triquetrous, 0.8-2.7 (-3) mm wide, glabrous, 

angles smooth, wire-like. Leaves 2-8; sheaths 

subinflated at apex, unwinged, hirsute to 

glabrescent, angles smooth or hirsute; ligule 

absent; contraligule obtuse or slightly concave, 

with hirsute margin, the inner band often hirsute 

in a longitudinal line medially; blades flattened to 

subplicate, 1-20 cm × 1.7-4 (-5) mm, abaxial and

372 


adaxial surface glabrous, margins, midvein, and 

lateral veins often hirsute, or sparsely so, 

subabruptly narrowed to an acute or acuminate, 

subflattened apex. Inflorescence terminal, 

interruptedly glomerate-spicate with no lateral 

branching, 8-22 cm × 7-10 mm, composed of 4- 

10, sessile, fascicles of spikelets; fascicles 4-8 mm 

diam., with 3-7 (-9) spikelets, reflexed at maturity; 

bractlets ciliate at base, margins with dark hairs; 

spikelets bisexual (androgynous), ovoid-ellipsoid, 

3.5-5 × 0.8-1.3 mm; staminate scales lanceolate, 

membranous; pistillate scale ovate to ovatelanceolate, 

2-2.5 (-3) × 1-1.5 mm (excluding awn), 

sparsely setose on sides, pale green to dark 

purplish brown, carina 1-nerved, setose, prolonged 

beyond the acute apex as a setose awn. Staminate 

flowers with 2 stamens, the anthers 1.7-2.2 mm 

long, with a lanceolate, bristly appendage at apex. 

Achene body subglobose to broadly obtusely 

trigonous in cross section, 1.4-1.8 × 1-1.3 mm, 

broadly obtuse to rounded at apex, apiculate, with 

a brownish or blackish apiculum, porose, smooth, 

white or grayish, with a stipe-like, trigonous base; 

style base deciduous; hypogynium inconspicuous, 

3-lobed, with setaceous awl-shaped lobes, these 

often impressed into margins of base. 


States, Panama, West Indies, South America, and 

tropical and southern Africa, including 

Madagascar. 

Distribution in Puerto Rico: In moist or wet 

sandy soils of savannas, pastures, meadows, grassy 

slopes, and roadside banks. Aibonito, Cayey, 

Dorado, Guayama, Lares, Mayagüez, Moca, 

Naguabo, Río Grande, San Juan, Vega Alta, and 

Vega Baja. 


Aibonito to Cayey, Chase s.n. (US). Dorado: W 

or Dorado, Woodbury s.n. (US). Lares: Sargent 

3253 (US). Moca: Sargent 366 (US). Naguabo: 

Caribbean National Forest, Axelrod & Grose 11224 

(UPRRP). Río Grande: El Yunque, Fosberg 44197 

(US); Sargent 554 (US). San Juan: Santurce, Heller 

& Heller 986 (US); Río Piedras, Stevenson 6488 

(US). Vega Alta: Bo. Sabana, Proctor & 

Concepción 41833 (US). 

5. Scleria eggersiana Boeck., Beitr. Cyper. 2: 41. 

1890. Type: Cuba. Eggers s.n. (holotype: B, 

destroyed). 

Scleria microcarpa var. latifolia Boeck., Linnaea 

38: 517. 1874. Type: Guadeloupe. Collector 

unknown (holotype: B, destroyed). 

Scleria microcarpa sensu Grisebach, Fl. Brit. W. 

I. 578. 1864, pro parte, non Nees von 

Esenbeck, 1834. 

Scleria mitis sensu Grisebach, Fl. Brit. W. I. 578. 

1864, non P. J. Bergius, 1765. 

Scleria grisebachii C. B. Clarke in Urban, Symb. 

Antill. 2: 150. 1900. Lectotype: Martinique. 

Duss 445 (NY!), here designated. 

Coarse, rhizomatous perennial, 1-3 m tall; 

rhizome short, horizontal, scaly, knotty at culm 

bases, 5-10 mm thick. Culm erect, approximate 

or remotely spaced on rhizome, obtusely trigonous 

to trigonous proximally, trigonous to triquetrous 

distally, stiff and hardened, (5-) 6-12 mm wide, 

glabrescent, smooth, retrorsely scabrid on angles. 

Leaves 5-8; sheaths broadly winged, glabrous, 

angles retrorsely scabrous or irregularly scabrid; 

ligule absent; contraligule triangular to lanceolate, 

the margin cartilaginous, crisped, ciliate-scabrous; 

blades subflattened to plicate, 20-70 × (1-)1.3-2.7 

cm, glabrescent, margins and abaxial midvein 

antrorsely scabrous, the distal margins often 

forming small, obliquely positioned ears, above 

these, the blade is acuminate to a blunt, 

subflattened apex. Inflorescence composed of a 

terminal and series of 2-5 (-8), narrowly rhombic, 

axillary partial panicles from the upper leaf-like 

bracts, the terminal one largest, 7-15 × 1.5-4 cm; 

bractlets linear, finely ciliate on margins; spikelets 

unisexual; staminate spikelets 3.5-4.2 × 1.2-1.6 

mm; staminate scales golden brown to light 

reddish brown; pistillate spikelets ovoid, 3-4 × 

2.5-4 mm; sterile pistillate spikelet scales 4, ovateorbicular, 

the uppermost 1.6-2.8 mm diam., 

yellowish brown, reddish brown-lineolate, carina 

1-nerved, shortly prolonged beyond the acute to 

broadly acute apex. Staminate flower with 3 

stamens, the anthers 1.4-2 mm long, with a 

triangular-subulate prickly apiculum. Achene body 

ovoid to subglobose or globose, 2-2.5 × 2.5-3 mm, 

rounded to subtruncate at apex, not apiculate, 

smooth, glabrous, white to grayish white, 

estipitate, eporose; style base deciduous or 

persistent, conic-cylindrical, 0.8-1.2 × 0.4-0.6 mm, 

finely antrorsely scabrid; hypogynium 3-lobed, 

adhering to and hidden by the spongy-thickened, 

tan to yellowish, darker reticulate-veined,


rugulose, shallowly 3-lobed cupula, 1.3-2.3 × 2.8- 

3.3 mm, which is densely ciliate on its rim, the 

hairs yellow-brown. 


Central America, West Indies, Trinidad, Tobago, 

and northern South America. 


areas of marshy habitats, forest margins, swales, 

drainage ditches, canals, pastures, and roadside 

ditches. Bayamón, Cataño, Loíza, Luquillo, 

Maunabo, Naguabo, and Río Grande. 

Common names: Puerto Rico: Cortadera, 

Cortadora, Cortadora de altura, Lambedora. 


Bayamón: Sintenis 1215 (US). Río Grande: Sierra 

de Luquillo, Sintenis 1443, 1714 (US); Bo. 

Herreras, Rd. 187, Proctor 45162 (US); Proctor 

& Rivera 48358 (US). 

6. Scleria georgiana Core, Brittonia 1: 243. 1934; 

Scleria gracilis Elliott, Sketch Bot. S. 

Carolina 2: 557. 1824, nom. illeg., non 

Richard, 1792. Type: United States; Georgia. 

Baldwin s.n. (holotype: CHARL; isotype: 

NY). 

Slender rhizomatous perennial, (12-) 18-65 

(-80) cm tall; rhizome horizontal, nodose, 2.5-4 

mm thick, often densely branched and interwoven 

with those of previous years. Culms erect, wiry, 

trigonous, glabrous, smooth on angles or 

sometimes sparsely antrorsely scabrous at apex, 

0.7-1.5 mm wide. Leaves 1-3; sheaths elongate, 

unwinged, hispidulous to glabrous, purplish; ligule 

absent; contraligule absent or essentially so, the 

orifice convex to truncate, or subdeltate on upper 

sheaths, hispidulous; blades triangular-channeled 

proximally, trigonous distally, lowermost on culm 

with short plicate or 2-parted blades, 2-30 cm × 

0.3-1 mm, to 1.5 mm wide at base, glabrous, edges 

of channel antrorsely scabrous, attenuate to 

triquetrous apex. Inflorescence composed of a 

single terminal fascicle of (-1) 2-8 (-14) spikelets 

at the summit of the culm, or sometimes a second 

smaller, subcontiguous fascicle below, 5-15 mm 

in diam.; involucral bracts 2-3, leaf like but 

reduced, the lowermost one elongate, 1-16 cm 

long, overtopping the inflorescence and appearing 

as a continuation of the culm; spikelets unisexual 

(female) or androgynous, the staminate portion 

(spikelet) borne at the base of the cupula; staminate 

spikelets 5-7 × 1.5-2.5 mm; staminate scales 

reddish brown; pistillate spikelets ovoid to ovoidlanceoloid, 

slightly compressed, 3.5-7 × 1.5-2.5 

mm; sterile pistillate spikelet scales ovate to ovatelanceolate, 

3.5-6 × 2-4 mm, light brown, finely 

reddish lineolate, stained with dark brown or 

reddish brown on distal sides and margins, carina 

1-nerved, prolonged beyond the acute to acuminate 

apex as a short straight to slightly excurved awn. 

Staminate flowers with 3 stamens, the anthers 3-4 

mm long, with a bristly apiculum. Achene ovoid 

or ovoid-ellipsoid, 2.5-2.7 × 1.7-2 mm, 

longitudinally ridged or indistinctly so, broadly 

obtuse to rounded at apex, apiculate, eporose, 

estipitate, milky white, sometimes stained (often 

uniformly so) with purple, the base trigonous, 

concave and bearing 2 iridescent pits on each side; 

style base deciduous; hypogynium absent. 


States, Belize, Greater Antilles, and Brazil. 

Distribution in Puerto Rico: Wet sands in 

savanna-like swales or seasonal pools. Vega Baja, 

known to occur only in the vicinity of Laguna 

Tortuguero. 


Manatí: Laguna Tortuguero, 03 Dec 1960, 

Woodbury s.n. (US); May 1960, Woodbury s.n. 

(US); 2 Feb 1970, Woodbury s.n. (US). Vega Baja: 

Bo. Algarrobo, just S of Laguna Tortuguero, 

Proctor et al. 45031 (US). 

7. Scleria havanensis Britton, Bull. Torrey Bot. 

Club. 42: 492. 1915. Type: Cuba. León 4731 



cm tall; rhizomes relatively stout, horizontal, 

nodose, 4-6 mm thick. Culms crowded along 

rhizome, erect, slender, trigonous proximally, 

triquetrous distally, glabrous, scabrid to smooth 

on angles, 1.2-2 mm wide. Leaves 4-8; sheaths 

unwinged, scabridulous to glabrescent, scabrid on 

angles, light brown, often tinged with red; ligule 

absent; contraligule short, convex to subtruncate, 

often callous-thickened medially, retrorsely 

pubescent, narrowly scarious on margin; blades 

plicate, 4-45 cm × 1.4-3.4 mm, often scabridulous 

on adaxial surface, margins antrorsely scabrid to 

smooth, the apex acuminate to blunt subflattened 

tip. Inflorescence composed of a single, small, 

terminal panicle, 1.5-3 cm in diam., or sometimes 

a second smaller lateral one below, from the upper

374 


reduced leaf-like bracts; lowest panicle branch 

often divergent and at a right angle to main axis; 

bract of terminal panicle erect, often exceeding 

panicle, up to 13 cm long; bractlets linearsetaceous, 

smooth to scabrid on margins; spikelets 

bisexual (androgynous); staminate spikelets 4-5 

× 1-1.5 mm; pistillate spikelets oblong-lanceolate, 

subcompressed, 3-5 × 1-2 (spreading to 5 mm); 

sterile pistillate spikelet scales 3, ovate-lanceolate, 

3-4.5 × 1.4-2.2 mm, light brown, finely reddish 

brown lineolate, brown at base or sometimes along 

the narrowly scarious margins, carina 3-nerved, 

often obscurely so, greenish, mucronate at the 

acute to acuminate apex. Staminate flowers with 

3 stamens, the anthers 1.9-2.6 mm long, with a 

very fine subulate-prickly apiculum. Achene body 

ovoid, obtusely trigonous to subrounded, 2-2.6 × 

1.6-2 mm, obtuse to acute at apex, short-apiculate, 

estipitate, eporose, shallowly reticulate to nearly 

smooth, glabrous, white; style base deciduous; 

hypogynium trigonous, with a pearl-colored 

pebbly crust on sides between the decurrent costae 

of the achene body. 


Puerto Rico. 

Distribution in Puerto Rico: On dry or damp 

serpentine soils or shale, in stony thickets, barrens, 

hillsides, forest edges and trails, and roadside 

slopes. Arecibo, Maricao, Mayagüez, Sabana 

Grande, San Germán, and Yauco. 


Arecibo: Santana, Vélez 1398, 1399 (US). 

Maricao: Rd. 119 to San Germán, km 15.3, 

González-Más 1657, 1659 (US); Forest Descanso 

trail, González-Más 1636 (US). Mayagüez: Las 

Mesas, near Mayagüez, Holm 41 (US). 

8.Scleria hirtella Sw., Prodr. 19. 1788; 

Hypoporum hirtellum (Sw.) Nees, Linnaea 9: 

303. 1834. Type: Jamaica. Swartz s.n. 

(holotype: BM; isotypes: B, M, S-Sw. R- 

5625). 

Scleria doradoensis Britton, Bull. Torrey Bot. Club 

50: 55. 1923. Type: Puerto Rico. Britton et 

al. 7064 (holotype: NY!; isotypes: UPR!, 

US!). 

Slender loosely caespitose annual, 10-55 

(-70) cm tall. Culms erect, unbranched, triquetrous, 

soft, flexible, glabrous, smooth on angles, 0.6-1.5 

mm wide. Leaves 3-5 per culm; sheaths elongate, 

unwinged, reddish or the uppermost tinged with 

red, often with sparse translucent hairs, the angles 

smooth; ligule absent; contraligule indistinct, with 

a tuft of translucent hairs, the convex margin 

sometimes with an inconspicuous scarious 

appendage; blades flattened-plicate, lowermost on 

culm with short plicate or 2-parted blades, 2-20 

cm × 1.5-3 mm, glabrous or essentially so, margins 

and abaxial midvein smooth proximally, antrorsely 

scabrous distally, the apex acuminate. 

Inflorescence terminal, interruptedly glomeratespicate 

with no lateral branching, composed of 3- 

10 (-13), sessile, axillary partial fascicles of 

spikelets, these 4-7 mm diam.; bractlets coarsely 

ciliate, often with purplish brown hairs; spikelets 

bisexual (androgynous), 2.5-3.5 × 1-2 mm; 

staminate scales brown, tinged with brown-black 

distally; sterile pistillate spikelet scales 3, ovate 

to ovate-lanceolate, 1.7-2.5 × 0.8-1.4 mm, 

(excluding awn), stramineous to golden brown, 

reddish lineolate, stained with brown-black on 

sides and margins, carina 1-nerved, often strigose, 

prolonged beyond the acute to acuminate apex as 

a curved, strigose awn. Staminate flower with 2 

stamens, the anthers 0.9-1.3 mm long, with a black 

prickly apiculum. Achene globose to subglobose, 

or obloid, rounded in cross section, 1.5-1.9 × 1- 

1.4 mm, smooth, sparsely and obscurely 

tuberculate, or distinctly verrucose-tuberculate, 

rounded to broadly rounded at apex, apiculate, 3to 

5-porose on each side, glabrous, shiny, bright 

white to gray, with a trigonous or triquetrous, stipelike 

base, the very base, just above its attachment, 

with a narrow, reddish or greenish, cellularreticulate 

band on each side; style base deciduous; 

hypogynium absent. 

General distribution: Central America, West 

Indies, Trinidad, and South America. 

Distribution in Puerto Rico: Wet, sandy soils 

in savannas, borders of lakes and marshes, grassy 

hillsides, forest edges, pasture, and roadside 

ditches. Aguada, Dorado, Manatí, Mayagüez, Vega 

Alta, Vega Baja, and Yabucoa. 

Note: The type of Scleria doradoensis is 

distinguished by the verrucose-papillate achenes, 

while those of S. hirtella are essentially smooth. 

However, intermediate achenes between these two 

stages are common, often occurring in the same 

population. The intermediate achenes exhibit 

various degrees of papillosity or verrucosity, but


the morphology of the trigonous base remains 

uniform. 


Aguada: Sargent 586 (US). Dorado: Vicinity of 

Dorado, N.L. Britton et al. 7064 (NY, UPR); Bo. 

Higuillar, Proctor 43225 (US); To Cerro Gordo, 

Rd. 693, km 13, González-Más 306 (US). Manatí: 

Bo. Tierras Nuevas Saliente, Axelrod et al. 10964 

(UPRRP). San Juan: Santurce, Heller & Heller 

591 (US). Vega Alta: Espinosa, Stevenson 2606 

(US). Vega Baja: Tortuguero Lagoon, Woodbury 

s.n., 12 Mar 1960 (NY); González-Más 1065 (US); 

González-Más & Reda 2164 (US). 

9. Scleria lithosperma (L.) Sw., Prodr. 18. 1788; 

Scirpus lithospermus L., Sp. Pl. 51. 1753. 

Lectotype: Rheede, Hort. Malab. 12: t. 48. 

1693, designated by Camelbeke & 

Goetghebeur, Taxon 49: 295. 2000. 

Scleria filiformis Sw., Prodr. 19. 1788; Scleria 

lithosperma var. filiformis (Sw.) Britton, Ann. 


Hispaniola. Swartz s.n. (holotype: probably 

S-Sw.). 

Scleria purpurea Poir. in Lamarck, Encycl. 7: 4. 

1806. Type: St. Thomas, U.S. Virgin Islands. 

Collector unknown (holotype: P-Lam.). 

Scleria krugiana Boeck., Beitr. Cyper. 1: 35. 1888. 

Type: Puerto Rico. Sintenis 4945 (holotype: 

B, destroyed; isotypes: L, NY!). 

Fig. 54. E-J 

Slender caespitose perennial, (20-) 50-90 

(-105) cm tall; rhizomes short, nodose, 2-4 mm 

thick. Culms ascending to erect, crowded together 

along nodes of the rhizome, trigonous, filiform, 

stiff, 0.5-2 mm wide, weakly antrorsely scabrous 

on the angles, pilose to essentially glabrous on 

sides. Leaves 6-10 per culm, shorter than the 

inflorescence; sheaths pilose to glabrescent, redtinged, 

purplish proximally; ligule absent; 

contraligule triangular or subrounded, the margin 

hispid; blades plicate, often inrolled along margins, 

2-25 (-35) cm × 1-3.5 mm, abaxially green, pilose 

to glabrescent, adaxially bluish green, shiny and 

glabrous except for the pilose midvein, antrorsely 

scabrous on margins and abaxial midvein, longacuminate 

to triquetrous apex. Inflorescence 

composed of a terminal and series of 1-3 remote, 

axillary, narrow partial panicles, the terminal one 

2-5 × 1-1.5 cm; bractlets subulate, to 3 cm long; 

panicle branches scabrous on margins; spikelets 

sessile or short-stalked, bisexual or unisexual; 

staminate spikelets 4-5 × 1 mm; fertile staminate 

scales 4-5 × 1 mm, brown; pistillate spikelets 

narrowly ovoid, 5-7 × 2-3 mm, typically bearing 

a reduced staminate spikelet just below attachment 

(cupula) of maturing fruit; sterile pistillate spikelet 

scales 4-5, ovate-lanceolate, 3-5 × 1.2-1.4 mm, 

thin, submembranous, glabrous to sparsely pilose, 

dark brown above, ciliolate along margins; carina 

1-nerved, lateral nerves indistinct, prolonged 

beyond the acuminate apex as a short to elongate 

mucro. Staminate flowers with 1 (-2) stamens, the 

anthers 1-2 mm long, with a triangular-subulate 

prickly apiculum. Achene ellipsoid to ellipsoidobovoid, 

obtusely trigonous, umbonate, 2-2.5 × 

1.5-1.8 mm, obtuse or sub-rounded at apex, 

apiculate, eporose, smooth, shiny, bone-white 

when mature, with a stipe-like, trigonous base 

bearing a depression between the costae on each 

side, the very base, just above its attachment, with 

a greenish or reddish cellular-reticulate ring 

surrounding it; hypogynium absent. 



