United States 

Department of 

Agriculture 

Forest Service 

Southern Forest 

Experiment Station 

New Orleans, 

Louisiana 

An Annotated List of the 

Flora of the Bisley Area, 

Luquillo Experimental Forest, 

Puerto Rico 

1987 to 1992 

General Technical Report 

SO-94 

AU~US~ 1993 Jesus Danilo Chinea, Renee J. Beymer, Carlos Rivera, lnes 

Sastre de Jesus, and F. N. Scatena 
\ 

PUERTO RlCO 

Figure 1. - Map of the Bisley watershed area in the northeastern Llcqrrillo Experimental Forest. The area szrrveyed incllrdes Bisley watersheds 1 

and 2 and areas along the Bisley road. Bisle-v 3, the control watershed, 

was itot sur~e~ved. 

(Odum and Pigeon 1970). Other studies have reported 

the plant species composition of various areas in Bisley (Basnet 1990; Garcia Montiel 1991; Heaton and 

Letourneau 1989; Migenis and Ackerman, in press). 

The most comprehensive tree species list of the Bisley 

area was assembled by Perez (1988) a few years 

before Hurricane Hugo. 

VEGETATION 

The forests of the Bisley area have been classified 

as subtropical wet forests (Ewe1 and Whitmore 1973), 

lower montane rain forests (Beard 19441, and singledominant forests (Richards 1966). The Bisley watersheds are covered by a secondary forest of the 

tabonuco type (Dacryodes excelsa Vahl). This forest 

type is part of the Dacryodes-Sloanea association of 

Puerto Rico, Hispaniola, and higher elevation islands 

of the Lesser Antilles. The three dominant species in 

the watersheds-D. excelsa, Sloanea berteriana 

Choisy, and Prestoea n~ontarza (R. Grah.) Nichols.- 

comprise 60 percent of the aboveground biomass, 51 

percent of the basal area, 49 percent of the stem density, and 57 percent of the importance value.' Details 

of the ecology and history of the area have been published elsewhere (Brown and others 1983; Garcia 

Montiel1991; Odum and Pigeon 1970; Scatena 1989). 

Floristic differences between major habitats in the 

Bisley area have been noted. For example, abundance 

and diversity of introduced vascular plants decrease 

with distance from the roads, trails, and streams. Introduced species are almost absent from the upper 

parts of the watersheds, which are the areas least affected by human disturbances. The abundance and 

diversity of bryophytes also increase toward stream 

banks, a trend similar to the observed by Churchill (in 

press) in Ecuador. 

Iscatena, F.N.; Lugo A.E. 1n.d.l Natural disturbances and the 

vegetation in two subtropical wet steepland watersheds of Puerto 

Rico. Illanuscript has been submitted to the "Journal of Ecology." 
Two growth forms dominate the bryophyte flora of 

Bisley: cushions (e.g., Leucobryum, Octoblepharum, 

Syrrhopodon, and Calymperes) and mats (e.g., 

Sematophyllum, Callicostella, Isopterygium, and 

Lejeunaceae) (Sastre de Jesus and Buck, in press). 

Pendent bryophytes are poorly represented. 

STATISTICAL SUMMARY 

The bryophyte flora of the Bisley area contains 52 

species in 40 genera and 21 families, while the vascular flora contains at least 284 species in 215 genera 

and 81 families (table 2). Among the vascular flora, 

Table 2.- Statistical summary of the known flora in the Bisley area 

of the Luquillo Experimental Forest, Puerto Rico, 1987 to 

1992 

Vegetation Families Genera Species 

category 

..---.-.----.-.--.-.--.Number ....................... 

Bryophyta 2 1 40 52 

Peteridophyta 8 27 43 

Gymnospermae 0 0 0 

Angiospermae 73 188 

Dicot herbs 29 

Monocot herbs 57 

Dicot vines 28 

Woody shrubs" 20 

Trees* 107 

Total 102 255 336 

"Categories based on Britton and Wilson (1923, 1925), Little and 

others (1974), Little and Wadsworth (1989), as well as the authors' 

field experience. 

Table 3.-Number of native, endemic, and introduced species 

among the plant groups in the Bisley area of the Luquillo 

Experimental Forest, Puerto Rico, 1987 to 1992 

Plant Introduced 

iTrOUP Natives Endemics* species Total 

-.------..----.--..-.--Number of species -----.---.-.----.---- 

B~YoP~Y~~ 52 0 0 52 

Pteridophyta 43 3 0 43 

Dicotyledones 164 26 17 181 

Monocotyledones 55 1 5 60 

the major growth forms are represented by 107 

species of trees, 20 species of shrubs, 28 species of 

dicotyledonous vines, 86 species of herbs, and 43 

species of ferns. 