Islands: Dry soils, often on limestone and 

serpentine, in rocky woodlands, wooded hillsides, 

thickets, forest trails, scrub forest, grassy slopes, 

barrens, rocky ravines, and roadsides. Arecibo, 

Barceloneta, Bayamón, Caja de Muertos, Cayo 

Ramos, Ceiba, Corozal, Dorado, Fajardo, Guánica, 

Guayama, Humacao, Isabela, Manatí, Maricao, 

Mayagüez, Mona Island, Orocovis, Patillas, 

Peñuelas, Ponce, Rincón, Río Grande, Sabana 

Grande, Salinas, San Germán, Vega Baja, Vieques, 

Yabucoa, and Yauco; St. Croix, St. James, St. John, 

St. Thomas, Tortola, and Virgin Gorda. 


Arecibo: Santana, Vélez 1396 (US). Bayamón: 

Liogier 10199 (US). Isabela: Reserva Forestal 

Bosque Guajataca, La Caballa Trail, Acevedo- 

Rdgz. & Siaca 11732 (UPR). Guánica: Guánica 

Forest along main road to Ranger Station, 

Acevedo-Rdgz. & Chinea 3025 (COL, JBSD, MO, 

US, NY, UPR, US). Mona Island: Camino del 

Diablo, Acevedo-Rdgz. et al. 4272 (SJ, US). 

Peñuelas: Sintenis 4753 (US). Ponce: 3 mi. W of 

Ponce, Heller 6244 (US). Rincón: Puerto Higiero, 

Atomic Energy Project Grounds, González-Más 

2045 (US). Sabana Grande: Susúa Reserve Forest, 

González-Más 2028 (US). San Germán: Sargent

376 


489 (US). Vega Baja: Liogier 9794 (US). Vieques 

Island: Calabaza to Ensenada, Shafer 2948 (US). 

Yabucoa: Sintenis 4945 (NY, US). Yauco: Susúa 

Forest Reserve, Axelrod et al. 4629 (US). ST. 

CROIX: District of Queen, Estate, Proctor 44970 

(US); Caledonia Gut, Fosberg 55396 (US); Signal 

hill, A.E. Ricksecker 440 (US). Frederiksted, 

Oersted s.n. (US). ST. JOHN: Bethamia to 

Rosenberg, N.L. Britton & Shafer 239 (US). ST. 

THOMAS: Oersted s.n. (US); Eggers s.n. (US); 

Cowells Hill, Eggers s.n. (NA); Raccoon Bay, N.L. 

Britton et al. 163 (US). TORTOLA: Scrub Island, 

Proctor 46566 (US). 

10. Scleria melaleuca Rchb. ex Schltdl. & Cham., 

Linnaea 6: 29. 1831. Type: Surinam. Weigelt 

s.n. (holotype: G; isotypes: F, G, HAL). 

Schoenus latifolius Vahl, Enum. Pl. 2: 226. 1805 

[non Scleria latifolia Sw., 1788]. Type: Puerto 

Rico. Herb. Ventenat s.n. (holotype: C-Vahl). 

Scleria pterota C. Presl in Oken, Isis 21: 268. 1828, 

nom. nud. 

Scleria pratensis Lindl. ex Nees in Martius, Fl. 

Bras. 2(1): 179. 1842. Type: Brazil. Macrae 

s.n. (holotype: probably at CGE). 

Scleria ottonis Boeck., Linnaea 38: 490. 1874. 

Type: Cuba. Otto 299 (holotype: B, 

destroyed). 


cm tall; rhizome short, horizontally creeping, 

thickened and nodose, 5-8 mm thick, purplish 

brown. Culms erect, borne singly along the 

rhizome, triquetrous, 2.5-5 (-8) mm wide, stiff and 

hardened, glabrous to sparsely pilose, scabrid on 

angles distally. Leaves 4-8, the upper ones 

equaling to exceeding the inflorescence; sheaths 

inflated towards apex, distal ones wing-angled, 

septate-nodulose to indistinctly so, retrorsely 

scabrous on margins, sparsely pilose to glabrous, 

purple-tinged at base; ligule absent; contraligule 

triangular, the thickened, cartilaginous margin 

hispid to glabrescent; blades flat, 20-60 cm × (3-) 

5-17 (-19) mm, 5-veined, obscurely septate near 

base, thin, glabrous or sparsely pilose proximally, 

antrorsely scabrous on margins and abaxial 

midvein, the apex short to abruptly acuminate with 

blunt tip. Inflorescence composed of a single 

terminal and 1-4 axillary, subcontiguous to remote, 

narrowly pryamidal partial panicles from the upper 

leaf-like bracts, the terminal one largest, 4-11 × 

1-4 cm; bractlets flattened, falcate, to 2 cm long; 

panicle branches wing-angled, scabrous on angles, 

often dark purplish; spikelets unisexual; staminate 

spikelets 3-5 × 1-1.5 mm; fertile staminate scales 

oblong-ovate to lanceolate, 3-4 × 1-2 mm; pistillate 

spikelets elliptic to elliptic-obovoid, 

subcompressed, 3-4 × 2-3 mm; sterile pistillate 

spikelet scales 3-4, widely-ovate to ovate-oblate, 

2.5-3 × 2-3.5 mm, glabrous, shiny, margins finely 

ciliate, brown, dark purple distally on sides, carina 

1-nerved, prolonged beyond the abruptly 

acuminate, cuspidate-divergent apex as a short, 

excurved mucro. Staminate flowers with 1 stamen, 

the anthers 1-1.3 mm long, with a dark brown, 

triangular-subulate, bristly apiculum. Achene body 

depressed-globose to globose, subumbonate, 1.5- 

2.2 (-2.9) × 2-2.5 mm, shorter than the scales, 

apiculate, eporose, estipitate, smooth or sometimes 

with tufts of fine hairs near base, white or 

variegated with purple; hypogynium 1-1.5 × 1.8- 

2 mm, 3-lobed, the lobes broadly rounded with 

entire margins, reflexed. 


America, South America, and the West Indies. 


Islands: Wet or moist places in wooded hillsides, 

forest edges, trails in secondary forests, roadsides 

ditches, roadbanks, grassy slopes, marshy areas, 

meadows, pastures, stream borders, lake margins, 

sand savannas, seepage areas, scrub forests, and 

waste grounds. Adjuntas, Aguas Buenas, Arecibo, 

Bayamón, Caguas, Canóvanas, Cataño, Cayey, 

Carolina, Ceiba, Cidra, Corozal, Dorado, Fajardo, 

Guayama, Guaynabo, Humacao, Isabela, Maricao, 

Maunabo, Mayagüez, Naguabo, Patillas, Río 

Grande, San Germán, San Juan, San Lorenzo, San 

Sebastián, Toa Baja, Trujillo Alto, Utuado, Vega 

Alta, Vega Baja, and Yabucoa; St. Croix, St. John, 

St. Thomas, Tortola, and Virgin Gorda. 

Common name: Puerto Rico: Cortadora 

blanca. 



10554 (UPRRP, US). Canóvanas: Bo. Cubuy, Trejo 

et al. s.n. (UPRRP). Cataño: Bo. Palo Seco, 

Axelrod & Díaz 12708 (UPRRP). Cayey: Rd. 184 

to Guavate-Carite, km 3.7, González-Más 1564 

(US). Cidra: Pueblo Viejo, Hioram 93 (US). 

Humacao: Santa Teresa, Liogier et al. 31348 (US). 

Isabela: Bo. Planas, Guajataca Forest Reserve, 

Axelrod et al. 11249 (UPRRP). Maricao: Rd. 120,


near San Antonio River, González-Más 411 (US). 

Mayagüez: Miradero, near aqueduct, González- 

Más 1762 (US). Naguabo: Bo. Florida, Rd. 191, 

km 27.5, González-Más 1388 (US); Sierra de 

Naguabo, along Río Cubuy to Meseta Falls, Shafer 

3154 (US). Río Grande: Along Rt. 186 bordering 

Caribbean National Forest, Kelloff et al. 372 (US) 

San Germán: Sargent 751 (US); San Juan: Río 

Piedras, Stevenson 75 (US); Santurce, Heller & 

Heller 15 (US). San Sebastián: Sargent 239 (US). 

Utuado: Vicinity of Utuado, N.L. Britton & Cowell 

888 (US). ST. CROIX: SE ridge, Mt. Eagle, Fosberg 

60776 (US); Lebanon Hill, A.E. Ricksecker 416 

(US). ST. JOHN: Bordeaux Mountain, Woodbury 57/ 

6543, 58/6544 (VINPS); Bethania to Rosenberg, 

N.L. Britton & Shafer 275 (US). ST. THOMAS: 

Crown, N.L. Britton & Marble 1342 (US); 

Charlotte Amalie, Millspaugh 506 (US); 

Signalhill, Eggers s.n. (NA); Eggers s.n. (US). 

TORTOLA: High Bush, N.L. Britton & Shafer 832 

(US). 

11. Scleria microcarpa Nees ex Kunth, Enum. 

Pl. 2: 341. 1837; Ophryoscleria microcarpa 

(Nees ex Kunth) Nees in Martius, Fl. Bras. 

2(1): 184. 1842. Type: Surinam. Weigelt s.n. 

(holotype: B, destroyed; isotypes: BM, BR, 

HAL). 

Scleria foliosa C. Wright in Sauvalle, Anales Acad. 

Ci. Méd. Habana 8: 154. 1871, non 

Hochstetter ex A. Richard, 1850; Scleria 

microcarpa var. foliosa C. B. Clarke in Urban, 

Symb. Antill. 2: 149. 1900. Type: Cuba. 

Wright 3807 (holotype: GH, isotype: NY). 

Slender or medium-sized rhizomatous 

perennial, 55-160 (-210) cm tall; rhizome short to 

elongate, horizontal, scaly, knotty at culm bases, 

3-6 mm thick. Culms erect to bending over at 

maturity, approximate to remote along rhizome, 

trigonous proximally, triquetrous distally, smooth 

and glabrous, scabrid on angles (at least distally), 

3-6 mm wide. Leaves 5-8; sheaths winged, 

glabrous or pubescent, the angles retrorsely 

scabrous, or often with mixed antrorse and retrorse 

barbs; ligule absent; contraligule lanceolate, 

glabrous, with straight veins, the margin ciliatescabrous 

to smooth; blades subflattened to plicate, 

2-45 cm × 4-13 (-16) mm, glabrous or pubescent, 

margins, abaxial midvein, and sometimes adaxial 

lateral veins antrorsely scabrous (at least distally) 

or with mixed antrorse and retrorse barbs 

proximally, acuminate or attenuate to blunt, 

subflattened tip. Inflorescence composed of a 

single terminal and 1-5 linear to linear-rhombic 


bracts, the terminal one 3-16 × 0.5-1 (-2) cm; 

bractlets linear-subulate, antrorsely scabrous on 

margins; branches narrowly winged; spikelets 

unisexual; staminate spikelets 1.5-2.5 × 0.8-1.3 

mm; staminate scales light brown; pistillate 

spikelets ovate, 2-3 × 1-1.5 mm; sterile pistillate 

spikelet scales 4, broadly ovate to rounded, 

curvate-keeled, 0.9-1.4 mm diam., straw-colored 

to dark brown, reddish brown lineolate, carina 1nerved, 

shortly prolonged at acute apex. Staminate 

flower with 3 stamens, the anthers 0.7-1.1 mm 

long, with a triangular-subulate, prickly apiculum. 

Achene body ovoid to ellipsoid-ovoid, 1-1.8 × 1.2- 

1.9 mm, slightly compressed, obtuse to subrounded 

at apex, estipitate, eporose, smooth, 

glabrous, white or greenish; style base persistent, 

short-conic, 0.3-0.5 × 0.2-0.3 mm, brown, finely 

antrorsely scabrid; hypogynium 3-lobed, adhering 

to and hidden by the spongy-thickened, tan to 

yellowish, dark brown mottled, shallowly 3-lobed 

cupula, 0.8-1.3 × 1.3-2 mm, which is finely ciliate 

on its rim, the hairs whitish or yellow-brown. 


Central America, West Indies, Trinidad, and South 

America. 

Distribution in Puerto Rico: In wet or moist 

often sandy soils in forest edges, second-growth 

forests, forest openings, marshy areas, lake 

borders, creek and stream edges, trails, scrubs, 

thickets, savannas pastures, and roadside ditches. 

Arecibo, Cabo Rojo, Carolina, Dorado, Fajardo, 

Florida, Luquillo, Manatí, Mayagüez, Naguabo, 

Patillas, Río Grande, San Juan, Toa Baja, Utuado, 



Cabo Rojo: Sintenis 78 b (US). Dorado: Bo. 

Higuillar, just E of Aeropuerto de Dorado, Proctor 

42447 (US). Río Grande: Luquillo Mountains, 

Wilson 218 (US). San Juan: Stone quarry 2 mi. E 

of Santurce, Heller & Heller 1362 (US); Río 

Piedras, Stevenson 3651 (US). Toa Baja: Sabana 

Seca, Primate Research Center, Trejo 1107 

(UPRRP).Vega Baja: Tortuguero Lake, González- 

Más 2160 (US). Bo. Puerto Nuevo, González-Más 

1094 (US).

378 


12. Scleria mucronata Poir. in Lamarck, Encycl. 

7: 3. 1806. Type: Puerto Rico. Ledrú 83 p.p. 

(holotype: P!; isotype: P). 

Scleria cubensis Boeck., Beitr. Cyper. 2: 42. 1890. 

Type: Cuba. Eggers 5113 (holotype: B, 

destroyed; isotype: P) 

Scleria microcarpa var. subeciliata C. B. Clarke 

in Urban, Symb. Antill. 2: 149. 1900. 

Lectotype: Jamaica. Hart 674 (US!), here 

designated. 

Scleria catalinae Britt., Bull. Torrey Bot. Club 42: 

489. 1915. Type: Cuba. Van Hermann 3272 



tall; rhizome thick and hardened, nodose. Culms 

erect, sharply trigonous to triquetrous distally, 3- 

9 mm wide, glabrous, smooth proximally, 

scabridulous on sides and angles distally. Leaves 

6-8; sheaths unwinged or only very narrowly so, 

scabrid to smooth on angles, glabrous; ligule 

absent; contraligule triangular or narrowly so, with 

rounded tip, glabrous, the margin thickened, often 

crenulate; blades subflattened or plicate, 30-80 cm 

× 9-37 mm, adaxial and abaxial surfaces glabrous, 

margins, abaxial midvein, and abaxial lateral 

nerves finely antrorsely scabrous to essentially 

smooth, abruptly narrowed at base, not decurrent 

or only very narrowly so along the lateral margins 

of the sheath, acuminate or subattenuate to 

subflattened, blunt tip. Inflorescence composed of 

a terminal and series of 2-4 narrowly rhombic, 


bracts, the terminal one largest, 9-28 × 2-6 (-8) 

cm; bractlets linear, scabrid on angles; panicle 

branches finely pubescent to glabrous; spikelets 

unisexual; staminate spikelets ovate to ovateelliptic, 

2.5-3.5 × 1-1.5 mm; staminate scales 

brown to light reddish brown, grayish puberulent; 

pistillate spikelets ovoid, 2-3 mm long; sterile 

pistillate spikelet scales 4, ovate-orbicular, the 

uppermost 2.5-3 mm in diam, brown, grayish 

puberulent to glabrous, finely brown-lineolate, 

carina 1-nerved, shortly prolonged beyond the 

acute to broadly acute apex. Staminate flower with 

2-3 stamens, the anthers 1-1.3 mm long, with a 

triangular-subulate, bristly apiculum. Achene body 

ovoid to subglobose or globose, 1.8-2.2 × 2.1-2.5 

mm, rounded to obtusely so or truncate at apex, 

smooth, glabrous, white to grayish white or bluegreen, 

estipitate, eporose; style base persistent, 

conic-cylindrical, 0.6-1 × 0.5-0.6 mm, finely 

reticulate, smooth or essentially so; hypogynium 

3-lobed, adhering to and hidden by the spongythickened, 

tan to yellowish, smooth, shallowly 3lobed 

cupula, 1.3-1.5 × 2.2-2.7 mm, which is 

sparsely and inconspicuously ciliate on its rim, the 

hairs very short and fine. 


Jamaica, and Puerto Rico. 

Distribution in Puerto Rico: Montane forests 

on lateritic soils and on serpentine substrate at 

Maricao; wet or damp areas on forest edges, trails, 

glades, river banks, creek margins, rocky slopes 

and ravines, thickets, hillsides, and roadbanks. 

Barranquitas, Cayey, Juana Díaz, Maricao, 

Naguabo, Río Grande, and Villalba. 


Barranquitas: Sargent 362 (US); La Torrecilla, 

Axelrod et al. 5327 (US); Monte Torrecilla, N.L. 

Britton et al. 5536 (US). Maricao: Bo. Indiera Fría, 

along track near Río Lajas, Proctor & Padrón 

45677 (US). Naguabo: Sierra de Luquillo: Summit 

of Loma La Mina to Río Blanco, Shafer 3384 (US). 

Río Grande: El Yunque, Sargent 544 (US). 

Villalba: near Salto Doña Juana, González-Más 

2195, 2196 (US). Juana Díaz: Sargent 3198 (US). 

13. Scleria muehlenbergii Steud., Nomencl. Bot., 

ed. 2, 2: 543. 1841; Scleria laxa Torr., Ann. 

Lyceum Nat. Hist. New York 3: 376. 1836, 

non R. Brown, 1810. Type: United States; 

South Carolina. Collector unknown (holotype: 

PH). 

Scleria hemitaphra Steud., Syn. Pl. Glumac. 2: 

169. 1855. Type: United States; Louisiana, 

Drummond Herb. s.n. (holotype: probably at 

P). 

Scleria reticularis var. pubescens Britton, Ann. 


Cuba. C. Wright 3800 (holotype: NY!; 

isotypes: GH, US!) 

Scleria stevensiana Britton, Bull. Torrey Bot. Club 

50: 55. 1923. Type: Puerto Rico; Lares. Steven 

& Hess 4944 (holotype: NY!). 

Scleria setacea sensu many authors, non Poiret, 

1806. 

Slender to medium-sized, caespitose 

perennial, 20-80 (-100) cm tall; rhizome short, 

nodose, 1-2 mm thick. Culms erect or ascending, 

triquetrous, retrorsely scabrid to smooth on angles, 

otherwise glabrous, 0.8-2.3 mm wide, to 0.5 mm 

wide distally. Leaves 2-5 (excluding upper leaf-


like panicle bracts, remote; sheaths inflated, 

narrowly winged, often hirsute proximally, 

remotely cross-veined, retrorsely ciliate-scabrid on 

angles, purplish proximally; ligule absent; 

contraligule obtuse to rounded, with straight veins, 

the margin with a short scarious appendage; blades 

subflattened to plicate, 4-30 (-37) cm × (1.4-) 1.7- 

4 (-5) mm, glabrous or nearly so, margins and 

abaxial midvein antrorsely scabrid, the apex 

acuminate or attenuate to blunt tip. Inflorescence 

composed of a terminal and series of 2-4 narrow 

partial panicles from the upper leaf-like bracts, on 

short to elongate peduncles, the terminal one 1.5- 

3 (-5) × 0.6-1.8 cm; spikelets unisexual; staminate 

spikelets 2.5-3.5 × 0.7-1 mm; staminate scales light 

brown to dark reddish brown; pistillate spikelets 

ovoid-lanceoloid, 4-6 × 1-1.7 mm; sterile pistillate 

spikelet scales 3, ovate to ovate-lanceolate, 2.7- 

4.5 × 2-2.5 mm, light brown to dark reddish brown 

or purplish, carina 1-nerved, antrorsely scabrous 

distally, prolonged beyond the acuminate apex as 

a mucro. Staminate flower with 2 stamens, the 

anthers 0.8-1.4 mm long, with a prickly apiculum. 

Achene body broadly ovoid to subglobose, 1.7- 

2.5 × 1.5-2 mm, irregularly reticulate-pitted or 

papillose-reticulate, with dense to sparse tufts of 

minute rusty hairs on the ridges, dingy white or 

grayish, estipitate, the apex rounded, abruptly 

apiculate; style base deciduous; hypogynium 

deeply 3-lobed, 0.7-1 × 1.2-1.5 mm, the lobes 

oblong, concave with a low ridge medially, 

appressed to base of achene body, with an acute 

to attenuate tooth at apex. 

General distribution: United States, Central 

America, West Indies, Trinidad, and South 

America. 