Ninety-three percent of the listed species are natives of Puerto Rico; only 22 species (all of them angiosperms) are introduced species (table 3). Thirty of 

the 314 native species (mostly dicotyledones) are endemic to Puerto Rico; these species make up 9 percent 

endemism among all plant species, or 11 percent endemism among the vascular plants. The largest 

families of flora in the Bisley area, those with 10 

species or more, are listed in table 4. 

Table 4.- The 9 largest families of flora (each had 10 or more 

species) in the Bisley area of the Luquillo Experimental 

Forest, Puerto Rico, 1987 to 1992 

Family Number of species 

Bryophyta 

Lejeuneaceae 

Pteridophyta 

Polypodiaceae 

Dicotyledones 

Lauraceae 

Leguminosae 

Melastomataceae 

Rubiaceae 

Compositae 

Monocotyledones 

Cyperaceae 

Orchidaceae 

Total 314 30 22 336 

"Endemics are a subgroup of native species. 
ANNOTATED LIST 

Nomenclature follows van der Wijk and others 

(1959-69) for mosses, Gradstein (1989) for hepatics, 

Proctor (1989) for ferns, Ackerman and Del Castillo 

(in press) for orchids, and Liogier and Martorell 

(1982) for most other species. 

The annotations vary somewhat, reflecting the different collaborators' comments. The absence of an annotation usually indicates that the species is uncommon or that it has not been observed after the original 

collection. 

Division: BRYOPHYTA 

Class: MUSCI (Mosses) 

Family: Fissidentaceae 

Fissidens inaequalis Mitt. Common in clay soil, 

mineralized clay. 

F. mollis Mitt. 

F. repandus Wils. ex Mitt. On riparian rocks. 

F. zollegeri Mont. In mineralized clay. 

Family: Dicranaceae 

Dicranella perrottetii (Mont.) Mitt. On clay soil of 

uprooted trees. 

Leucoloma cruegerianum (C. Mull.) Jaeg. On tree 

trunks, rarely on logs. 

Family: Leucobryaceae 

Leucobrym martianum (Hornsch.) Hampe ex C. 

Mull. On bases of tree trunks and logs. 

Octoblepharum albidum Hedw. On bases of tree 

trunks and logs. 

Family: Calymperaceae 

Claymperes erosum C. Miill. On tree trunks and 

logs. 

C. nicaraguense Ren. & Card. On logs. 

Syrrhopodon incompletus var. berteroanus (Brid.) 

Reese. On tree trunks and logs. 

S. lgulatus Mont. On logs. 

S. parasiticus var. parasiticus Florsch. On logs. 

Family: Bryaceae 

Bryum truncorum Brid. On riparian rocks. Archegonia observed in May. 

Family: Rhizogoniaceae 

Pyrrhobryum spiniforme (Hedw.) Mitt. On tree 

trunks. Sporophytes produced in February. 

Family: Bartramiaceae 

Philonotis elongata (Dism.) Crum et Steere. On wet 

soil, rocks, rarely on logs; also on soil of uprooted 

trees. 

Family: Orthotrichaceae 

Groutiella apiculate (Hook.) Crum et Steere. On 

logs. Sporophytes observed in April. 

Family: Neckeraceae 

Neckeropsis disticha (Hedw.) Kindb. On bases of tree 

trunks and logs. Sporophytes produced March to 

April. 

N. undulata (Hedw.) Reichdt. On bases of tree 

trunks, logs, and rocks. 

Family: Callicostaceae 

Callicosta evanescens (C. Mull.) Crosby. On logs in 

stream. 

Callicostella depressa (Hedw.) Jaeg. Common on 

logs. 

C. pallida (Hornsch.) hgstr. Mostly found on logs. 

Sporophytes produced in February. 

Crossomitrium patrisiae (Brid.) C. Mull. On logs in 

shady areas. 

Cyclodictyon albicans (Hedw.) Broth. On logs. 

C. varians (Sull.) 0. Kuntze 

Family: Leucomiaceae 

Leocomium strumosum (Hornsch.) Mitt. On logs and 

rocks. 

Family: Thuidiaceae 

Thuidium urceolatum Lor. On logs and soil. 
Family: Sematophyllaceae Order: METZGERIALES 

Sematophyllum adnatum (Michx.) Britt. On logs. 

S. subpinatum (Brid.) Britt. On logs. Family: Aneuraceae 

Taxithelium planum (Brid.) Mitt. 

Trichosteleum sentosum (Sull.) Jaeg. Riccadia digitiloba (Spruce) Pagan. On logs and 

rocks. 

Family: Hypnaceae 

Family: Metzgeriaceae 

lsopterygium tenerum (Sw.) Mitt. On logs. Sporophytes observed in February. Metzgeria uncigera Evans. On logs and rocks. 