Distribution in Puerto Rico: On moist or wet 

sands around pond margins, savannas and 

savanna-like swales, and roadsides. Aguada, 

Añasco, Dorado, Manatí, Moca, Rincón, Vega 


Note: Scleria muehlenbergii has been treated 

by many authors as a synonym of the closely 

related S. reticularis Michx. However, a recent 

proposal (Camelbeke, Reznicek, & Goetghebeur, 

Taxon 52: 355. 2003) was made to conserve the 

name Scleria reticularis Michx. with a conserved 

type based on a discovery that the type material 

of that name is referable to a different species, 

Scleria triglomerata Michx. If the proposal is 

accepted by the spermatophyte committee, the 

proposed type will fix the name in a strict sense to 

apply to an element which is restricted to the 

eastern United States. After examining the 

specimen proposed for conservation (North 

Carolina. Leblond 1651 (NCU) and other material 

of S. reticularis sensu stricto, we concluded that 

this taxon was not conspecific with the Puerto 

Rican material which we are here applying in the 

strict sense to the more widespread S. 

muehlenbergii. 


Aguada: Sargent 596 (US). Dorado: Vicinity of 

Dorado, N.L. Britton et al. 6654 (US). Manatí: 

Laguna Tortuguero, May 1960, Woodbury s.n. 

(US). Vega Alta: Bo. Sabana, Proctor et al. 44146 

(US). Vega Baja: Bo. Algarrobo, just S of Laguna 

Tortuguero, Proctor et al. 45030 (US). 

14. Scleria scindens Nees ex Kunth, Enum. Pl. 2: 

343. 1837. Type: Martinique. Kohaut 269 

(holotype: probably at AWH). 

Scleria chlorantha Boeck., Linnaea 38: 506. 1874. 

Type: Dominican Republic. Mayerhoff s.n. 


Scleria scaberrima Boeck., Beitr. Cyper. 2: 41. 

1890. Type: St. Vincent. Eggers s.n. (holotype: 


Fig. 56. C-E 

Robust perennial, 0.5-3.5 m tall, often 

growing in large colonies; rhizome stout, nodose. 

Culms erect or ascending, vine-like and 

clambering, solitary from nodes of the rhizome, 

sharply 3-angled, (2-) 3.2-8 cm wide, mixed 

retrorsely-antrorsely scabrid on angles, sparsely 

strigose on sides, glabrescent at base. Leaves 

numerous, overtopping the inflorescence; sheaths 

subloose, inflated near summit, appressed-strigose, 

glabrescent; ligule absent; contraligule convex to 

truncate, with straight veins, the margin 

cartilaginous, often ciliate; blades subflattened or 

plicate, 30-90 cm × 5-15 mm, obscurely septate, 

stiff, somewhat coriaceous, blunt to sharply 

retrorsely scabridulous on margins and abaxial 

midvein, the apex acuminate to subflattened, blunt 

tip. Inflorescence composed of a single, dense, 

pyramidal panicle, 3-8 (-10) cm diam., or 

sometimes with 1 or 2 additional smaller panicles 

in the axils of the upper leaf-like bracts; bracts 

subtending the terminal panicle typically 3, 

overtopping the inflorescence; bractlets setaceous, 

ciliate-scabrous; panicle branches minutely 

pubescent, glabrescent; staminate spikelets

380 


narrowly oblong-ovoid, cylindrical to slightly 

compressed, ca. the same length as the pistillate; 

staminate scales many, the 2-3 at base sterile, like 

the pistillate, 3-4 × 2-3 mm; pistillate spikelets 

ovoid-elliptic, 4-5 × 2-3 mm, spreading with 

developing achene; sterile pistillate spikelet scales 

4, ovate-orbicular, the uppermost one 3.5-4 mm 

in diam., cupuliform, glabrescent, green, finely 

reddish brown lineolate, ciliate on margins, carina 

1-nerved, mucronate at the short-acuminate to 

acute apex. Staminate flowers with 3 stamens, the 

anthers 1.8-2.2 mm long, with a triangularsubulate, 

prickly apiculum. Achene body globose, 

2.3-2.6 × 2.3-2.7 mm, minutely apiculate, shiny 

white to purple-tinged; hypogynium 1-1.7 × 2.3- 

2.7 mm, 3-lobed, with weakly revolute, rounded, 

entire, spreading to divergent lobes which are not 

appressed to base of achene body. 


Virgin Islands, and the Lesser Antilles. 

Distribution in the Virgin Islands: Secondary 

vegetation along roadsides, edges of forested 

areas, hillsides, and streams. St. John, Tortola, and 

Virgin Gorda. 

Common name: Virgin Islands: Razor-grass. 

Selected specimens examined: ST. JOHN: Coral 

Bay Quarter; along dirt road to Bordeaux 

Mountain, Acevedo-Rdgz. et al. 5102 (K, MO, NY, 

UPRRP, US). TORTOLA: Sage Mountain, Fishlock 

91, 379 (NY); High Bush, N.L. Britton & Shafer 

822 (NY, US). 

15. Scleria secans (L.) Urb., Symb. Antill. 2: 169. 

1900; Schoenus secans L., Syst. Nat. ed. 10: 

865. 1759. Lectotype: Jamaica. Sloane, Voy. 

Jamaica 1: t. 77, f. 1. 1707, designated by T. 

Koyama in R.A. Howard, Fl. Lesser Antill. 

3 : 309. 1979. 

Scleria reflexa Kunth in Humb., Bonpl., & Kunth., 

Nov. Gen. Sp. 1 [quarto ed.]: 232. 1816; 

Mastigoscleria reflexa (Kunth) Nees in 

Martius, Fl. Bras. 2(1): 177. 1842. Type: 

Venezuela. Humboldt & Bonpland s.n. 


Scleria flagellum sensu many authors, non Swartz, 

1788. 

Fig. 56. F, G 

Vine-like, rhizomatous perennial, elongating 

to 5-10 m, often forming dense thickets; rhizome 

coarse, knotty. Culms scandent, scrambling and 

climbing over secondary vegetation, frequently 

branched from upper nodes, trigonous proximally, 

triquetrous distally, often hirsute on sides medially 

to glabrous, retrorsely spinulose on angles, 1.8-4 

mm wide, to 6 mm wide at base. Leaves numerous; 

sheaths tight, unwinged or essentially so, hirsute 

medially, the angles smooth or retrorsely 

spinulose; ligule a dense band of hairs; contraligule 

deltate with obtuse apex, often densely hirsute with 

whitish trichomes, the margin often bearing a 

broad scarious appendage with anastomosing 

veins, or sometimes wanting; blades plicate or 

subflattened, 10-40 cm × 2-7 mm, adaxial surface 

hirsute at base, abaxial surface sparsely hirsute to 

glabrous, margins, abaxial midvein, and often 

adaxial main lateral nerves retrorsely spinulose, 

gradually tapering to and attenuate, subflattened 

apex. Inflorescence composed of a terminal and 

series of 1-3, axillary, pyramidal partial panicles, 

from the upper leaf-like bracts, the terminal one 

largest, 2.5-6 (-8) × 1.3-4.5 cm, the lower 

becoming successively smaller or wanting; 

bractlets and rachis ciliate-hirsute on margins or 

angles; spikelets unisexual; staminate spikelets 3- 

4.5 × 0.8-1.2 mm; proximal staminate scales 

greenish, stained with reddish or blackish purple 

on margins and sometimes sides distally; pistillate 

spikelets ovoid, 5-8 × 1.5-2 mm (spreading to 5 

mm with developing achene); sterile pistillate 

spikelet scales 4, ovate to ovate-lanceolate, 2.5-6 

× 2-4 mm, light brown to brown, reddish or 

blackish purple on margins and sometimes sides 

distally, carina 1-nerved (at least distally), 

mucronate at the acute to acuminate apex. 

Staminate flowers with 3 stamens, the anthers 2- 

2.5 mm long, with a lanceolate-setaceous 

appendage at apex, often bearing some appressed 

trichomes. Achene body ovoid to oblong-ovoid, 

rounded, 2.8-4 × 2.4-2.9 mm, obtuse at apex, white 

or sometimes green-mottled, smooth, glossy, 

glabrous or essentially so, estipitate, eporose; style 

base deciduous or its base sometimes persisting 

as a bulbous, antrorsely scabrid tubercle; 

hypogynium discoid, obscurely 3-lobed, 0.3-1 × 

1.5-2 mm, the lobes entire, reflexed, irregularly 

wrinkled, reddish lineolate. 


America, West Indies, Trinidad, Tobago, and South 

America. 

Distribution in Puerto Rico: Secondary 

thickets in openings, along trails, and borders of


Fig. 56. A-B. Scleria canescens. A. Fertile branch, with details of the node and leaf margin. B. Achene with scales and achene 

showing hypogynium. C-E. Scleria scindens. C. Branch, showing detail of the node. D. Inflorescence. E. Achene showing 

hypogynium. F-G. Scleria secans. F. Fertile branch, with detail of the node. G. Achene with hypogynium. From Acevedo-Rdgz., 

P. 2003, Bejucos y plantas trepadoras de Puerto Rico e Islas Vírgenes, Smithsonian Institution.

382 


lower montane forest, 440-700 m elevation. Also 

collected at Laguna Tortuguero (no Municipality 

given) by Woodbury s.n., 3 Mar 1960 (UPR). 

Bayamón, Caguas, Cayey, Guayama, Luquillo, 

Patillas, and Río Grande. 

Common name: Puerto Rico: Cortadora de 

altura. 


Bayamón: Bo. Guaraguao Arriba, Proctor 41703 

(SJ). Caguas: Kuntze 316 (NY); N.L. Britton & 

Cowell 1402 (US, NY). Cayey: Sierra de Cayey, 

Bo. Guavate, Proctor 47938 (SJ). Guayama: Carite 


Luquillo: Bo. Sabana, Caribbean National Forest, 

Axelrod 5848 (UPRRP). Patillas: on property of 

Tropic Ventures, Taylor 7847 (UPRRP); Carite 

Forest Reserve, Rt. 184, Axelrod 5873 (UPRRP). 

Río Grande: Sierra de Luquillo, Proctor & 

Alemany 47720 (SJ). 

16. Scleria verticillata Muhl. ex Willd., Sp. Pl. 

4: 317. 1805; Hypoporum verticillatum 

(Muhl. ex Willd.) Nees, Linnaea 9: 303. 1834. 

Type: United States; Pennsylvania. Willdenow 

s.n. (holotype: B-Willd. 17326). 

Slender caespitose annual, 10-70 cm tall, with 

aromatic roots. Culms ascending to erect, 

triquetrous to subtrigonous with smooth, pale ribangles, 

firm, flexuous, glabrous, 0.4-2 mm wide. 

Leaves 5-8 per culm; sheaths elongate, unwinged, 

subloose, glabrous to sparsely pilose, the basal 

ones often reddish purple or purple-black; ligule 

absent; contraligule small, truncate or slightly 

convex, hirsute or glabrous, or wanting; blades 

plicate to subflattened, 3-30 cm × 0.7-3 mm, 

glabrous, margins and abaxial midvein smooth 

proximally, finely antrorsely scabrous distally, 

attenuate to apex. Inflorescence terminal, 

interruptedly glomerate-spicate, unbranched or 

sometimes with 1-2 lateral branches present, 

composed of 3-10 (-15), sessile fascicles of 

spikelets; bractlets filiform, glabrous; rachis 

narrowly wing-angled; fascicles 3-7 mm diam., 

erect or patent, not reflexed; spikelets bisexual 

(androgynous), 2.5-4 × 1-2 mm; staminate scales 

lanceolate; sterile pistillate spikelet scales 3, ovatelanceolate 

to oblong-lanceolate, 2.2-3 × 0.8-1.2 

mm, glabrous, stramineous or purplish, carina 3nerved, 

greenish, prolonged beyond the 

acuminate-cuspidate apex as an excurved mucro. 

Staminate flower with 1 stamen, the anther 1-1.5 

mm long, with a lanceolate-subulate prickly 

apiculum. Achene subglobose to depressed 

globose, 1-1.5 × 1-1.3 mm, verrucose, rounded to 

subtruncate at apex, apiculate, 4- or 5-porose on 

each side basally, glabrous, white to grayish or 

purple-tinged, with a trigonous, stipe-like base 

bearing a depression between the costae on each 

side, the very base, just above attachment, with a 

narrow, greenish or reddish cellular-reticulate band 

on each side; style base deciduous; hypogynium 

absent. 



Distribution in Puerto Rico: Wet sands in 

savanna-like swales or seasonal pools. Manatí and 

Vega Baja. Known to occur only in the vicinity of 

Laguna Tortuguero. 


Without locality, 1970, Woodbury s.n. (US). 


Scleria bracteata Cav., Icon. 5: 34. 1799. Reported 

for Puerto Rico by Kartesz (1998) in “A 

Synonymized Checklist of the Vascular Flora 

of the United States, Puerto Rico, and the 

Virgin Islands” (http://www.csdl.tamu. edu/ 

FLORA/b98/check98.htm). This record was 

based on a pers. comm. from John Fairey 

(CLEM) in a letter to John Kartesz dated May 

11, 1979. No specimens were seen during the 

preparation of this treatment nor has it been 

recently collected in the flora area. 

Scleria mitis P. J. Bergius, Kongl. Vetensk. Acad. 

Handl. 26: 145. 1765. Cited for Puerto Rico 

by Core (1936) based on Sintenis 1215 (US). 

However, two sheets of this collection seen 

are both clearly Scleria eggersiana Boeck. 

Even though one of the specimens has 

immature achenes, the cilia along the margin 

of the hypogynium are brown and the style 

base is finely antrorsely scabrid, characteristic 

of S. eggersiana. Scleria mitis has dense, 

black cilia on the margin of the hypogynium 

and coarse, appressed, retrorse black 

trichomes on the style base. Urban (1903) 

cites Sintenis 1714 and 6639, and Eggers & 

Rensch 1180b as S. mitis from Puerto Rico. 

Sintenis 1714 and 1215 are S. eggersiana. 

Sintenis 6639 and Eggers & Rensch 1180b 

were not seen. No collections of this species 

from Puerto Rico and the Virgin Islands were


seen during the preparation of this treatment 

nor has it been recently collected in the flora 

area. 

Scleria oligantha Michx., Fl. Bor.-Amer. 2: 167. 

1803. Cited for Puerto Rico by Liogier & 

Martorell (1982: 221; 2000: 246). No 

specimens were seen during the preparation 

of this treatment nor has it been recently 

collected in the flora area. 

Scleria pauciflora Muhl. ex Willd., Sp. Pl. 4: 318. 

1805. Cited for Puerto Rico by Liogier & 

Martorell (1982: 221; 2000: 246). No 

specimens are known to substantiate this 

record. Records of this species are probably 

based on misidentifications of specimens of 

Scleria brittonii Core at UPR. 

Scleria purdiei C. B. Clarke, Kew Bull. Addit. Ser. 

8: 57. 1908. Reported for Puerto Rico by 

Kartesz (1998) in “A Synonymized Checklist 

of the Vascular Flora of the United States, 

Puerto Rico, and the Virgin Islands” (http:// 

www.csdl.tamu.edu/FLORA/b98/ 

EXCLUDED GENERA 

check98.htm). This record was based on a 

pers. comm. from John Fairey (CLEM) in a 

letter to John Kartesz dated May 11, 1979. 

No specimens were seen during the 



Scleria scabriuscula Schltdl., Linnaea 20: 544. 

1847. Reported for Puerto Rico by Kartesz 

(1998) in “A Synonymized Checklist of the 

Vascular Flora of the United States, Puerto 

Rico, and the Virgin Islands” (http:// 

www.csdl.tamu.edu/FLORA/b98/ 

check98.htm). This record was based on a 

pers. comm. from John Fairey (CLEM) in a 

letter to John Kartesz dated May 11, 1979. 

No specimens were seen during the 



Scleria triglomerata Michx., Fl. Bor.-Amer. 2: 

168. 1803. Records of this species are based 

on misidentifications of Scleria havanensis 

Britton. 

Isolepis disticha Boeck., Flora 41: 415. 1858. (syn: Scirpus subdistichus Boeck. which is a later 

name based on the same collection). The identity of this taxon, based on Bertero 248 (B) from St. 

Thomas, is unknown. According to Britton & P. Wilson (1923), C. B. Clarke examined the specimen 

and thought it might perhaps be Scirpus cernuus Vahl (Isolepis cernua (Vahl) Roem. & Schult.), a plant 

otherwise not known to occur in the West Indies. This plant was described as having subdistichous 

spikelet scales, while I. cernua has spirally imbricate spikelet scales. In their list of nomina exclusa, 

Muasya and Simpson (2002) indicate that the taxon is a Bulbostylis. As to description, this plant fits 

Bulbostylis pauciflora quite well except that the achene of Scirpus subdistichus is described as having a 

short, hard mucro at its apex and there is no mention of the leaves having a pilose sheath apex. Bulbostylis 

achenes typically have a persistent, bulbous style base at apex and the leaf sheaths have pilose apicies. 

The description of the achene as described for Scirpus subdistichus better fits Cyperus which typically 

has distichous spikelet scales, a puncticulate achene epidermis, and an apiculate achene apex. 

Family 33. TYPHACEAE Cat-tail Family 

Typhaceae Juss., Gen. Pl. 25. 1789, nom. conserv. 

by G. R. Proctor and M. T. Strong 

Tall, erect, perennial, monoecious herbs of marshy or aquatic habitats, with creeping rhizomes and 

erect stems and leaves. Leaves basal, long-linear, alternate, distichous, sheathing at base, glabrous. 

Flowers unisexual and densely arranged in separate cylindrical spikes on the same stalk, the staminate

384 


Fig. 57. A-F. Typha domingensis. A. Habit. B. Inflorescence. C. Cluster of young anthers. D. Bract. E. Fertile pistillate flower. F. 

Fertile ovary. (A, B, D, E, and F, from Proctor 50610; C, from Proctor 46202).


Fig. 58. A. Podocarpus coriaceus, fruit. B. Pinus caribaea. C. Zamia amblyphyllidia. D. Zamia portoricensis. E. Alocasia 

macrorrhizos. F. Alocasia plumbea. G. Anthurium crenatum, habit. (A-E, G, unvouchered; F, from Acevedo-Rdgz. 13377). All 

photos by P. Acevedo-Rdgz.

386 


Fig. 59. A. Anthurium crenatum, infructescence B. Anthurium × selloum. C. Colocasia esculenta. D. Dieffenbachia seguine. E- 

F. Dracontium asperum. E. Habit. F. Inflorescence. G-H. Xanthosoma sagittifolium. G. Leaf. H. Inflorescence. I. Xanthosoma 

undipes. (A, from Acevedo-Rdgz. 2857; B, from Acevedo-Rdgz. 3810; C, unvouchered; D, from Acevedo-Rdgz. 13418; E-F, 

from Acevedo-Rdgz. 13402; G-H, from Acevedo-Rdgz. 13414; I, from Acevedo-Rdgz. 13413). All photos by P. Acevedo-Rdgz.


Fig. 60. A-B, D. Xanthosoma undipes. A. Detail of inflorescence. B. Cross section of petiole at invagination area. D. Base of 

petiole showing invagination. C. Xanthosoma violaceum, cross section of petiole at invagination area. E. Philodendron hederaceum. 

F. Syngonium podophyllum, inflorescence. G. Smilax domingensis, fruit. H. Smilax coriacea, inflorescence. I. Rajania cordata, 

leaf. (A, from Acevedo-Rdgz. 13413; B and D, from Acevedo-Rdgz. 13752; C, from Acevedo-Rdgz. 13753; E, from Acevedo- 

Rdgz. 2880; F, from Acevedo-Rdgz. 4061; G, from Acevedo-Rdgz. 6341; H, from Acevedo-Rdgz. 9330; I, unvouchered). All 

photos by P. Acevedo-Rdgz.

388 


Fig. 61. A. Curculigo capitulata, inflorescence. B. Trimezia steyermarkii, flower. C. Crocosmia × crocosmiiflora, inflorescence. 

D. Eucharis amazonica. E. Agave missionum. F. Hymenocallis latifolia, inflorescence. G. Agave sisalana. H. Crinum zeylanicum, 

inflorescence. (A-B, unvouchered; C, from Acevedo-Rdgz. 9424; D, from Acevedo-Rdgz. 9394; E, unvouchered; F, from Acevedo- 

Rdgz. 14051; G-H, unvouchered). All photos by P. Acevedo-Rdgz.


Fig. 62. A-B. Furcraea tuberosa. A. Habit. B. Leaf margin. C-D. Yucca aloifolia. C. Habit D. Flowers. E. Sansevieria hyacinthoides. 

F. Aloe vera. G. Sabal causiarum. H-I. Thrinax morrisii. H. habit. I. Inflorescence. (A-B, unvouchered; C-D, from Acevedo- 

Rdgz. 2008; E, from Acevedo-Rdgz. 3859; F-I, unvouchered). All photos by P. Acevedo-Rdgz.

390 


Fig. 63. A. Aploleia monandra, flower. B. Gibasis geniculata. C. Tripogandra serrulata, inflorescence. D. Xiphidium caeruleum. 

E. Eichhornia crassipes, inflorescence. F. Pontederia cordata. G. Musa group ABB “mafafo”. H. Musa group AAB “manzano”. 

I. Heliconia caribaea, inflorescence. (A, from Acevedo-Rdgz. 7911; B, from Acevedo-Rdgz. 13409; C, from Acevedo-Rdgz. 

10760; D-I, unvouchered). All photos by P. Acevedo-Rdgz.


Fig. 64. A. Alpinia zerumbet, inflorescence. B. Alpinia purpurata, inflorescence. C. Zingiber zerumbet, inflorescence. D. 

Hedychium coronarium, flowers. E. Renealmia jamaicensis var. puberula, inflorescence. F. Costus speciosus, inflorescence. G. 

Canna indica. H. Calathea lutea. (A-B, unvouchered; C, from Acevedo-Rdgz. 13419; D, unvouchered; E, from Acevedo-Rdgz. 

6224; F, unvouchered; G, from Acevedo-Rdgz. 10705; H, from Acevedo-Rdgz. 10767). All photos by P. Acevedo-Rdgz.

392 


Fig. 65. A. Bromelia pinguin, habit. B. Guzmania monostachia, habit. C. Guzmania berteroniana, habit. D. Tillandsia utriculata, 

habit. E. Werauhia sintenisii, habit. F. Cyperus papyrus, habit. G. Cladium jamaicense, habit. H. Rhynchospora ciliata (A, F, 

unvouchered; B, from Acevedo-Rdgz. 10739; C, from Acevedo-Rdgz. 10641; D, from Acevedo-Rdgz. 1971; E, from Acevedo- 

Rdgz. 10518). Photos A-F by P. Acevedo-Rdgz.; G & H by R.A. Howard.


spike terminal, the pistillate one below, contiguous or separated by a short interval from the staminate 

spike; perianth of bristles. Staminate flowers with 1-7 (usually 3) stamens with free or united filaments, 

the anthers linear (all staminate structures soon deciduous). Pistillate flowers with 1-locular ovary, 

elongate filiform style and linear to lance-ovate or spatulate stigma. Sterile flowers often present, these 

on long hairy stalks ending in a swollen, abortive ovary. Fruit a minute, 1-seeded achene. A cosmopolitan 

family of 10 species, characteristic of non-saline to brackish wetland sites in nearly all temperate and 

tropical regions. All the species are included in the single genus Typha. 

TYPE: Typha L. 

Typha Sp. Pl. 971. 1753. 

1. TYPHA 

With characteristics of the family. A single species occurs in Puerto Rico and the Virgin Islands. 

LECTOTYPE: Typha latifolia L., designated by P. Wilson in Britton, N. Amer. Fl. 17: 3. 1909. 

1. Typha domingensis Pers., Syn. Pl. 2: 532. 1807. 

Type: Hispaniola. Collector unknown 

(holotype: probably at P). 

Typha angustifolia sensu Britton & P. Wilson, 

1923, non Linnaeus, 1753. 

Fig. 57. A-F 

Stems 1.6-2.5 m tall, 7-14 (-16) mm wide at 

base, 3-6 mm wide at apex, the sheathing bases 

(13-) 15-30 mm wide, terete to subcompressed, 

smooth, finely ribbed. Leaves 6-10; sheaths 

elongate, abruptly narrowed into the leaf blade at 

summit or rarely with a short, rounded auricle, both 

sheaths and blades red-lineolate at adaxial junction 

of sheath and blade; blades plano-convex or 

flattened, linear, 0.8-2.5 m × 7-12 (-15) mm, 

glaucous, smooth, spongy-thickened, yellowish 

green to pale brown, glabrous. Inflorescences 

exceeding the leaves; spikes pale brown to 

cinnamon-colored; staminate spike 20-35 × 0.9- 

1.8 cm long, shortly separated from the pistillate 

spike, with very numerous clavellate hairs 

subtending the flowers and fruits; pistillate spike 

14-23 × 0.8-1.8 cm. Staminate flowers with 2-3 (- 

4) stamens, the filaments connate or sometimes 

distinct, the anthers 2-3.2 (-3.5) mm long. Sterile 

(abortive) pistils clavate, subcompressed, rounded 

at the apex, reticulate, 0.8-1.2 × 0.3-0.6 mm. 

Fertile achenes minute, narrowly elliptic. 


Mexico, Central America, Bermuda, Bahamas, 

Greater Antilles, Cayman Islands, Trinidad, and 

chiefly coastal South America. Apparently absent 

from the Lesser Antilles, despite reports. 


Islands: This species grows in marshy or wetland 

habitats from low to middle elevations (Sea level 

to 325 m or more); it is far more common and 

widespread than the vouchered records indicate, 

because specimens are often not easy to collect. 

Recorded from Arecibo, Cabo Rojo, Carolina, 

Cataño, Ceiba, Guánica, Humacao, San Juan, and 

Vieques; St. Croix, St. Thomas, and Tortola. 

Common name: Puerto Rico: Eneas, Yerba de 

eneas. 

Note: The species of Typha are considered 

very useful in many parts of the world. The fibrous 

stems and leaves are used for thatch, soft matting, 

ropes and baskets. The downy wool of the 

inflorescence can be applied to wounds like cotton. 

The young shoots are edible, tasting when cooked 

like asparagus. The pollen, which is produced in 

great abundance, can be used like flour for baking 

a type of bread; it is also highly inflammable. 


Cabo Rojo: Sintenis 688 (US). San Juan: Martín 

Peña, Stevenson 1232 (US). Guánica: Sintenis 

3936 (US).

394 


CULTIVATED FAMILIES 

ALLIACEAE 

The following taxa of Allium were listed by Britton & P. Wilson (1923) as cultivated and generally 

grown for food in Puerto Rico or the Virgin Islands: A. ascalonicum Bory & Chaub., of Syrian origin, 

was occasionally grown in the Virgin Islands; A. cepa L., a native of Persia, was sometimes grown for 

food in Puerto Rico; A. fistulosum L., from Siberia, was cultivated in the Virgin Islands; A. porrum L., 

European, was planted for food in Puerto Rico and Virgin Islands gardens; A. sativum L., European, was 

cultivated for food; and A. scorodoprasum L., of European origin, was cultivated in St. Thomas. 

COLCHICACEAE 

Gloriosa superba L., native of the Old World tropics, was listed by Britton & P. Wilson (1923) as 

cultivated on St. Croix. It has also been recorded for St. Thomas: Rose s.n. (US). 

CONVALLARIACEAE 

Aspidistra lurida Ker Gawl., of Chinese origin, was occasionally grown for ornament in Puerto 

Rico gardens (Britton & P. Wilson, 1923). 

CYCLANTHACEAE 

Carludovica palmata Ruiz & Pav., native of Peru, is occasionally planted for ornament in Puerto 

Rico (Britton & P. Wilson, 1923). 

LILIACEAE 

Chlorophytum comosum (Thunb.) Jacques (native to South Africa), Hemerocallis fulva (L.) L. (native 

to Europe), Lilium auratum Lindl. (native to Japan), L. lancifolium Thunb. (native to Japan), L. longiflorum 

Thunb. (native to Japan and China), and L. regale E. H. Wilson (native to China) were cited by Liogier 

& Martorell (1982) as cultivated in Puerto Rico. 

NOLINACEAE 

Dasylirion glaucophyllum Hook., a native of Mexico, was cultivated in St. Thomas at Louisenhoj 

(Britton & P. Wilson, 1923). 

PANDANACEAE 

The following species of Pandanus were cited by Liogier & Martorell (1982) as cultivated in Puerto 

Rico. Pandanus baptistii Misonne (native to New Britain Island), P. odoriferus (Forssk.) Chiov. (native 

to southeast Asia and the Philippines), P. pacificus Mast. (native to New Guinea, Moluccas and Marianas), 

P. tectorius Parkinson (native to southeast Asia, Australia, Philippines and Tahiti), P. utilis Bory (native 

to Madagascar, also planted in St. Croix and St. Thomas), and P. veitchii Mast. (native to Polynesia). 

STRELITZIACEAE 

Ravenala madagascariensis Sonn. and Strelitzia reginae Aiton, were cited by Martorell et al. (1981) 

as cultivated in Puerto Rico. 

TACCACEAE 

Tacca chantrieri André, native to Malesia, was cited by Liogier & Martorell (1982) as cultivated in 

Puerto Rico.


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Natl. Herb. 51: 1-483. 

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

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Adams, C. D. 1972. Flowering plants of Jamaica. 

University of the West Indies, Mona, Jamaica. 

848 pp. 

Bello y Espinosa, D. 1880-1883. Apuntes para la 

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Nat. 10: 231-304 (1880); 12: 103-130 (1883). 

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398 

A 

Abildgaardia Vahl 241 

Abildgaardia monostachyos (L.) Vahl 241 

Abildgaardia ovata (Burm.f.) Kral 241 

Acontias cubensis sensu Bello 49 

Acontias hastifolius Schott 49 

Acontias helleborifolius (Jacq.) Schott 49 

Acrista monticola O. F. Cook 144 

Acrocomia Mart. 137 

Acrocomia aculeata (Jacq.) Lodd. 137 

Acrocomia aculeata sensu Britton & P. Wilson 137 

Acrocomia media O. F. Cook 137 

Acrocomia sclerocarpa sensu Bello 137 

Actinophloeus macarthurii (H. Wendl.) Becc. 152 

Aechmea Ruiz & Pav. 201 

Aechmea fasciata (Lindl.) Baker 201 

Aechmea lingulata (L.) Baker 202 

Aechmea nudicaulis (L.) Griseb. 202 

Aechmea paniculata Ruiz & Pav. 201, 203 

Aechmea plumieri Baker 202 

Agathis robusta (F.J. Müll.) F.M. Bailey 17 

Agavaceae Endl. 116 

Agave L. 117 

Agave americana L. 117 

Agave beauleriana Jacobi 121 

Agave cubensis Jacq. 121 

Agave eggersiana Trel. 118 

Agave evadens Trel. 121 

Agave jacquiniana Schult. 121 

Agave minor Proctor 118 

Agave missionum Trel. 118 

Agave neglecta Small 121 

Agave portoricensis Trel. 118 

Agave sisalana Perrine 120 

Agave tuberosa Mill. 122 

Agave vivipara L. 120 

Aglaonema commutatum Schott 52 

Aglaonema pictum Kunth 52 

Aiphanes Willd. 138 

Aiphanes acanthophylla (Mart.) Burret 138 

Aiphanes aculeata Willd. 138 

Aiphanes minima (Gaertn.) Burret 139 

Aletris fragrans L. 130 

Alisma L. 58 

Alisma berteroi Spreng. 59 

Alisma bolivianum Rusby 59 

Alisma rostratum Nutt. 59 

Alisma tenellum Mart. 60 

Alisma tenellum f. latifolius Seub. 60 

Alismataceae Vent. 58 

Alliaceae 386 

Allium ascalonicum Bory & Chaub. 386 

Allium cepa L. 386 

Allium fistulosum L. 386 

Allium porrum L. 386 

Allium sativum L. 386 

Allium scorodoprasum L. 386 

Alocasia (Schott) G. Don 22 


INDEX TO SCIENTIFIC NAMES 

Alocasia cucullata (Lour.) G. Don 22 

Alocasia macrorrhizos (L.) G. Don 23 

Alocasia plumbea Van Houtte 23 

Alocasia undipes K. Koch 50 

Aloe L. 133 

Aloe arborescens Mill. 135 

A1oe barbadensis Mill. 134 

Aloe ciliaris Haw. 135 

Aloe ferox Mill. 135 

Aloe fragrantissima Jacq. 130 

Aloe grandidentata Salm-Dyck 135 

Aloe greenii Baker 135 

Aloe hyacinthoides L. 130 

Aloe hyacinthoides ß guineensis L. 131 

Aloe obscura Mill. 135 

A1oe perfoliata var. vera L. 134 

Aloe vera (L.) Burm. f. 134 

Aloe vulgaris Lam. 134 

Alpinia Roxb. 179 

Alpinia antillarum sensu Urban 184 

Alpinia antillarum var. puberula (Gagnep.) Moscoso 184 

Alpinia aromatica Aubl. 183 

Alpinia elatior Jack 180 

Alpinia exaltata (L. f.) Roem. & Schult. 182 

Alpinia galanga (L.) Willd. 179 

Alpinia occidentalis Sw. 183 

Alpinia purpurata (Vieill.) K. Schum. 180 

Alpinia sanderae Sander 180 

Alpinia speciosa (J.C. Wendl.) K. Schum. 179 

Alpinia spicata Jacq. 190 

Alpinia zerumbet (Pers.) B. L. Burtt & R. M. Sm. 179 

Amaryllidaceae J. St. Hil. 106 

Amaryllis L.106 

Amaryllis atamasco L. 113 

Amaryllis aurea Ruiz & Pav. 113 

Amaryllis belladonna L. 116 

Amaryllis equestris Aiton 109 

Amaryllis punicea Lam. 109 

Amaryllis reginae L. 109 

Amaryllis tubispatha sensu Ker Gawler 115 

Amaryllis zeylanica L. 108 

Amomum alpina Rottb. 182 

Amomum littorale J. König 180 

Amomum montanum J. König 185 

Amomum zerumbet L. 185 

Amomum zingiber L. 185 

Anacharis canadensis (Michx.) Planch. 64 

Anacharis densa (Planch.) Vict. 64 

Ananas Mill. 203 

Ananas ananas (L.) Voss 203 

Ananas comosus (L.) Merr. 203 

Ananas monstrosus (Carr.) L. B. Sm. 203 

Ananas sativus Schult. f. 203 

Ananassa monstrosa Carr. 203 

Aneilema geniculata (Jacq.) Woodson 161 

Angiosperms 63, 68 

Anthurium Schott 24 

Anthurium acaule (Jacq.) Schott 24 

Anthurium acaule sensu authors 25


Anthurium acaule var. brevipes Engl. 25 

Anthurium acaule var. portoricense Kuntze 25 

Anthurium andreanum Linden 29 

Anthurium cordatum (L.) Schott 24 

Anthurium cordifolium sensu Eggers 24 

Anthurium crenatum (L.) Kunth 25 

Anthurium dominicense Schott 26 

Anthurium dominicense var. sintenisii Engl. 26 

Anthurium huegelli sensu Bello 25 

Anthurium huegelli sensu Eggers 25 

Anthurium huegelli sensu Millspaugh 

Anthurium lanceolatum Kunth ex Bello 25 

Anthurium longilaminatum Engl. 29 

Anthurium scandens (Aubl.) Engl. 26 

Anthurium × selloum K. Koch 28 

Aploleia Raf. 154 

Aploleia monandra (Sw.) H. E. Moore 154 

Apteria Nutt. 93 

Apteria aphylla (Nutt.) Barnhart 93 

Apteria aphylla var. hymenanthera (Miq.) Jonker 93 

Apteria hymenanthera Miq. 93 

Apteria setacea Nutt. 93 

Araceae Juss. 21 

Araucaria angustifolia (Bertol.) Kuntze 17 

Araucaria bidwillii Hook. 17 

Araucaria heterophylla (Salisb.) Franco 17 

Archontophoenix cunninghamiana (H. Wendl.)H. Wendl. & 

Drude 153 

Areca Linneaus 136 

Areca catechu L. 152 

Areca madagascariensis Mart. 152 

Arecastrum romanzoffianum (Cham.) Becc. 153 

Arecaceae Schultz Sch. 135 

Arenga pinnata (Wurmb) Merr. 153 

Arenga saccharifera Labill. 152 

Arikuryroba schizophylla (Mart.) L. H. Bailey 153 

Arum L. 21 

Arum auritum L. 46 

Arum bicolor Aiton 29 

Arum colocasia L. 30 

Arum cucullatum Lour. 22 

Arum esculentum L. 30 

Arum grandifolium Jacq. 38 

Arum hederaceum Jacq. 40 

Arum helleborifolium Jacq. 49 

Arum lingulatum L. 42 

Arum macrorrhizon L. 23 

Arum nigrum Vell. 51 

Arum sagittifolium L. 48, 50 

Arum seguine Jacq. 31 

Asparagaceae Juss. 125 

Asparagopsis Kunth 126 

Asparagopsis setaceus Kunth 127 

Asparagus L. 126 

Asparagus aethiopicus L. 126 

Asparagus densiflorus sensu authors 126 

Asparagus draco L. 129 

Asparagus officinalis L. 128 

Asparagus plumosus Baker 128 

Asparagus setaceus (Kunth) Jessop 127 

Asparagus sprengeri sensu authors 126 

Asparagus terminalis L. 129 

Asphodelaceae Juss. 133 

Asphodelus L. 133 

Aspidistra lurida Ker Gawl. 386 

Astrocaryum standleyanum L. H. Bailey 153 

Atamosco carinata (Herb.) P. Wilson 113 

Atamosco rosea (Lindl.) Green 115 

Atamosco tubispatha sensu Britton & P. Wilson 115 

Athyrocarpus persicariifolius sensu Britton & P. Wilson 161 

B 

Bactris acanthophylla Mart. 138 

Bactris gasipaes Kunth 153 

Bactris pavoniana sensu Bello 139 

Banksia speciosa J. König 190 

Belamcanda Adans. 101 

Belamcanda chinensis (L.) A. DC. 101 

Belamcanda punctata Moench. 101 

Bihai bihai of Britton & P. Wilson 175 

Bihai bihai (L.) Griggs 175 

Bihai borinquena (Griggs) Griggs 175 

Bihai rostrata (Ruiz & Pav.) Griggs 177 

Billbergia fasciata Lindl. 201 

Billbergia pyramidalis (Sims.) Lindl. 231 

Borassus flabellifer L. 152 

Bromelia L. 204 

Bromelia ananas L. 203 

Bromelia comosa L. 203 

Bromelia karatas L. 204, 206 

Bromelia lingulata L. 202 

Bromelia nudicaulis L. 202 

Bromelia pinguin L. 204 

Bromelia plumieri (Morren) L. B. Sm. 206 

Bromeliaceae Juss. 199 

Bulbostylis Kunth 242 

Bulbostylis antillana (Britton) Fernald 243 

Bulbostylis capillaris (L.) C. B. Clarke 242 

Bulbostylis capillaris subsp. antillana (Britton) 

T. Koyama 243 

Bulbostylis capillaris subsp. insulana M.T. Strong 243 

Bulbostylis curassavica (Britton) Kük. ex Urb. 244 

Bulbostylis curassavica var. pallescens Kük. & Ekman ex 

Urb. 244 

Bulbostylis ekmanii Kük. 245 

Bulbostylis floccosa var. β “(?)” pumilo C. B. Clarke 244 

Bulbostylis hirtella sensu Urban 248 

Bulbostylis junciformis (Kunth) C. B. Clarke 248 

Bulbostylis langsdorffiana (Kunth) C.B. Clarke 248 

Bulbostylis papillosa Kük. 245 

Bulbostylis pauciflora (Liebm.) C. B. Clarke 245 

Bulbostylis portoricensis (Britton) Fernald 245 

Bulbostylis stenocarpa Kük. 245, 247 

Bulbostylis subaphylla C. B. Clarke 247 

Bulbostylis vestita (Kunth) C. B. Clarke 247, 248 

Burmannia L. 93 

Burmannia capitata (Walter ex J. F. Gmel.) Mart. 94 

Burmannia disticha L. 93 

Burmanniaceae Blume 92 

Butia capitata (Mart.) Becc. 152 

C 

Caladium Vent. 29 

Caladium bicolor (Aiton) Vent. 29 

Caladium brasiliense Desf. 49 

Caladium colocasia (L.) W. Wight 30

400 


Caladium maculatum Lodd. 31 

Calathea G. Mey. 195 

Calathea allouia (Aubl.) Lindl. 195 

Calathea crotalifera S. Watson 196 

Calathea discolor G. Mey. 195 

Calathea insignis Petersen 196 

Calathea lutea (Aubl.) Schult. 196 

Calathea ornata (Lindl.) Körn. 197 

Calathea quadratispica Woodson 196 

Calathea zebrina (Sims) Lindl. 197 

Calla palustris L. 52 

Callisia Loefl. 156 

Callisia fragrans (Lindl.)Woodson 156 

Callisia monandra (Sw.) Schult. 154 

Callisia repens (Jacq.) L. 156 

Callisia umbellulata Lam. 154 

Calyptrogyne occidentalis sensu Britton & P. Wilson 140 

Calyptrogyne quisqueyana (L. H. Bailey) León 140 

Calyptrogyne rivalis (O. F. Cook) León 140 

Calyptronoma Griseb. 139 

Calyptronoma occidentalis (Sw.) H. E. Moore 140 

Calyptronoma quisqueyana L. H. Bailey 140 

Calyptronoma rivalis (O. F. Cook) L. H. Bailey 140 

Campelia zanonia (L.) Kunth 165 

Canna L. 191 

Canna coccinea sensu Britton & P. Wilson 192 

Canna edulis Ker Gawl. 194 

Canna glauca L. 192 

Canna indica L. 191, 192 

Canna jaegeriana Urb. 194 

Canna lambertii Lindl. 192 

Canna lutea Mill. 194 

Canna pertusa Urb. 194 

Canna portoricensis Bouché 192 

Canna sylvestris Roscoe 192 

Cannaceae Juss. 191 

Caraguata berteroniana Schult. f. 208 

Caraguata grandiflora Baker 208 

Caraguata macrostachya Bello 228 

Caraguata sintenisii Baker 230 

Carex L. 248 

Carex cladostachya Wahlenb. 250 

Carex cladostachya f. polystachya (Sw. ex Wahlenb.) C. B. 