Family: Pallaviciniaceae 

C1ass:HEPATICAE (hepatics) 

Order: JUNGERMANNIALES 

Family: Lepidoziaceae 

Bazzania schwanecheana (Hampe & Gott.) Trev. On 

logs. w 

Telaranea nematodes (Gott. ex Aust.) Howe. On logs 

and leaf litter. 

Pallavicinia lyellii (Hook.) S. Gray 

Division: PTERIDOPHYTA 

(Ferns and fern-allies) 

Family: Psilotaceae 

Family: Geocalycaceae Psilotum nudum (L.) P. Beauv. Over bridge on Bisley Road at Bisley 3. 

Lophocolea bidentata (L.) Dum. On logs and rocks. 

Family: Lycopodlaceae 

Family: Plagiochilaceae 

Plagiochila sp. 

Lycopidium cernum L. Outside watersheds, along 

roadsides. 

Family: Lejeuneaceae Family: Selaginellaceae 

Archilejeunea parviflora (Nees) Schiffn. On logs and Selaginella krugii Hieron. Entire area. Endemic. 

tree trunks. 

Ceratolejeunea patentissima (Hampe & Gott.) 

Evans. On logs and tree trunks. Family: Marattiaceae - 

C. valida Evans 

Cheilolejeunea rigidula (Nees ex Mont.) Schust. On Danaea elliptica J.E. Smith 

logs. D. nodosa (L.) J.E. Smith. Entire area. 

~olura rynchophora Jov.-Ast. On leaves of shrubs 

beside creek; common after rainy season. Family: Gleicheniaceae 

Cyclolejeunea luteola (Spruce) Grolle 

Lejeunea sp. On logs. Dicranopteris flexuosa (Schrad.) Underw. 

Macrolejeunea cerina (Lehm. & Lindenb.) Gradst. Gleichenia bifida (Willd.) Spreng. Along roadsides. 

On logs - and tree trunks. 

Marchesinia brachiata (Sw.) Schiffn. On fern leaves 

beside creek. Family: Hymenophyllaceae 

Microlejeunea acutifolia Steph. 

M. bullata (Tayl.) Steph. 

Odorztolejeunea lunulata (Web.) Schiffn. 

Prionolejeunea innovata Evans 

Trichomanes rigidum Sw. On high, shady slopes. 
Family: Cyatheaceae 

Cyathea horrida (L.) J.E. Smith. Entire area. 

C. arborea (L.) J.E. Smith. Entire area. 

C. borinquena (Maxon) Domin. Endemic. 

C. portoricensis Spreng. ex Kuhn. Entire area, except sunny areas. Endomic. 

Family: Polypodiaceae 

Adiantum latifolium Lam. Entire area. 

A. pyramidale (L.) Willd. In the watersheds. 

Arachniodes chaerophylloides (Poir.) Proctor 

Asplenium auritum Sw. 

A. salicifolium L. 

A. serratum L. In the watersheds. 

Blechnum occidentale L. In the watersheds. 

Bolbitis aliena (Sw.) Alston. Rare in the watersheds. 

B. nicotianifolia (Sw.) Alston. Very common in the 

watersheds. 

Ctenitis subincisa (Willd.) Ching 

Dennstaedtia bipinnata (Cav.) Maxon. Entire area. 

D. obtusifolia (Willd.) Moore. Entire area. 

Elaphoglossum sp. Entire area. 

Hemidictym marginatum (L.) K. Presl. Inside and 

outside the watersheds; near creek beds. 

Hypolepis repens (L.) K. Presl. Common in sunny, 

disturbed areas. 

Lonchitis hirsuta L. Riparian. 

Nephrolepis exaltata (L.) Schott 

Nephrolepis rivularis (Vahl) Mett. ex Krug. In the 

watersheds, along roadsides. 

Odontosoria aculeata (L.) J. Smith. Entire area, in 

drier areas. 

Pityrogramma calomelanos (L.) Link var. aureoflava 

(Hooker) Weatherby ex Bailey 

P. calomelanos (L.) Link var. calomelanos. Entire 

area. 

Polybotrya cervina (L.) Kaulf. Entire area, not common. 

Polypodium aureum L. 

P. crassifolium L. Upper slopes of watersheds. 

P. piloselloides L. In the watersheds, epiphytic. 

P. spp. Many epiphytic species. 

Pteris altissima Poir. In the watersheds. 

Tectaria trifoliata (L.) Cav. Entire area. 

Thelypteris balbisii (Spreng.) Ching 

T. deltoidea (Sw.) Proctor. Entire area. 

T. reticulate (L.) Proctor. Sunny areas, roadsides. 

Division: EMBRYOPHSTA 

Class: ANGIOSPERMAE (flowering plants) 

Subclass: DICOTYLEDONES 

Family: Casuarinaceae 

Casuarina equisetifolia J.R. & G. Forst. Along roadsides. Introduced from Australia. 