Clarke ex Lindm. 250 

Carex dussiana Boeck. 250 

Carex hirta L. 249 

Carex longii Mack. 250 

Carex ovata Burm.f. 241 

Carex polystachya Sw. ex Wahlenb. 250 

Carex pulicaris L. 249 

Carludovica palmata Ruiz & Pav. 386 

Caryota mitis Lour. 152 

Caryota urens L. 152 

Catopsis Griseb. 206 

Catopsis floribunda L. B. Sm. 206 

Catopsis nitida (Hook.) Griseb. 207 

Catopsis nitida (Hook.) Griseb. 206 

Catopsis nitida sensu Duss 206 

Catopsis nutans sensu Britton & P. Wilson 206 

Catopsis sessiliflora (Ruiz & Pav.) Mez 207 

Caulinia guadalupensis Spreng. 68 

Caulinia ovalis R. Br. 65 

Chamaecyparis funebris (Endl.) Franco 17 

Chamaedorea elegans Mart. 153 

Chevalliera lingulata (L.) Griseb. 202 

Chrysalidocarpus madagascariensis Becc. 153 

Corypha umbraculifera L. 153 

Cryosophila warscewiczii (H. Wendl.) Bartlett 153 

Chlorophytum comosum (Thunb.) Jacques 386 

Chrysalidocarpus lutescens H. Wendl. 152 

Cipura plicata (Sw.) Griseb. 104 

Cladium P. Browne 251 

Cladium jamaicense Crantz 251 

Cladium mariscus subsp. jamaicense (Crantz) Kük. 251 

Cladium occidentale sensu Grisebach 251 

Cladium restioides (Sw.) Benth. ex C. B. Clarke 327 

Coccothrinax Sarg. 140 

Coccothrinax alta (O. F. Cook) Becc. 141 

Coccothrinax argentea (Lodd.) Sarg. 141 

Coccothrinax argentea sensu Britton & P. Wilson 141 

Coccothrinax barbadensis (Lodd. ex Mart.) Becc. 141 

Coccothrinax eggersiana Becc. 141 

Coccothrinax eggersiana var. sanctae-crucis Becc. 141 

Coccothrinax jucunda Sarg. 141 

Coccothrinax latifrons (O. F. Cook) Becc. 141 

Coccothrinax sanctae-thomae Becc. 141 

Cocos L. 142 

Cocos aculeata Jacq. 137 

Cocos nucifera L. 142 

Cocos plumosa Lodd. ex Hook. 142 

Cocops rivalis O. F. Cook 140 

Colchicaceae 386 

Colocasia Schott 30 

Colocasia antiquorum Schott 30 

Colocasia esculenta (L.) Schott 30 

Commelina L. 157 

Commelina benghalensis L. 158 

Commelina cayennensis Rich. 158 

Commelina communis L. 158 

Commelina diffusa Burm. f. 158 

Commelina elegans Kunth 159 

Commelina erecta L. 159 

Commelina longicaulis Jacq. 158 

Commelina nudiflora L. 162 

Commelina rufipes var. glabrata (D. R. Hunt) Faden & D. 

R. Hunt 159 

Commelina scapiflora Roxb. 162 

Commelina serrulata Vahl 166 

Commelina virginica sensu authors 159 

Commelina zanonia L. 165 

Commelinaceae R. Br. 153 

Commelinopsis glabrata D. R. Hunt 159 

Commelinopsis persicariifolia Delile 161 

Convallaria fruticosa L. 129 

Convallariaceae 386 

Cordyline Comm. ex Juss. 129 

Cordyline cannifolia R. Br. 129 

Cordyline cylindrica (Bojer ex Hook.) Britton 131 

Cordyline fruticosa (L.) A. Chev. 129 

Cordyline guineensis (L.) Britton 131 

Cordyline terminalis (L.) Kunth 128 

Corypha elata Roxb. 152 

Corypha minor Jacq. 149 

Corypha umbraculifera L. 153 

Costaceae Nakai 186 

Costus L. 187 

Costus arabicus L. 187 

Costus cylindricus Jacq. 190


Costus glabratus sensu Bello 185 

Costus guanaiensis var. macrostrobilus (K. Schum.) 

Maas 187 

Costus macrostrobilus K. Schum. 187 

Costus malortieanus H. Wendl. 188 

Costus scaber Ruiz & Pav. 188 

Costus speciosus (J. König) Sm. 190 

Costus spicatus (Jacq.) Sw. 190 

Costus spicatus var. [β] pubescens Griseb. 190 

Costus zerumbet Pers. 179 

Crinum L. 107 

Crinum amabile Donn 107 

Crinum amabile Donn ex Ker-Gawl. 107 

Crinum americanum L. 108 

Crinum asiaticum L. 107 

Crinum erubescens Aiton 108 

Crinum giganteum Andr. 108 

Crinum latifolium var. zeylanicum (L.) Hook. f. 108 

Crinum zeylanicum (L.) L. 108 

Crocosmia Planch. 102 

Crocosmia aurea Planch. 102 

Crocosmia × crocosmiiflora (Lemoine ex Morren) N. E. 

Br. 102 

Crocosmia pottsii (Baker) N. E. Br. 102 

Cupressus arizonica Greene 17 

Cupressus lusitanica Mill. 17 

Cupressus marcrocarpa Hartw. 17 

Cupressus sempervirens L. 17 

Curculigo Gaertn. 97 

Curculigo capitulata (Lour.) Kuntze. 97 

Curculigo orchioides Gaertn. 97 

Curculigo recurvata Dryand. 97 

Curculigo scorzonerifolia (Lam.) Baker 97 

Curcuma longa L. 186 

Curima colophylla O. F. Cook 139 

Cyclanthaceae 386 

Cymodocea C. Koenig 71 

Cymodocea manatorum Asch. 73 

Cymodoceaceae N. Taylor 71 

Cyperaceae Juss. 236 

Cyperus L. 253 

Cyperus acicularis (Schrad. ex Nees) Steud. 180 

Cyperus aggregatus (Willd.) Endl. 256 

Cyperus alternifolius of many authors 273 

Cyperus alternifolius subsp. flabelliformis (Rottb.) 

Kük. 273 

Cyperus angulatus Nees 292 

Cyperus aristatus Rottb. 287 

Cyperus articulatus L. 259 

Cyperus balbisii Kunth 286 

Cyperus bipartitus Torr. 261 

Cyperus brevifolius (Rottb.) Hassk. 318 

Cyperus brizaeus Vahl 260 

Cyperus bromoides Willd. ex Link 292 

Cyperus brunneus Sw. 260 

Cyperus brunneus sensu Grisebach 282 

Cyperus calcicola Britton 291 

Cyperus capillaris J. König ex Roxb. 277 

Cyperus caribaeus Pers. 241 

Cyperus cayennensis (Lam.) Britton 258 

Cyperus comosus (Kunth) Poir. 271 

Cyperus complanatus (Retz.) Willd. 308 

Cyperus compresso-triqueter Boeck. 276 

Cyperus compressus L. 261 

Cyperus compressus var. capillaceus C. B. Clarke 261 

Cyperus confertus Sw. 262, 284 

Cyperus confertus sensu Grisebach 266 

Cyperus coriifolius Boeck. 294 

Cyperus croceus Vahl 262 

Cyperus cubanus Liebm. 280 

Cyperus cubensis Steud. 267 

Cyperus cuspidatus Kunth 294 

Cyperus cuspidatus sensu Britton & P. Wilson 287 

Cyperus densicaespitosus Mattf. & Kük. ex Kük. 321 

Cyperus depauperatus Vahl 305 

Cyperus difformis L. 263 

Cyperus digitatus Roxb. 264 

Cyperus discolor Boeck. 260 

Cyperus distans L.f. 264 

Cyperus eggersii Boeck. 265 

Cyperus ehrenbergii Kunth 269 

Cyperus elatus Rottb. 264 

Cyperus elegans L. 266 

Cyperus esculentus L. 253, 267 

Cyperus esculentus var. macrostachyus Boeck. 267 

Cyperus evaginatus Boeck. 260 

Cyperus fastuosus Ham. 280 

Cyperus ferax Rich. 280 

Cyperus ferax var. acicularis (Schrad. ex Nees) Kük. 280 

Cyperus ferrugineus Poir. 294 

Cyperus filicinus Vahl 294 

Cyperus filiformis Sw. 268, 289 

Cyperus filiformis var. densiceps Kük. 269 

Cyperus flabelliformis Rottb. 273 

Cyperus flavescens L. 268 

Cyperus flavus (Vahl) Nees 258 

Cypreus flavus var. aggregatus (Willd.) Kük. 258 

Cyperus flexuosus Vahl 269 

Cyperus flexuosus sensu Grisebach 280 

Cyperus flexuosus var. insignis (Kunth) Kük. 269 

Cyperus floridanus Britton ex Small 269 

Cyperus formosus Vahl 293 

Cyperus fuligineus Chapm. 270 

Cyperus giganteus Vahl 271 

Cyperus giganteus var. comosus (Kunth) Kük. 271 

Cyperus globulosus sensu authors 262 

Cyperus gracilis R. Br. 290 

Cyperus granularis (Boeck.) Britton 290 

Cyperus hahnianus Boeck. 283 

Cyperus hartii Boeck. 290 

Cyperus haspan L. 271 

Cyperus haspan subsp. juncoides (Lam.) Kük. 271 

Cyperus hermaphroditus (Jacq.) Standl. 294 

Cyperus humilis Kunth 294 

Cyperus imbricatus Retz. 272 

Cyperus imbricatus var. elongatus (Boeck.) Kük. 272 

Cyperus inflexus Muhl. 287 

Cyperus insignis Kunth 269 

Cyperus involucratus Rottb. 273 

Cyperus iria L. 274 

Cyperus juncoides Lam. 271 

Cyperus kingsii C.D. Adams ex Proctor 270 

Cyperus krugii Boeck. 260 

Cyperus kyllinga Endl. 319 

Cyperus ligularis L. 275 

Cyperus laevigatus L. 274 

Cyperus lutescens Torr. & Hook. 267 

Cyperus luzulae (L.) Rottb. ex Retz. 294

402 


Cyperus martinicensis Boeck. 276 

Cyperus michauxianus Schult. 280 

Cyperus monostachyos L. 241 

Cyperus mutisii (Kunth) Andersson 276 

Cyperus mutisii var. martinicensis (Boeck.) Kük. 276 

Cyperus mutisii var. semitribrachiatus (Böck) Kük. 276 

Cyperus nanus Willd. 277, 288 

Cyperus nanus var. subtenuis Kük. 288 

Cyperus obtusatus (J. Presl & C. Presl) Mattf. & Kük. 322 

Cyperus ochraceus Vahl 279 

Cyperus ochraceus var. excelsior Kük. 279 

Cyperus ochraceus var. minor Kük. 279 

Cyperus ochreatus Boeck. 276 

Cyperus odoratus L. 266, 280 

Cyperus odoratus sensu Britton & P. Wilson 283 

Cyperus ottonis Boeck. 260 

Cyperus papyrus L. 281 

Cyperus parvispiculatus Boeck. 280 

Cyperus pedunculatus (R. Br.) J. Kern 327 

Cyperus peruvianus F. N. Williams 322 

Cyperus peruvianus var. foliatus (Kük.) Kük. 321 

Cyperus planifolius Rich. 269, 282 

Cyperus planifolius var. brunneus (Sw.) Kük. 260 

Cyperus platystachyus Griseb. 290 

Cyperus poeoides Ham. 280 

Cyperus polystachyos Rottb. 283 

Cyperus polystachyos var. polystachyos 284 

Cyperus pseudo-bromoides Boeck. 292 

Cyperus pseudo-variegatus Boeck. 284 

Cyperus pulguerensis M. T . Strong 284 

Cyperus purpurascens Vahl 282 

Cyperus radiatus Vahl 272 

Cyperus radiatus var. elongatus Boeck. 272 

Cyperus retroflexus Buckley 294 

Cyperus rotundus L. 267, 284 

Cyperus sanctae-crucis Liebm. 280 

Cyperus semitribrachiatus Boeck. 276 

Cyperus sesquiflorus (Torr.) Mattf. & Kük. 319 

Cyperus sintenisii Boeck. 275 

Cyperus sphacelatus Rottb. 286 

Cyperus squarrosus L. 287, 294 

Cyperus subtenuis (Kük.) M.T. Strong 288 

Cyperus surinamensis Rottb. 289, 294 

Cyperus surinamensis var. formosus (Vahl) Kük. 293 

Cyperus swartzii Boeck. 289 

Cyperus swartzii var. elongatus Kük. 290 

Cyperus swartzii var. granularis (Desf. ex Boeck.) 

Kük. 290 

Cyperus tenuis Sw. 290 

Cyperus tenuis sensu Grisebach 288 

Cyperus tenuis var. capillaris (Sw.) Kük. 277 

Cyperus trigonus Boeck. 275 

Cyperus unifolius Boeck. 291 

Cyperus unioloides R. Br. 292 

Cyperus urbanii Boeck. 292 

Cyperus vahlii (Schrad. ex Nees) Steud. 269 

Cyperus virens Michx. 293 

Cyperus viscosus Sw. 266 

Cyrtanthus uniflorus Ker Gawl. 116 

Cyrtospadix bicolor (Aiton) Britton & P. Wilson 29 

D 

Dasylirion glaucophyllum Hook. 366 

Dendropogon usneoides (L.) Raf. 225 

Dichromena ciliata Vahl 337 

Dichromena colorata (L.) Hitchc. 340 

Dichromena contracta (Nees) Steud. 340 

Dichromena filiformis C. B. Clarke 335 

Dichromena filiformis (Vahl) Kunth 346 

Dichromena hispidula (Vahl) Kunth 343 

Dichromena leucocephala Michx. 340 

Dichromena microcephala Bertero ex Spreng. 356 

Dichromena persooniana Nees 364 

Dichromena pura Nees 337 

Dichromena pusilla (Sw.) Kunth 335 

Dichromena radicans sensu Urban 356 

Dichromena radicans sensu Britton & P. Wilson 356 

Dichromena velutina Kunth 362 

Diclidium aciculare Schrad. ex Nees 280 

Diclidium vahlii Schrad. ex Nees 269 

Dicotyledons 11 

Dictyosperma album (Bory) H. Wendl. & Drude ex 

Scheff. 153 

Dictyosperma rubrum H. Wendl. & Drude 152 

Dieffenbachia Schott 31 

Dieffenbachia maculata (Lodd.) G. Don 31 

Dieffenbachia plumieri Schott 31 

Dieffenbachia seguine (Jacq.) Schott 31 

Dioscorea L. 82 

Dioscorea alata L. 83 

Dioscorea altissima Lam. 85 

Dioscorea bulbifera L. 85 

Dioscorea cayennensis Lam. 86 

Dioscorea floribunda M. Martens & Galeotti 88 

Dioscorea friedrichsthalli R. Knuth 88 

Dioscorea latifolia Benth. 88 

Dioscorea pilosiuscula Bertero ex Spreng. 86 

Dioscorea polygonoides Humb. & Bonpl. ex Willd. 86 

Dioscorea rotundata Poir. 88 

Dioscorea sativa L. 83 

Dioscorea trifida L. f. 88 

Dioscoreaceae R. Br. 82 

Diplanthera beaudettei Hartog 72 

Diplanthera tridentata Steinh. 72 

Diplanthera wrightii (Asch.) Asch. 72 

Dracaena L. 129 

Dracaena draco (L.) L. 129, 130 

Dracaena fragrans (L.) Ker Gawl. 130 

Dracaena goldieana Bullen ex Mast. & T. Moore 130 

Dracaena surculosa Lindl. 130 

Dracaena terminalis (L.) L. 129 

Dracaenaceae Salisb. 128 

Dracontium L. 33 

Dracontium asperum K. Koch 33 

Dracontium lancifolium Jacq. 46 

Dracontium pertusum L. 36 

Dracontium polyphyllum L. 33 

Dracontium polyphyllum sensu authors 33 

Dracontium scandens Aubl. 26, 27 

Dyckia sulphurea K. Koch. 231 

Dypsis lutescens (H. Wendl.) Beentje & J. Dransf. 152 

E 

Echinodorus Rich. ex Engelm. 59 

Echinodorus berteroi (Spreng.) Fassett 59 

Echinodorus bolivianus (Rusby) Holm-Niels. 59


Echinodorus cordifolius sensu Bello 59 

Echinodorus latifolius (Seub.) Rataj 60 

Echinodorus rostratus (Nutt.) Engelm. 59 

Echinodorus tenellus var. latifolius (Seub.) Fassett 60 

Egeria densa Planch. 64, 65 

Eichhornia Kunth 170 

Eichhornia azurea (Sw.) Kunth 170, 171 

Eichhornia crassipes (Mart.) Solms 170 

Eichhornia diversifolia (Vahl) Urb. 171 

Elaeis guineensis Jacq. 152 

Elaeis occidentalis Sw. 140 

Elaeis oleifera (Kunth) Cortés ex Prain 

Eleocharis R. Br. 294 

Eleocharis acicularis var. lindheimeri C. B. Clarke ex 

Britton 304 

Eleocharis acicularis var. radicans (A. Dietr.) Britton 304 

Eleocharis albivaginata var. flaccida (Rchb. ex Spreng. f.) 

Boeck.99 

Eleocharis albivaginata var. humilis Boeck. 299 

Eleocharis ambigens Fernald 298 

Eleocharis atropurpurea (Retz.) J. Presl & C. Presl 306 

Eleocharis atropurpurea sensu Urban 297 

Eleocharis bahamensis Boeck. 300, 306 

Eleocharis capitata R. Br. 300 

Eleocharis caribaea (Rottb.) S.F. Blake 300 

Eleocharis cellulosa Torr. 296 

Eleocharis chaetaria sensu C. B. Clarke 305 

Eleocharis confervoides (Poir.) G. C. Tucker 304, 306 

Eleocharis constricta Schult. 297 

Eleocharis debilis Kunth 297 

Eleocharis depauperata (Vahl) Kunth 305 

Eleocharis elegans (Kunth) Roem. & Schult. 297 

Eleocharis fallax Weath. 298 

Eleocharis flaccida (Rchb. ex Spreng. f.) Urb. 299 

Eleocharis flavescens (Poir.) Urb. 299 

Eleocharis geniculata (L.) Roem. & Schult. 300, 306 

Eleocharis geniculata sensu Britton & P. Wilson 297 

Eleocharis geniculata sensu C. B. Clarke 297 

Eleocharis geniculata var. minor (Vahl) Roem. & 

Schult. 300 

Eleocharis interstincta (Vahl) Roem. & Schult. 301 

Eleocharis lindheimeri (C. B. Clarke ex Britton) 

Svenson 304 

Eleocharis microcarpa sensu Britton & P. Wilson 301 

Eleocharis minima sensu C. B. Clarke 301 

Eleocharis minutiflora Boeck. 301, 302 

Eleocharis montana (Kunth) Roem. & Schult. 302 

Eleocharis mutata (L.) Roem. & Schult. 302 

Eleocharis nervosa Kük. 298 

Eleocharis nigrescens var. minutiflora (Boeck.) 

Svenson 301 

Eleocharis nodulosa (Roth) Schult. 302 

Eleocharis ocreata (Nees) Steud. 299 

Eleocharis ocreata var. flaccida (Rchb. ex Spreng. f.) 

Boeck. ex C. B. Clarke 299 

Eleocharis ocreata var. humilis (Boeck.) C. B. Clarke 299 

Eleocharis oligantha C. B. Clarke 303, 306 

Eleocharis pachycarpa E. Desv. 306 

Eleocharis pachystyla (C. Wright) C. B. Clarke 304 

Eleocharis palustris (L.) Roem. & Schult. 295 

Eleocharis radicans (A. Dietr.) Kunth 304 

Eleocharis retroflexa (Poir.) Urb. 305 

Eleocharis rostellata (Torr.) Torr. 305 

Eleocharis shaferi Britton 297 

Eleocharis sintenisii Boeck. 300 

Eleocharis sintenisii sensu authors 297 

Eleocharis yunquensis Britton 297 

Eleogenus ocreatus Nees 299 

Eleogiton radicans A. Dietr. 304 

Eleutherine anomala Herb. 104 

Eleutherine bulbosa (Mill.) Urb. 104 

Eleutherine Herb. 102 

Eleutherine plicata (Sw.) Bold. 104 

Elodea Michx. 64 

Elodea canadensis Michx. 64 

Elodea densa (Planch.) Casp. 64 

Epipremnum Schott 34 

Epipremnum aureum (André) G. S. Bunting 34 

Epipremnum mirabile Schott 34 

Epipremnum pinnatum (L.) Engl. 34 

Epipremnum pinnatum ‘Aureum’ Nicolson 34 

Etlingera Giseke 180 

Etlingera elatior (Jack) R. M. Sm. 180 

Eucharis Planch. & Linden 108 

Eucharis amazonica Linden ex Planch. 108, 109 

Eucharis candida Planch. & Linden 108 

Eucharis grandiflora sensu authors 109 

Euterpe globosa sensu authors 144 

Euterpe montana Graham 144 

Euterpe oleracea sensu Boldingh 144 

F 

Fimbristylis Vahl 307 

Fimbristylis annua (All.) Roem. & Schult. 309 

Fimbristylis annua var. diphylla (Retz.) Kük. 309 

Fimbristylis autumnalis var. complanata (Retz.) Kük. 308 

Fimbristylis castanea of authors 314 

Fimbristylis complanata (Retz.) Link 308 

Fimbristylis curassavica (Britton) Alain 244 

Fimbristylis cymosa R. Br. 308 

Fimbristylis cymosa subsp. spathacea (Roth) T. 

Koyama 308 

Fimbristylis dichotoma (L.) Vahl 307, 309 

Fimbristylis diphylla (Retz.) Vahl 309 

Fimbristylis divaricata Ham. 343, 344 

Fimbristylis ferruginea (L.) Vahl 311 

Fimbristylis ferruginea var. compacta Kük. 311 

Fimbristylis glomerata Urb. 308 

Fimbristylis harrisii (Britton) C.D. Adams 247 

Fimbristylis hirsuta Hochst. ex Steud. 343 

Fimbristylis hirtella Ham. 344 

Fimbristylis hirtella Vahl 344 

Fimbristylis inaguensis Britton 311 

Fimbristylis littoralis Gaudich. 312 

Fimbristylis miliacea (L.) Vahl 312, 313 

Fimbristylis monostachyos (L.) Hassk. 241 

Fimbristylis obtusifolia sensu C. B. Clarke 309 

Fimbristylis ovata (Burm.f.) J. Kern 241 

Fimbristylis papillosa (Kük.) Alain 245 

Fimbristylis portoricensis (Britton) Alain 245 

Fimbristylis quinquangularis (Vahl) Kunth 313 

Fimbristylis sintenisii Boeck. 308 

Fimbristylis spadicea (L.) Vahl 314 

Fimbristylis spathacea Roth 308 

Fimbristylis vestita (Kunth) Hemsl. 247 

Fimbristylis villosa (Poir.) Roem. & Schult. 309 

Fuirena Rottb. 314

404 

Fuirena robusta Kunth 315 

Fuirena simplex Vahl 317 

Fuirena squarrosa Michx. 316 

Fuirena umbellata Rottb. 315, 316 

Furcraea Vent.121 

Furcraea andina Trel. 124 

Furcraea antillana A. Alvarez 121 

Furcraea cubensis (Jacq.) Vent. 121 

Furcraea elegans Todaro 124 

Furcraea foetida (L.) Haw. 124 

Furcraea hexapetala (Jacq.) Urb. 121 

Furcraea hexapetala sensu Urban 122 

Furcraea macrophylla Baker 124 

Furcraea selloa K. Koch cv. “marginata” 124 

Furcraea stratiotes J. B. Petersen 122 

Furcraea tuberosa (Mill.) W.T. Aiton 122 

G 


Gaussia H. Wendl. 142 

Gaussia attenuata (O. F. Cook) Becc. 143 

Gaussia portoricensis Hemsl. 143 

Gaussia princeps H. Wendl. 143 

Galatea bulbosa (Mill.) Britton 104 

Gibasis Raf. 161 

Gibasis geniculata (Jacq.) Rohweder 161 

Gibasis pulchella (Kunth) Raf. 161 

Gladiolus communis L. 106 

Gladiolus cuspidatus Jacq. 106 

Gladiolus x gandavensis Van Houtte 106 

Gloriosa superba L. 386 

Grantia brasiliensis (Wedd.) Mac Mill. 56 

Guillainia purpurata Vieill. 180 

Guzmania Ruiz & Pav. 208 

Guzmania berteroniana (Schult. f.) Mez in C. DC. 208, 211 

Guzmania erythrolepis Brongn. ex Planch. 211 

Guzmania erythrolepis sensu Britton & P. Wilson 208 

Guzmania lingulata (L.) Mez 209 

Guzmania lingulata var. concolor Proctor & Cedeño- 

Mald. 209 

Guzmania lingulata var. splendens (Planch.) Mez 209 

Guzmania monostachia (L.) Rusby ex Mez 209 

Guzmania sintenisii (Baker) Mez 230 

Guzmania tricolor Ruiz & Pav. 208, 210 

Gymnosiphon Blume 95 

Gymnosiphon aphyllus Blume 95 

Gymnosiphon germaini Urb. 95 

Gymnosiphon niveus (Griseb.) Urb. 95 

Gymnosiphon portoricensis Urb. 95, 96 

Gymnosiphon sphaerocarpus Urb. 96 

Gymnosperms 11 

H 

Haemodoraceae R. Br. 267 

Haemodorum Sm. 267 

Halodule Endl. 72 

Halodule beaudettei (Hartog) Hartog 72 

Halodule tridentata (Steinh.) Endl. ex Unger 72 

Halodule wrightii Asch. 72 

Halophila Thouars 65 

Halophila aschersonii Ostenf. 65, 67 

Halophila baillonis Asch. 65 

Halophila baillonis sensu Britton & P. Wilson 66 

Halophila decipiens Ostenf. 65 

Halophila engelmannii Asch. 65, 66 

Halophila ovalis (R. Br.) Hook. 65 

Haloschoenus hispidulus (Vahl) Nees 343 

Hapalanthus repens Jacq. 165 

Haworthia tessellata Haw. 135 

Hedychium J. König 181 

Hedychium coronarium J. König 181 

Hedychium flavescens Carey ex Roscoe 182 

Hedychium flavum Roxb. 181 

Helianthium tenellum sensu Britton & P. Wilson 60 

Heliconia L. 174 

Heliconia acuminata Rich. 177 

Heliconia bihai (L.) L. 174, 175 

Heliconia borinquena Griggs 175 

Heliconia caribaea Lam. 175 

Heliconia humilis of authors 177 

Heliconia latispatha Benth. 178 

Heliconia marginata (Griggs) Pittier 178 

Heliconia psittacorum L. f. 175 

Heliconia platystachys Baker 178 

Heliconia psittacorum H spathocircinata cv. “Golden 

Torch” 177 

Heliconia rostrata Ruiz & Pav. 177 

Heliconia stricta Huber 177 

Heliconia subulata Ruiz & Pav. 178 

Heliconiaceae Nakai 174 

Hemerocallis fulva (L.) L. 386 

Hemicarpha micrantha (Vahl) Pax 324 

Hemicarpha subsquarrosa (Muhl.) Nees 324 

Heteranthera Ruiz & Pav. 171 

Heteranthera diversifolia Vahl 171 

Heteranthera limosa (Sw.) Willd. 172 

Heteranthera reniformis Ruiz & Pav. 171,, 172 

Hippeastrum Herb. 109 

Hippeastrum equestre Herb. 109 

Hippeastrum puniceum (Lam.) Kuntze 109 

Hippeastrum reginae (L.) Herb. 109 

Hohenbergia Schult. & Schult. f. 211 

Hohenbergia antillana Mez 212 

Hohenbergia attenuata Britton 212 

Hohenbergia portoricensis Mez 212 

Hohenberiga stellata Schult. f. 211 

Hohenbergia tetaensis Proctor & Cedeño-Mald. 212 

Howea belmoreana (C. Moore & F. Muell.) Becc. 252 

Hydrocharis L. 63 

Hydrocharis spongia Bosc 67 

Hydrocharitaceae Juss. 63 

Hydrocleys Rich. 57 

Hydrocleys commersonii Rich. 58 

Hydrocleys nymphoides (Willd.) Buchenau 58 

Hydromystria laevigata (Humb. & Bonpl. ex Willd.) 

Hunz. 67 

Hydromystria stolonifera G. Mey. 67 

Hymenocallis Salisb. 110 

Hymenocallis caribaea (L.) Herb. 110 

Hymenocallis declinata (Jacq.) Sweet 110 

Hymenocallis expansa (Sims) Herb. 110 

Hymenocallis latifolia (Mill.) M. Roem. 110 

Hymenocallis littoralis (Jacq.) Salisb. 110 

Hymenocallis praticola Britton & P. Wilson 112 

Hymenocallis speciosa (L.f. ex Salisb.) Salisb. 112 

Hymenocallis undulata Herb. 112 

Hyophorbe verschaffeltii H. Wendl. 152


Hyospathe pubigera Griseb. & H. Wendl. 144 

Hypaelyptum argenteum Vahl 324 

Hypoëlytrum berterii Spreng. 59 

Hypoporum hirtellum (Sw.) Nees 374 

Hypoporum verticillatum (Muhl. ex Willd.) Nees 382 

Hypoxis L. 97 

Hypoxis decumbens L. 100 

Hypoxis hirsuta (L.) Coville 99 

Hypoxis juncea Sm. var. wrightii Baker 397 

Hypoxis scorzoneraefolia Lam. 97 

Hypoxis wrightii (Baker) Brackett 100 

Hypoxidaceae R. Br. 12 

I 

Inodes causiarum O. F. Cook 149 

Inodes glauca Dammer 149 

Iria monostachyos (L.) Kuntze 241 

Iridaceae Juss. 100 

Iris L. 100 

Isolepis cernua (Vahl) Roem. & Schult. 383 

Isolepis disticha Boeck. 383 

Isolepis vestita Kunth 247 

Ixia chinensis L. 101 

Ixia xiphidium Loefl. 168 

J 

Juncaceae Juss. 234 

Juncellus laevigatus (L.) C. B. Clarke 274 

Juncus L. 234 

Juncus acutus L. 234 

Juncus dichotomus Elliott 236 

Juncus effusus L. 236 

Juncus parviflorus Poir. 341 

Juncus tenuis Willd. 234 

Juncus tenuis var. dichotomus (Elliott) A. W. Wood 236 

Juncus tenuis Willd. var. tenuis 236 

K 

Kaempferia rotunda L. 186 

Kyllinga Rottb. 317 

Kyllinga aphylla (Vahl) Kunth 321 

Kyllinga brevifolia Rottb. 318 

Kyllinga brevifolia Roem. & Schult. 335 

Kyllinga capillaris (Sw.) Griseb. 277 

Kyllinga cayennensis Lam. 258 

Kyllinga cruciformis Schrad. ex Schult. 318 

Kyllinga filiformis Sw. 289 

Kyllinga flexuosa Boeck. 321 

Kyllinga granularis Boeck. 290 

Kyllinga monocephala sensu G. Forster 319 

Kyllinga monocephala sensu Thunberg 318 

Kyllinga nemoralis (J. R. Forst. & G. Forst.) Dandy ex 

Hutchinson & Dalz. 317, 318 

Kyllinga obtusata J. Presl & C. Presl 322 

Kyllinga odorata Vahl 319 

Kyllinga odorata sensu Duss 321 

Kyllinga peruviana Lam. 322 

Kyllinga peruviana var. foliata Kük. 321 

Kyllinga pumila Michx. 321 

Kyllinga pungens Link 322 

Kyllinga sesquiflora Torr. 319 

Kyllinga tibialis Poit. ex Ledeb. 321 

Kyllinga triceps sensu G. Forster 319 

Kyllinga vaginata Lam. 322 

L 

Lagenocarpus Nees 323 

Lagenocarpus guianensis Nees 323 

Lagenocarpus portoricensis Britton 323 

Lagenocarpus tremulus sensu Urban 323 

Languas speciosa (J.C. Wendl.) Small 179 

Latania loddigesii Mart. 153 

Lemna L. 53 

Lemnaceae Gray 52 

Lemna aequinoctialis Welw. 53 

Lemna minor L. 53 

Lemna minor sensu Grisebach 53 

Lemna minuta Kunth 54 

Lemna minuscula Herter 54 

Lemna oblonga Phil. 56 

Lemna perpusilla sensu Britton & P. Wilson 53 

Lemna perpusilla sensu Liogier & Martorell 54 

Lemna perpusilla var. trinervis Austin ex A. Gray 53 

Lemna polyrhiza L. 55 

Lenma punctata G. Mey. 55 

Lemna trinervis (A. Gray) Small 53 

Lemna valdiviana Phil. 54 

Lenticula polyrhiza (L.) Lam. 55 

Leucojum capitulatum Lour. 97 

Licuala grandisH. Wendl. 152 

Liliaceae 386 

Lilium auratum Lindl. 386 

Lilium lancifolium Thunb. 386 

Lilium longiflorum Thunb. 386 

Lilium regale E. H. Wilson 386 

Limnobium Rich. 67 

Limnobium laevigatum (Humb. & Bonpl. ex Willd.) 

Heine 67 

Limnobium spongia (Bosc) Steud. 67 

Limnobium spongia subsp. laevigatum (Humb. & Bonpl. 

ex Willd.) Lowden 67 

Limnobium stoloniferum (G. Mey.) Griseb. 67 

Limnocharis Humb. & Bonpl. 57 

Limnocharitaceae Takht. ex Cronquist 57 

Lipocarpha R. Br. 324 

Lipocarpha argentea (Vahl) R. Br. 324 

Lipocarpha micrantha (Vahl) G. C. Tucker 324 

Lipocarpha senegalensis (Lam.) T. & H. Durand 324 

Livistona chinensis R. Br. 152 

Lobelia aphylla Nutt. 93 

M 

Machaerina Vahl 326 

Machaerina restioides (Sw.) Vahl 326, 327 

Maranta L. 197 

Maranta allouia Aubl. 195 

Maranta arundinacea L. 197 

Maranta cassupo Jacq. 195 

Maranta divaricata sensu authors 198 

Maranta galanga L. 179 

Maranta geniculata (L.) Lam. 199 

Maranta gibba Sm. 198 

Maranta indica Tussac 197

406 


Maranta lutea Aubl. 196 

Marantaceae Petersen 194 

Mariscus aggregatus Willd. 258 

Mariscus aphyllus Vahl 321 

Mariscus brizaeus (Vahl) C. B. Clarke 260 

Mariscus brunneus (Sw.) C. B. Clarke 260 

Mariscus capillaris (Sw.) Vahl 277 

Mariscus cayennensis (Lam.) Urb. 258 

Mariscus confertus (Sw.) C. B. Clarke 262 

Mariscus echinatus sensu C. B. Clarke 262 

Mariscus elatus Vahl 294 

Mariscus filiformis Spreng. 289 

Mariscus flabelliformis Kunth 290 

Mariscus flavus Vahl 258 

Mariscus floridensis C. B. Clarke 269 

Mariscus fuligineus (Chapm.) C. B. Clarke 270 

Mariscus globulosus sensu Urban 262 

Mariscus gracilis Vahl 290 

Mariscus incompletus (Jacq.) Urb. 294 

Mariscus incompletus sensu Urban 276 

Mariscus jamaicensis (Crantz) Britton 251 

Mariscus ligularis (L.) Urb. 275 

Mariscus mutisii Kunth 276 

Mariscus pedunculatus (R. Br.) T. Koyama 327 

Mariscus planifolius (Rich.) Urb. 282 

Mariscus purpurascens (Vahl) C. B. Clarke 282 

Mariscus rufus Kunth 275 

Mariscus squarrosus (L.) C. B. Clarke 287 

Mariscus swartzii A. Dietr. 289 

Mariscus tenuis C. B. Clarke 290 

Mariscus urbanii (Boeck.) C. B. Clarke 292 

Martinezia acanthophylla (Mart.) Becc. 138 

Martinezia caryotifolia Kunth 152 

Mastigoscleria reflexa (Kunth) Nees 380 

Molineria hortensis Britton 116 

Monocotyledons 18 

Monstera Adans. 36 

Monstera adansonii Schott 

Monstera deliciosa Liebm. 36 

Monstera pertusa (Roxb.) Schott 36 

Montbretia × crocosmiiflora Lemoine ex Morren 102 

Moraea northiana Schneev. 104 

Moraea plicata Sw. 102, 104 

Murdannia Royle 162 

Murdannia nudiflora (L.) Brenan 162 

Murdannia scapiflora (Roxb.) Royle 162 

Musa L. 173 

Musa acuminata Colla 173 

Musa balbisiana Colla 173 

Musa bihai L. 174, 175 

Musa humilis Aubl. 177 

Musa paradisiaca L. 173 

Musa sapientum L. 173 

Musa velutina H. Wendl. & Drude 174 

Musaceae Durande 173 

Myrosma cannifolia L. f. 199 

N 

Nageia coriacea (Rich. & A. Rich.) Kuntze 12 

Najas L. 68 

Najas guadalupensis (Spreng.) Magnus 68 

Najas major sensu Bello 68 

Najas marina L. 68 

Narcissus tazetta L. 116 

Neomarica Sprague 104 

Neomarica caerulea (Ker Gawl.) Sprague 104 

Neomarica northiana (Schneev.) Sprague 104 

Neoregelia carolinae (Beer) L. B. Sm. 231 

Neoregelia marmorata (Baker) L. B. Sm. 231 

Neovriesea macrostachya (Bello) Britton 238 

Neowashingtonia filifera (Linden ex André) Sud. 152 

Neowashingtonia robusta (H. Wendl.) Britton 152 

Nicolaia elatior (Jack) Horan. 180 

Nolinaceae 386 

O 

Oncostylis pauciflora Liebm. 245 

Oncostylis vestita (Kunth) Nees 247 

Ophryoscleria microcarpa (Nees ex Kunth) Nees 377 

Orchidaceae 8 

Oreodoxa oleracea sensu Bello 144 

Oreodoxa regia Kunth 146 

P 

Pancratium caribaeum L. 110 

Pancratium declinatum Jacq. 110 

Pancratium expansum Sims 110 

Pancratium latifolium Mill. 110 

Pancratium littorale Jacq. 110 

Pancratium speciosum L.f. ex Salisb. 112 

Pandanaceae 386 

Pandanus baptistii Hort. 386 

Pandanus odoriferus (Forssk.) Chiov. 386 

Pandanus pacificus J.H. Veitch 386 

Pandanus tectorius Du Roi 386 

Pandanus utilis Bory 386 

Pandanus veitchii Dallière 386 

Papyrus comosus Kunth 271 

Philodendron Schott 36 

Philodendron angustatum Schott 38 

Philodendron bipinnatifidum Schott ex Endl. 44 

Philodendron consanguineum Schott 38 

Philodendron fuertesii Krause 38 

Philodendron giganteum Schott 40, 52 

Philodendron grandifolium (Jacq.) Schott 38 

Philodendron hederaceum (Jacq.) Schott 40 

Philodendron isertianum Schott 42 

Philodendron krebsii Schott 38 

Philodendron lingulatum (L.) K. Koch 42 

Philodendron marginatum Urb. 38 

Philodendron micans Klotzsch ex K. Koch 42 

Philodendron nechodomae Britton 34 

Philodendron ornatum Schott 44 

Philodendron oxycardium sensu Britton & P. Wilson 42 

Philodendron radiatum Schott 44 

Philodendron scandens K. Koch & Sello 42 

Philodendron scandens ssp. isertianum (Schott) G.S. 

Bunting 42 

Philodendron urbanianum Krause 38 

Philodendron wrightii Griseb. 38 

Phoenix canariensis Chabaud 152 

Phoenix dactylifera L. 152 

Phoenix reclinata Jacq. 152 

Phoenix roebelinii O’Brien 153 

Phoenix rupicola T. Anders. 152


Phoenix sylvestris (L.) Roxb. 153 

Piaropus diversifolius (Vahl) P. Wilson 171 

Pinaceae Adans. 13 

Pinus L. 13 

Pinus caribaea Morelet 14 

Pinus occidentalis Sw. 14 

Pinus sylvestris L. 14 

Piperaceae 27 

Pistia L. 44 

Pistia occidentalis Blume 44 

Pistia stratiotes L. 44 

Pitcairnia L’Hér. 213 

Pitcairnia angustifolia Sol. 213, 214 

Pitcairnia angustifolia var. simplicior Proctor & Cedeño- 

Mald. 214 

Pitcairnia bromeliifolia L’Hér. 213 

Pitcairnia gracilis Mez 213 

Pitcairnia jareckii Proctor & Cedeño-Mald. 215 

Pitcairnia latifolia Aiton 213 

Pitcairnia ramosa J. Jacq. 213 

Platystachys bulbosa (Hook.) Beer 219 

Pleomele fragrans (L.) Salisb. 130 

Poaceae 8 

Podocarpaceae Endl. 12 

Podocarpus L’Hérit. ex Pers. 12 

Podocarpus coriaceus Rich. & A. Rich 12 

Podocarpus elongatus (W. T. Aiton) L’Hérit. ex Pers. 12 

Podocarpus macrophyllus var. maki Endl. 13 

Polianthes tuberosa L. 125 

Pontederia L. 168 

Pontederia azurea Sw. 170 

Pontederia cordata L. 172 

Pontederia crassipes Mart. 170 

Pontederia limosa Sw. 172 

Pontederiaceae Kunth 168 

Potamogeton L. 76 

Potamogeton epihydrus sensu Britton & P. Wilson 78 

Potamogeton fluitans sensu Britton & P. Wilson 78 

Potamogeton foliosus Raf. 77 

Potamogeton gramineus sensu Michaux 77 

Potamogeton illinoensis Morong 77 

Potamogeton insulanus Hagstr. 78 

Potamogeton natans L. 76 

Potamogeton nodosus Poir. 77 

Potamogeton nuttallii var. portoricensis (Graebn.) 

Graebn. 77 

Potamogeton occidentalis Sieber ex Cham. & Schltdl. 77 

Potamogeton pauciflorus Pursh 76 

Potamogeton pennsylvanicus var. portoricensis Graebn. 77 

Potamogeton plantaginea sensu Bello 78 

Potamogeton pulchelliformis Hagstr. 77 

Potamogetonaceae Dumort. 76 

Pothomorphe 27 

Pothos acaulis Jacq. 24 

Pothos aureus André 34 

Pothos cordatus L. 24 

Pothos crenatus L. 25 

Pothos pinnatus L. 34 

Prestoea Hook. f. 143 

Prestoea acuminata var. montana (Graham) An. Hend & 

Galeano 144 

Prestoea montana (Graham) G. Nicholson 144 

Prestoea pubigera (Griseb. & H. Wendl.) Hook. f. 144 

Pritchardia pacifica Seem. &.H. Wendl. 152 

Protasparagus Oberm. 126 

Protasparagus aethiopicus (L.) Oberm. 126 

Protasparagus setaceus (Kunth) Oberm. 127 

Pseudophoenix H. Wendl. ex Sarg. 145 

Pseudophoenix sargentii H. Wendl. ex Sarg. 145 

Pseudophoenix vinifera (Mart.) Becc. 146 

Psilocarya nitens (Vahl) A. W. Wood 352 

Psilocarya portoricensis Britton 352 

Psilocarya rufa Nees 362 

Psilocarya velutina (Kunth) Nees 362 

Ptychomeria nivea Griseb. 95 

Ptychomeria portoricensis (Urb.) Schltr. 95 

Ptychomeria sphaerocarpa (Urb.) Schltr. 96 

Ptychosperma elegans (R. Br.) Blume 153 

Ptychosperma macarthurii (H. Wendl. ex H.J. Veitch) H. 

Wendl. ex Hook. f. 153 

Pycreus angulatus (Nees) Nees 292 

Pycreus ferrugineus (Poir.) C. B. Clarke 294 

Pycreus flavescens (L.) P. Beauv. ex Rchb. 268 

Pycreus laevigatus (L.) Nees 274 

Pycreus odoratus sensu Urban 283 

Pycreus polystachyos (Rottb.) P. Beauv. 283 

Pycreus polystachyos var.hahnianus (Boeck.) C. B. 

Clarke 283 

Pycreus unioloides (R. Br.) Urb. 292 

R 

Rajania L. 88 

Rajania cordata L. 90 

Rajania cordata L. var. microcarpa Uline ex Knuth 90 

Rajania flexuosa Bello 83 

Rajania hastata L. 90 

Rajania sintenisii Uline 90 

Rajania venosa R. Knuth 90 

Ravenala madagascariensis Sonn. 386 

Rectanthera fragrans (Lindl.) O. Deg. 156 

Remirea Aubl. 327 

Remirea maritima Aubl. 327 

Remirea pedunculata R. Br. 327 

Renealmia L. f. 182 

Renealmia alpinia (Rottb.) Maas 182 

Renealmia antillarum var. puberula Gagnep. 184 

Renealmia aromatica (Aubl.) Griseb. 183 

Renealmia bracteosa Griseb. 182 

Renealmia exaltata L. f. 182 

Renealmia exaltata var. β gracilis K. Schum. 182 

Renealmia jamaicensis var. puberula (Gagnep.) Maas 184 

Renealmia monostachia L. 209 

Renealmia occidentalis (Sw.) Sweet 183 

Renealmia polystachia (L.) L. 222 

Renealmia recurvata L. 223 

Renealmia usneoides L. 225 

Raphia farinifera (Gaertn.) Hyl. 153 

Rhapis excelsa (Thunb.) A. Henry ex Rehder 152 

Rhapis flabelliformis L’Her. 152 

Rhoeo discolor (L’Heritier) Hance 165 

Rhoeo spathacea (Sw.) Stearn 165 

Rhopadostylis baueri (Hook.) H. Wendl. & Drude 152 

Rhynchospora Vahl 329 

Rhynchospora alba (L.) Vahl 330, 331 

Rhynchospora aurea Vahl 341 

Rhynchospora bahamaensis Britton 350, 351

408 


Rhynchospora berteroi (Spreng.) C. B. Clarke 335 

Rhynchospora blauneri Britton 336, 363 

Rhynchospora borinquensis Britton 351 

Rhynchospora brachychaeta C. Wright 336, 363 

Rhynchospora brachychaeta sensu Small 363 

Rhynchospora breviseta (Gale) Channell 336, 364 

Rhynchospora bruneri Britton 355 

Rhynchospora ciliata (G. Mey.) Kük. 337 

Rhynchospora colorata (L.) H. Pfeiff. 340 

Rhynchospora contracta (Nees) J. Raynal 340 

Rhynchospora corymbosa (L.) Britton 341 

Rhynchospora cymosa sensu Torrey 358 

Rhynchospora cymosa sensu Urban 358 

Rhynchospora cyperoides (Sw.) Mart. 348 

Rhynchospora depressirostris M.T. Strong 342, 343, 364 

Rhynchospora distans (Michx.) Vahl 346 

Rhynchospora distans var. β gracillima (C. Wright) 

Kük. 362 

Rhynchospora distans var. tenuis (Baldwin ex A. Gray) 

Britton 362 

Rhynchospora divaricata (Ham.) M.T. Strong 343, 344 

Rhynchospora divergens Chapm. ex M.A. Curtis 363 

Rhynchospora domingensis Urb. 344, 345 

Rhynchospora elongata Boeck. 361 

Rhynchospora fascicularis (Michx.) Vahl subsp. 

fascicularis 346 

Rhynchospora fascicularis (Michx.) Vahl subsp. 

fascicularis var. fascicularis 346 

Rhynchospora fascicularis subsp. fascicularis var. distans 

(Michx.) Chapm. 346 

Rhynchospora filiformis Vahl 335, 346 

Rhynchospora gigantea Link 347 

Rhynchospora gigantea var. latifolia H. Pfeiff. 364 

Rhynchospora glauca Vahl 359 

Rhynchospora globularis var. pinetorum (Small) Gale 364 

Rhynchospora globularis var. recognita Gale 358, 364 

Rhynchospora gracillima C. Wright 362 

Rhynchospora hispidula (Vahl) Boeck. 343, 344 

Rhynchospora holoschoenoides (Rich.) Herter 348 

Rhynchospora intermixta C. Wright 355 

Rhynchospora jamaicensis Britton 349 

Rhynchospora leucocephala (Michx.) Boeck. 340 

Rhynchospora lindeniana Griseb. 350 

Rhynchospora lindeniana var. bahamaensis (Britton) 

Gale 350 

Rhynchsopora longiflora of authors 344 

Rhynchsopora longiflora C. Presl 345, 363 

Rhynchospora longiflora var. domingensis (Urb.) Kük. 344 

Rhynchospora longispicata Boeck. 347 

Rhynchospora luquillensis Britton 333 

Rhynchospora macra (C. B. Clarke ex Britton) Small 363 

Rhynchospora marisculus Nees 351 

Rhynchospora marisculus sensu Britton & P. Wilson 353 

Rhynchospora micrantha sensu C. B. Clarke 341 

Rhynchospora micrantha Vahl 357 

Rhynchospora micrantha var. contracta (Nees) Kük. 340 

Rhynchospora microcarpa Baldwin ex A. Gray 352 

Rhynchospora microcephala (Bertero ex Spreng.) Kük. 356 

Rhynchospora miliacea (Lam.) A. Gray 363 

Rhynchospora nervosa subsp. ciliata (Vahl) T. Koyama 337 

Rhynchospora nervosa (Vahl) Boeck. 339, 363 

Rhynchospora nervosa var. ciliata (Vahl) Kük. 337 

Rhynchospora nervosa var. subfiliformis (H. Pfeiff.) 

Kük. 335 

Rhynchospora nitens (Vahl) A. Gray 352 

Rhynchospora odorata C. Wright ex Griseb. 353 

Rhynchospora oligantha var. breviseta Gale 336 

Rhynchospora oligantha A. Gray 363 

Rhynchospora oligantha sensu Kükenthal 336 

Rhynchospora pallida sensu C. B. Clarke 363 

Rhynchospora pedersenii Guagl. 364 

Rhynchospora perplexa Britton 364 

Rhynchospora pinetorum Britton & Small 364 

Rhynchospora plumosa Elliott 354 

Rhynchospora podosperma C. Wright 347 

Rhynchospora polyphylla sensu Grisebach 355 

Rhynchospora polyphylla (Vahl) Vahl 350 

Rhynchospora pura (Nees) Griseb. 337 

Rhynchospora pusilla Chapm. ex M. A. Curtis 355 

Rhynchospora pusilla (Sw.) Griseb. 335 

Rhynchospora racemosa C.Wright 355 

Rhynchospora radicans subsp. microcephala (Bertero ex 

Spreng.) W.W. Thomas 356 

Rhynchospora rariflora (Michx.) Elliott 357 

Rhynchospora recognita (Gale) Kral 358, 364 

Rhynchospora rufa (Nees) Boeck. 362 

Rhynchospora rugosa (Vahl) Gale 330, 359 

Rhynchospora section Dichromena (Michx.) Vahl 335 

Rhynchospora setacea (P. J. Bergius) Boeck. 360 

Rhynchospora setacea Vahl 357 

Rhynchospora sola Gale 364 

Rhynchospora stellata (Lam.) Griseb. 340 

Rhynchospora subfiliformis H. Pfeiff. 335 

Rhynchospora tenerrima Nees ex Spreng. 360 

Rhynchospora tenuis Willd. ex Link 364 

Rhynchospora tenuis sensu Britton & P. Wilson 342 

Rhynchospora trichodes C. B. Clarke 343, 344 

Rhynchospora uniflora Boeck. 360 

Rhynchospora vahliana Griseb. 337 

Rhynchospora velutina (Kunth) Boeck. 362 

Rhynchospora wrightiana Boeck. 362 

Roystonea O. F. Cook 146 

Roystonea borinquena O. F. Cook 147 

Roystonea hispaniolana L. H. Bailey 147 

Roystonea hispaniolana f. altissima Moscoso 147 

Roystonea oleracea (Jacq.) O.F. Cook 146 

Roystonea peregrina L. H. Bailey 147 

Roystonea regia (Kunth) O. F. Cook 146, 147 

Roystonea venezuelana L. H. Bailey 147 

Ruppia didyma Sw. 75 

Ruppia L. 75 

Ruppia anomala Ostenf. 75 

Ruppia didyma Sw. 75 

Ruppia maritima L. 75 

Ruppiaceae Horan. 74 

S 

Sabal Adans. 147 

Sabal adansonii Guers. 149 

Sabal beccariana L. H. Bailey 149 

Sabal bermudiana L. H. Bailey 149 

Sabal blackburnianum sensu Britton & P. Wilson 149 

Sabal causiarum (O. F. Cook) Becc. 149 

Sabal haitiensis Becc. 149 

Sabal minor (Jacq.) Pers. 149 

Sabal palmetto (Walter) Lodd. ex Schult. f. 150 

Sabal princeps Becc. 149


Sabal questeliana L. H. Bailey 150 

Sabal umbraculifera sensu Bello 150 

Sagittaria L. 60 

Sagittaria acutifolia sensu Bello 61 

Sagittaria acutifolia sensu Grisebach 61 

Sagittaria intermedia Micheli 61 

Sagittaria lancifolia L. 61 

Sagittaria latifolia Willd. 63 

Sagittaria sagittifolia L. 60 

Salvinia laevigata Humb. & Bonpl. ex Willd. 67 

Sansevieria Thunb. 130 

Sansevieria cylindrica Bojer ex Hook. 131 

Sansevieria guineensis (L.) Willd. 131 

Sansevieria hyacinthoides (L.) Druce 130, 131 

Sansevieria metallica Gérome & Labroy 131 

Sansevieria cf. pearsonii N. E. Br. 132 

Sansevieria thyrsiflora Petagna 131 

Sansevieria thyrsiflora Thunb. 131 

Sansevieria trifasciata Prain 131, 133 

Schoenoplectus (Rchb.) Palla 364 

Schoenoplectus americanus (Pers.) Volkart ex Schinz & 

Keller 365 

Schoenoplectus lacustris (L.) Palla 365 

Schoenoplectus olneyi (A. Gray ex Engelm. & A. Gray) 

Palla 365 

Scoenoplectus tabernaemontani (C.C. Gmel.) Palla 366 

Schoenoplectus validus (Vahl) Á. Löve & D. Löve 365 

Schoenus albus L. 330, 333 

Schoenus capillaris Sw. 277 

Schoenus ciliaris Michx. 337 

Schoenus ciliatus G. Mey. 337, 364 

Schoenus cladium Sw. 251 

Schoenus coloratus L. 319, 340 

Schoenus cymosus Muhl. 358 

Schoenus cyperoides Sw. 348 

Schoenus distans Michx. 346 

Schoenus fascicularis Michx. 345 

Schoenus hispidulus Vahl 343, 344 

Schoenus holoschoenoides Rich. 348 

Schoenus latifolius Vahl 376 

Schoenus pusillus Sw. 335 

Schoenus rariflorus Michx. 357 

Schoenus restioides Sw. 326, 327 

Schoenus rugosus Vahl 359 

Schoenus secans L. 380 

Schoenus setaceus P. J. Bergius 360 

Schoenus stellatus Lam. 340 

Scirpus americanus Pers. 365 

Scirpus annuus All. 309 

Scirpus capillaris L. 242 

Scirpus capitatus sensu Grisebach 300 

Scirpus caribaeus Rottb. 300 

Scirpus cernuus Vahl 383 

Scirpus complanatus Retz. 308 

Scirpus constrictus (Schult.) Griseb. 297 

Scirpus corymbosus L. 341 

Scirpus debilis Lam. 311 

Scirpus dichotomus L. 307, 309 

Scirpus diphyllus Retz. 309 

Scirpus elegans Kunth 297 

Scirpus ferrugineus L. 311 

Scirpus ferrugineus var. debilis (Lam.) Poir. 311 

Scirpus flaccidus Rchb. ex Spreng. f. 299 

Scirpus flavescens Poir. 299 

Scirpus geniculatus L. 300 

Scirpus geniculatus var. minor Vahl 300 

Scirpus glomeratus L. 319 

Scirpus glomeratus Retz. 308 

Scirpus hirtus Griseb. 248 

Scirpus interstinctus Vahl 301 

Scirpus intricatus L. 287 

Scirpus lacustris L. 365 

Scirpus lacustris sensu C.B. Clarke 365 

Scirpus lacustris subsp. validus (Vahl) T. Koyama 365 

Scirpus lithospermus L. 375 

Scirpus melanocarpus Griseb. 304 

Scirpus micranthus Vahl 324 

Scirpus miliaceus L. 312, 313 

Scirpus montanus Kunth 302 

Scirpus mutatus L. 302 

Scirpus nitens Vahl 352 

Scirpus nodulosus Roth 302 

Scirpus obtusifolius sensu Grisebach 308 

Scirpus ocreatus (Nees) Griseb. 299 

Scirpus olneyi A. Gray ex Engelm. & A. Gray 365 

Scirpus pachystylus C. Wright 304 

Scirpus palustris L. 295 

Scirpus quinquangularis Vahl 313 

Scirpus radicans Poir. 304 

Scirpus retroflexus Poir. 305 

Scirpus retroflexus sensu Grisebach 303 

Scirpus rostellatus Torr. 305 

Scirpus senegalensis Lam. 324 

Scirpus spadiceus L. 314 

Scirpus subdistichus Boeck. 383 

Scirpus subsquarrosus Muhl. 324 

Scirpus validus Vahl 365 

Scirpus vestitus (Kunth) Rchb. ex Boeck. 247 

Scirpus villosus Poir. 309 

Scirpus viscosus (Sw.) Lam. 266 

Scleria P. J. Bergius 366 

Scleria bracteata Cav. 382 

Scleria brittonii Core 369, 383 

Scleria canescens Boeck. 370 

Scleria catalinae Britt. 378 

Scleria chlorantha Boeck. 379 

Scleria ciliata Michx. 370, 371 

Scleria ciliata var. glabra (Chapm.) Fairey 369 

Scleria cubensis Boeck. 378 

Scleria distans Poir. 371 

Scleria doradoensis Britton 374 

Scleria eggersiana Boeck. 372, 382 

Scleria filiformis Sw. 375 

Scleria flagellum sensu authors 380 

Scleria flagellum-nigrorum P. J. Bergius 368 

Scleria foliosa C. Wright 377 

Scleria georgiana Core 373 

Scleria glabra (Chapm.) Britton 369 

Scleria gracilis Elliott 373 

Scleria grisebachii C. B. Clarke 372 

Scleria guianensis (Nees) Steud. 323 

Scleria havanensis Britton 373, 383 

Scleria hemitaphra Steud. 378 

Scleria hirtella Sw. 374 

Scleria krugiana Boeck. 375 

Scleria latifolia Sw. 376 

Scleria laxa Torr. 378 

Scleria lithosperma (L.) Sw. 375 

Scleria lithosperma var. filiformis (Sw.) Britton 375

410 


Scleria melaleuca Rchb. ex Schltdl. & Cham. 376 

Scleria microcarpa Nees ex Kunth 377 

Scleria microcarpa sensu Grisebach 372 

Scleria microcarpa var. foliosa C. B. Clarke 377 

Scleria microcarpa var. latifolia Boeck. 372 

Scleria microcarpa var. subeciliata C. B.Clarke 378 

Scleria micrantha Poir. 341 

Scleria microdiscus Steud. 335 

Scleria mitis P. J. Bergius 382 

Scleria mitis sensu Grisebach 372 

Scleria mucronata Poir. 378 

Scleria muehlenbergii Steud. 378, 379 

Scleria nutans Willd. ex Kunth 371 

Scleria oligantha Michx. 383 

Scleria ottonis Boeck. 376 

Scleria pauciflora Muhl. ex Willd. 382 

Scleria pauciflora var. glabra Chapm. 369 

Scleria pratensis Lindl. ex Nees 376 

Scleria pterota C. Presl 376 

Scleria purdiei C. B. Clarke 383 

Scleria purpurea Poir. 375 

Scleria reflexa Kunth 380 

Scleria reticularis Michx. 379 

Scleria reticularis var. pubescens Britton 378 

Scleria scaberrima Boeck. 379 

Scleria scabriuscula Schltdl. 383 

Scleria scindens Nees ex Kunth 379 

Scleria secans (L.) Urb. 380 

Scleria setacea sensu authors 378 

Scleria setacea Poir. 360 

Scleria stevensiana Britton 378 

Scleria triglomerata Michx. 379, 383 

Scleria verticillata Muhl. ex Willd. 382 

Setcreasea pallida Rose 164 

Setcreasea purpurea Boom 164 

Sisyrinchium L. 105 

Sisyrinchium bermudiana L. 105 

Sisyrinchium bulbosum Mill. 104 

Sisyrinchium exile E. P. Bicknell 105 

Sisyrinchium latifolium Sw. 102, 104 

Sisyrinchium micranthum Cav. 105 

Sisyrinchium rosulatum E. P. Bicknell 105 

Smilacaceae Vent. 78 

Smilax L. 79 

Smilax aspera L. 79 

Smilax coriacea Bello 79 

Smilax coriacea Spreng. 79 

Smilax coriacea var. ilicifolia (Desv. ex Ham.) O. E. 

Schulz 79 

Smilax domingensis Willd. 80 

Smilax guianensis var. subarmata O. E. Schulz 79 

Smilax havanensis Jacq. 80 

Smilax havanensis var. portoricensis A. DC. 79 

Smilax ilicifolia Desv. ex Ham. 79 

Smilax ilicifolia var. sublappacea sensu O. E. Schulz 79 

Smilax lappacea sensu O. E. Schulz 79 

Smilax rotundifolia Bello 79 

Smilax rotundifolia L. 82 

Smilax subarmata O. E. Schulz 79 

Spathiphyllum Schott 45 

Spathiphyllum lancifolium (Jacq.) Schott 46 

Spathiphyllum wallisii Regel 46 

Spermodon filiformis (Vahl) Nees 346 

Spirodela Schleid. 54 

Spirodela polyrhiza (L.) Schleid. 55 

Spirodela punctata (G. Mey.) C.H.Thomps. 55 

Spironema fragrans Lindl. 156 

Sprekelia formosissima (L.) Herb. 116 

Stenophyllus antillanus Britton 243 

Stenophyllus curassavicus Britton 244 

Stenophyllus harrisii Britton 247 

Stenophyllus portoricensis Britton 245 

Stenophyllus subaphyllus (C. B. Clarke) Britton 247 

Stenophyllus vestitus (Kunth) Britton 247 

Stratiotes nymphoides Willd. 58 

Strelitzia reginae Aiton 386 

Strelitziaceae 386 

Syagrus coronata (Mart.) Becc. 153 

Syagrus romanzoffiana (Cham.) Glassman 152 

Syngonium Schott 46 

Syngonium auritum (L.) Schott 46, 48 

Syngonium podophyllum Schott 46 

Syringodium Kütz. 73 

Syringodium filiforme Kütz. 73 

T 

Tacca chantrieri André 386 

Taccaceae 386 

Taetsia fruticosa (L.) Merr. 129 

Taxus elongata W. T. Aiton 12 

Thalassia Banks & Sol. ex K. D. Koenig 70 

Thalassia testudinum Banks & Sol. ex K. D. Koenig 70 

Thalia L. 198 

Thalia angustifolia C. Wright ex. Griseb. 199 

Thalia geniculata L. 199 

Thalia trichocalyx Gagnep. 199 

Thecophyllum sintenisii (Baker) Mez 230 

Thelmatophace polyrhiza (L.) Godr. 55 

Thrinax Sw. 150 

Thrinax argentea sensu Eggers 141 

Thrinax microcarpa Sarg. 151 

Thrinax morrisii H. Wendl. 151 

Thrinax parviflora Sw. 151 

Thrinax ponceana O. F. Cook 151 

Thrinax praeceps O. F. Cook 151 

Thrincoma alta O. F. Cook 141 

Thringis latifrons O. F. Cook 141 

Thringis laxa O. F. Cook 141 

Thryocephalon nemorale J. R. Forst. & G. Forst. 317, 318 

Tigridia pavonia (L. f.) DC. 106 

Thuja orientalis L. 17 

Tillandsia L. 215 

Tillandsia aloifolia Hook. 221 

Tillandsia angustifolia Sw. 221, 222 

Tillandsia ariza-juliae L. B. Sm. & Jiménez 218 

Tillandsia borinquensis Cedeño-Mald. & Proctor 218 

Tillandsia breviscapa A. Rich. 222 

Tillandsia bromoides Mez 223 

Tillandsia bulbosa Hook. 218, 219 

Tillandsia caraguata D. Dietr. 208 

Tillandsia domingensis Mez 227 

Tillandsia fasciculata Sw. 219 

Tillandsia fasciculata var. venosispica Mez 220 

Tillandsia festucoides Brongn. ex Mez 220 

Tillandsia festucoides sensu Britton & P. Wilson 218 

Tillandsia festucoides sensu Liogier & Martorell 218 

Tillandsia flexuosa Sw. 221


Tillandsia gladioflora H. Wendl. 229 

Tillandsia juncea (Ruíz & Pav.) Poir. 228 

Tillandsia × lineatispica Mez 221 

Tillandsia monostachia (L.) L. 209 

Tillandsia nitida Hook. 206, 207 

Tillandsia polystachia (L.) L. 222 

Tillandsia pruinosa Sw. 222 

Tillandsia psittacina Hook. 228 

Tillandsia pulchella Hook. 224 

Tillandsia pulchra Hook. 224 

Tillandsia ramosa Bello 228 

Tillandsia ramosa Sweet 225 

Tillandsia recurvata (L.) L. 223 

Tillandsia recurvata f. major André 223 

Tillandsia ringens Griseb. 230 

Tillandsia sessiliflora Ruiz & Pav. 207 

Tillandsia setacea Sw. 223 

Tillandsia sintenisii Baker 225 

Tillandsia sublaxa Baker 227 

Tillandsia subulata Vell. 224 

Tillandsia tenuifolia L. 224 

Tillandsia tenuifolia sensu Jacquin 221 

Tillandsia uniflora Kunth 223 

Tillandsia usneoides (L.) L. 225 

Tillandsia utriculata L. 207, 225 

Tillandsia valenzuelana A. Rich. 227 

Tillandsia variabilis Schltdl. 227 

Torulinium confertum Desv. ex Ham. 280 

Torulinium eggersii (Boeck.) C. B. Clarke 265 

Torulinium ferax (Rich.) Urb. 280 

Torulinium filiforme (Sw.) C. B. Clarke 268 

Torulinium flexuosum (Vahl) T. Koyama 269 

Torulinium michauxianum (Schult.) C. B. Clarke 280 

Torulinium odoratum (L.) Hooper 280 

Torulinium vahlii C. B. Clarke 269 

Tradescantia L. 164 

Tradescantia discolor L’Heritier 165 

Tradescantia elongata G. Mey. 166 

Tradescantia geniculata Jacq. 161 

Tradescantia monandra Sw. 154 

Tradescantia multiflora Jacq. 166 

Tradescantia pallida (Rose) D. R. Hunt 164 

Tradescantia pendula (Schnizl.) D. R. Hunt 166 

Tradescantia portoricensis Bello 159 

Tradescantia pulchella Kunth 161 

Tradescantia spathacea Sw. 165 

Tradescantia virginica L. 164 

Tradescantia zanonia (L.) Sw. 165 

Tradescantia zebrina Hort ex Bosse 165 

Trichelostylis complanata (Retz.) Nees 308 

Trichelostylis quinquangularis (Vahl) Nees 313 

Trimezia Salisb. ex Herb. 105 

Trimezia martinicensis sensu Liogier & Martorell 106 

Trimezia meridensis Herb. 105 

Trimezia steyermarkii R. C. Foster 106 

Tripogandra Raf. 166 

Tripogandra elongata (G. Mey.) Woodson 166 

Tripogandra multiflora Raf. 166 

Tripogandra serrulata (Vahl) Handlos 166 

Tripterella capitata (Walter ex J. F. Gmel.) Michx. 94 

Tritonia aurea Pappe ex Hook. 102 

Tritonia crocosmiiflora (Lemoine ex Morren) 

Nicholson 102 

Typha L. 393 

Typha angustifolia sensu Britton & P. Wilson 393 

Typha domingensis Pers. 393 

Typha latifolia L. 393 

Typhaceae Juss. 383 

V 

Veitchia merrillii (Becc.) H. E. Moore 153 

Vogelia capitata Walter ex J. F. Gmel. 94 

Vriesea Lindl. 228 

Vriesea aloifolia (Hook.) Beer 221 

Vriesea guadalupensis sensu Boldingh 230 

Vriesea macrostachya (Bello) Mez 228 

Vriesea psittacina (Hook.) Lindl. 228 

Vriesea ringens (Griseb.) Harms 230 

Vriesea sintenisii (Baker) L. B. Sm. & Pittendr. 230 

W 

Washingtonia filifera (Linden ex André) H. Wendl. 152 

Washingtonia robusta H. Wendl. 152 

Websteria confervoides (Poir.) S.S. Hooper 306 

Werauhia J. R. Grant 229 

Werauhia gladioflora (H. Wendl.) J. R. Grant 229 

Werauhia proctorii Cedeño-Mald. 230 

Werauhia ringens (Griseb.) J. R. Grant 230 

Werauhia sintenisii (Baker) J. R. Grant 230 

Wittmackia lingulata (L.) Mez 202 

Wolffia Horkel ex Schleid. 56 

Wolffia punctata Griseb. 56 

Wolffia brasiliensis Wedd. 56 

Wolffia michelii Schleid. 56 

Wolffia welwitschii Hegelm. 57 

Wolffiella (Hegelm.) Hegelm. 56 

Wolffiella lingulata sensu Liogier & Martorell 57 

Wolfiella oblonga (Phil.) Hegelm. 56 

Wolffiella welwitschii (Hegelm.) Monod 57 

Wolffiopsis welwitschii (Hegelm.) Hartog & Plas 57 

X 

Xanthosoma Schott 48 

Xanthosoma angustisectum Engl. 49 

Xanthosoma atrovirens K. Koch & Bouché 50 

Xanthosoma brasiliense (Desf.) Engl. 49 

Xanthosoma caracu K. Koch & Bouché 51 

Xanthosoma hastatum Eggers 49 

Xanthosoma hastifolium K. Koch 49 

Xanthosoma helleborifolium (Jacq.) Schott 49 

Xanthosoma helleborifolium var. angustisectum (Engl.) 

Engl. 49 

Xanthosoma jacquinii sensu Schott 50 

Xanthosoma nigrum (Vell.) Stellf. 51 

Xanthosoma sagittifolium (L.) Schott 48, 50 

Xanthosoma sylvestre Bello 29 

Xanthosoma undipes (K. Koch) K. Koch 50 

Xanthosoma violaceum Schott 51 

Xiphidium Aubl. 167 

Xiphidium albidum Lam. 168 

Xiphidium caeruleum Aubl. 167, 168 

Xiphidium floribundum Sw. 168 

Xyridaceae C. A. Agardh 231 

Xyris L. 232 

Xyris brevifolia sensu Elliott 233

412 

Xyris caroliniana sensu authors 233 

Xyris communis Kunth 233 

Xyris curtissii Malme 232 

Xyris difformis var. curtissii (Malme) Kral 232 

Xyris elliottii Chapm. 233 

Xyris indica L. 232 

Xyris jupicai Rich. 233 

Xyris neglecta Small 232 

Xyris serotina var. curtissii (Malme) Kral 232 

Y 

Yucca L. 124 

Yucca aloifolia L. 124 

Yucca elephantipes E. Regel ex Trel. 125 

Yucca gloriosa L. 125 

Yucca guatemalensis Baker 125 

Z 

Zamiaceae Horan. 14 

Zamia L. 15 

Zamia allison-armourii Millsp. 17 

Zamia amblyphyllidia D.W. Stev. 16 

Zamia angustifolia var. floridana (A. DC.) Regel 17 

Zamia debilis L. f. 17 

Zamia erosa O.F. Cook & G.N. Collins 17 


Zamia floridana A. DC. 17 

Zamia integrifolia L. f. 17 

Zamia latifoliolata Prenl. 17 

Zamia latifoliolata sensu Britton & P. Wilson 16 

Zamia media Jacq. 17 

Zamia portoricensis Urb. 16 

Zamia pumila L. 15, 17 

Zantedeschia aetiopica (L.) Link 52 

Zebrina pendula Schnizl. 165 

Zephyranthes Herb. 112 

Zephyranthes atamasco (L.) Herb. 113 

Zephyranthes aurea Baker 113 

Zephyranthes carinata Herb. 113 

Zephyranthes citrina Baker 113 

Zephyranthes eggersiana Urb. 113 

Zephyranthes grandiflora Lindl. 113 

Zephyranthes proctorii Acev.-Rodr. & M. T. Strong 115 

Zephyranthes puertoricensis Traub 115 

Zephyranthes rosea Lindl. 115 

Zingiber Mill. 184 

Zingiber cassumunar Roxb. 185 

Zingiber montanum (J. König) Link ex Dietr. 185 

Zingiber officinale Roscoe 185, 186 

Zingiber purpurem Roscoe 185 

Zingiber zerumbet (L.) Roscoe ex Sm. 185 

Zingiberaceae Adans. 178

Abeto 128 

Adelfa 113 

Aguja de Adán 125 

Ala de pájaro 128 

Aloe 135 

Aloe Family 133 

Amapola 109 

Amaranta 198 

Amaryllis Family 106 

Anunciación 109 

Arrowroot Family 194 

Asparagus Family 125 

Banana Family 173 

Bandera española 194 

Barbas de úcar 225 

Bayoneta 125, 129 

Bayoneta española 125 

Bejuco de agua 36 

Bejuco de calabazón 38, 42 

Bejuco de guaraguao 90 

Bejuco de membrillos 80 

Belleza 157 

Bihao 183, 184 

Bloodroot Family 167 

Boca de dragón 179 

Botecitos 175 

Botoncillo 339 

Burmannia Family 90 

Cabellera de la reina 127 

Calabazón cimarrón 42 

Caña amarga 190 

Caña india 188 

Canastillo 127 

Caoba del país 13 

Caobilla 13 

Caribbean pine tree 14 

Cat-tail Family 383 

Cenizo 173 

Century Plant Family 116 

Chamaluco 173 

Chamaluco enano 173 

Chucho 131 

Chufas 267 

Cocomacaco 130 

Coco 142 

Cocotero 142 

Coconut 142 

Cocuisa 122, 131 

Cocuiza 120 

Cohítre 158 

Cohítre azul 159, 162 

Cohítre blanco 165 

Cohítre enano 157 

Cohítre morado 154, 164, 166 

Cola de paloma 168 

Colorado 173 

Colorado blanco 173 

Coquí 100, 286 

Coquí blanco 339 

Coquillo 786 


INDEX TO COMMON NAMES 

Coroso 138 

Cortadera 282, 327, 373 

Cortadora 282, 327, 370, 373 

Cortadora blanca 376 

Cortadora de altura 327, 373, 382 

Cortadora de ciénaga 251 

Costus Family 186 

Cuarenteno 173 

Cuatrofilos 173 

Cuchara 67 

Datil 173 

Ditch-grass Family 74 

Dracena 130 

Drecina 130 

Duckweed Family 52 

Duende amarillo 113 

Duende blanco 115 

Duende rojo 115 

Duende rosado 113 

Dulce nieve 181 

Dumb cane 31 

Dunguey 79, 85, 86 

Dunguey blanco 79 

Eneas 384 

Erizo 214 

Espárrago de jardín 127 

Flecha de agua 61 

Flor de agua 170, 171 

Flor de Culebra 25 

Forrongo 173 

Frog’s bit Family 63 

Gorgojo 97 

Greenbriar 79 

Guapa 33 

Guarán 173 

Guáyaro 90 

Guineito 173 

Guineo cimarrón 175 

Guineo enano 173 

Guineo gigante 173 

Guineo silvestre 175 

Gunda 88 

Helecho plumoso 128 

Heliconia Family 174 

Hícamo 88 

Hierba de Manatí 70 

Hoja de costado 25 

Hoja de sal 197 

Indian-shot Family 191 

Iris Family 100 

Jacinto de agua 170 

Jartón 173 

Jazmín cimarrón 181 

Jazmín del río 181 

Jengibre amargo 186 

Jengibre colorado 185 

Jengibre de jardin 183 

Judío errante 166 

Junco 301, 366 

Junco cimarrón 260

414 

Junco de agua 276 

Junco de aparejos 301 

Junco de ciénaga 271, 301 

Junco de espiga 301 

Junco de playa 329 

Junco fino 296 

Junco pico 342 

Junquillo 298 

Junquito 310 

Karata 120 

Lady bug 110 

Lágrimas de la Virgen 104 

Lambedora 282, 327, 373 

Lenga de gallina 157 

Lengua de chucho 131 

Lengua de vaca 131 

Lentejilla de agua 52 

Lerenes 196 

Lino de agua 78 

Lirio 108, 112 

Lirio blanco 112 

Lirio de playa 112 

Lirio rojo 109 

Lirio turco 109 

Llantén de agua 59 

Llerenes 196 

Llume 143 

Mafafo 173 

Maguey 120, 122 

Maguey criollo 122 

Malanga 30 

Malanga cimarrona 29 

Malanga trepadora 48 

Malango 173 

Manaca 140 

Manatee-grass Family 79 

Mano poderosa 168 

Manzano 173 

Mapuey 88 

Maraca 194 

Maraca amarilla 192 

Maraca boba 192 

Maraca de pantano 192 

Maraca montuna 194 

Maraca morada 194 

Maraca roja 194 

Maravilla 101 

Maricongo 173 

Mariposa 104 

Marunguey 16, 17 

Maranta 198 

Mariposa blanca 181 

Maroon jancole 25 

Mata de huevo 125 

Matagallina 88 

Maya 204 

Moco de pavo 25 

Monte cristo 173 

Monte cristo enano 173 

Morado 172 

Morado verde 173 

Ñame 83 

Ñame de agua 83 

Ñame blanco 83 


Ñame de Guinea 86 

Ñame dunguey 85 

Ñame gulembo 90 

Ñame mapuey 88 

Ñame morado 88 

Ñame vino 88 

Narciso 181, 1884 

Narciso colorado 184 

Nardo 181 

Nidos de gungulén 223 

Ninfa 104 

Niño 173 

Paisaje 42 

Pajón de costa 292 

Paleta de pintor 29 

Palm Family 135 

Palma de abanico 141 

Palma de cocos 142 

Palma de coroso 138 

Palma de coyor 139 

Palma de escoba 150, 152 

Palma de lluvia 143 

Palma de mar 70 

Palma de Sierra 144 

Palma de sombrero 150 

Palma de yaguas 147 

Palma plateada 141 

Palma real 147 

Palma real puertorriqueña 147 

Palmita de jardín 17 

Pámpano 175, 197 

Pandereta 152 

Paragüita 274 

Parásita 209, 222, 229 

Philodendron Family 36 

Piche 173 

Pimienta Angola 179 

Piña 204 

Píña cortadora 214 

Píña de cuervo 214 

Pineapple 204 

Pineapple Family 204 

Pine Family 13 

Pino antillano 14 

Pino hondureño 14 

Piñón 222 

Pirigallo 209, 229 

Pitisilén 198 

Plátano común 173 

Plátano congo 173 

Plátano de indio 175 

Plátano enano173 

Podocarp 13 

Podocarp Family 12 

Pondweed Family 76 

Rábano 31 

Rábano cimarrón 31 

Raíz de zarzaparilla 80 

Rasca garganta 38 

Razor-grass 380 

Red yam 83 

Rush Family 234 

Saeta de agua 61 

Sábila 135

Sangría 165 

Sanguinaria 165 

Sansiviera redonda 131 

Sarsaparilla Family 78 

Scrub-bush 25 

Sedge family 236 

Serrucho 251 

She-dragon Family 128 

Sisal 120 

Spider Lily 112 

Spiderwort Family 153 

Star Grass Family 96 

Tomillo de agua 64 

Water-hyacinth Family 168 

Water-plantain Family 58 

Water-Poppy Family 57 

Water yam 83 

White lily 110 

White yam 83 

Yam Family 82 

Yautía 50 

Yautía amarilla 50 


Yautía blanca 50 

Yautía cimarrona 23, 29 

Yautía de palma 51 

Yautía lila 51 

Yautía madera 52 

Yautía malanga 30 

Yautía morada 51 

Yautía ornamental 23 

Yautía rascana 51 

Yellow-eyed grass Family 231 

Yerba Coqui 286 

Yerba de eneas 384 

Yerba de estrella 339 

Yerba de pato 53 

Yerba de zanja 5 

Yerba manatí 73 

Yuquilla 198 

Zábila 135 

Zamia Family 14 

Zarzaparilla 80 

Zingiber Family 178

abstract - Department of Botany - Smithsonian Institution

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