Family: Piperaceae 

Lepianthes peltat um (L.) Rafinesque. Entire area. 

Peperomia spp. Several species in the watersheds. 

Piper aduncum L. Along roadsides, in sunny areas. 

P. glabrescens (Miq.) C. DC. Entire area. 

P. hispidum Sw. Entire area. 

P. jacquemontianum Kunth. Along roadsides, near 

the creek in Bisley 1. 

Family: Chloranthaceae 

Hedyosmum arborescens Sw. In the watersheds before Hurricane Hugo. 

Family: Ulmaceae 

Trema micranthum (L.) Blume. Along roadsides; on 

high, disturbed slopes. 

Family: Moraceae 

Artocarpus altilis (S. Park.) Fosb. In drainages 

along road. Introduced from islands of the South 

Pacific. 

Cecropia peltata L. Entire area. 

Ficus citrifolia P. Miller. Inside and outside the 

watersheds. 

Family Urticaceae 

Pilea inaequalis (Juss. ex Poir.) Wedd. Entire area. 

P. krugii Urban. On upper slopes of Bisley 2. Endemic. 

Pilea obtusata Liebm. 

Urera baccifera L. Wedd. Entire area. 

U. sp. In the watersheds. 

Family: Phytolaccaceae 

Phytolacca rivinoides Kunth & Bouch6. Entire area. 

Trichostigma octandrum (L.) H. Walt 
Family: Menispermaceae Family: Leguminosae 

Subfamily: Minosoideae 

Cissampelos pareira L. Entire area. 

Family: Magnoliaceae 

Magnolia splendens Urban. In the watersheds, especially on ridges. Endemic. 

Family: Annonaceae 

Guatteria car-ibaea Urban. In the watersheds. 

Family: Lauraceae 

Aniba bracteata (Nees) Mez. Entire area. 

Beilschmiedia pendula (Sw.) Hemsl. In the water- - - sheds. 

Licaria triandra (Sw.) Kostermans 

Ocotea floribunda (Sw.) Mez. In the watersheds. 

0. giobosa (Aubl.) Schlecht. & Cham. 

0. LeucoxyZon (Sw.) Mez. Common in entire area. 

0. membranacea (Sw.) Howard 

0. moschata (Meissn.) Mez. In the watersheds. Endemic. 

0. portoricensis Mez. In the watersheds. Endemic. 

0. sinienisii (Mez) Alain. In the watersheds. 

Persea americana Miller. Along roadsides. Introduced from Mexico. 

Family: Chrysobalanaceae 

HirteLLa rugosa Pers. Entire area. Endemic. 

H. triandra Sw.2 Observed in Bisley 1 before Hurricane Hugo, but not found afterwards. 

Family: Rosaceae 

Rubus rosifolius Smith. Along roadsides. Introduced 

from Southeast Asia. 

Family: Connaraceae 

Roz~rea surinamensis Miq. Entire area. 

'~arcia, Diane. 1989. Personal communication (letter). On file 

with: International Institute of Tropical Forestry, U.S. Department of Agriculture, Forest Service, Rio Piedras, PR 00928. 

Inga fagifolia (L.) Willd. Entire area. 

I. vera Willd. Entire area. 

Mimosp pudica L. Along roadsides. 

Family: Leguminosae 

Subfamily: Caesalpiniodeae 

Cassia aeschinomene DC. Along roadsides. 

Family: Leguminosae 

Subfamily: Papilionoideae 

Andira inermis (W. Wright) HBK. Entire area. 

Desmodium adscendens (Sw.) DC. 

Neorudolphia volubilies (Willd.) Britton. Entire 

area. Endemic. 

Ormosia krugii Urban. Entire area. 

Pterocarpus officinalis Jacq. Inside and outside the 

watersheds. 

Pueraria phaseoloides (Roxb.) Benth. Along roadsides. Introduced from Asia. 

Family: Rutaceae 

Citrus X paradisi Macfad. Along roadsides. Introduced from other islands of the West Indies. 

Ravenia urbanii Engler. In the watersheds, mostly 

on high ridges. Endemic. 

Zanthoxylum rnartinicense (Lam.) DC. Entire area. 

Family: Burseraceae 

Dacryodes excelsa Vahl. Entire area. 

Tetragastris balsamifera (Sw.) Kuntze. In the watersheds. 

Family: Meliaceae 

Guarea glabra Vahl. Entire area. 

G. guidonia (L.) Sleumer. Entire area. 

Khaya nyasica Stapf. ex Baker. Entire area. Introduced from Africa. 

Swietenia macrophylla G. King. Entire area. Introduced from Mexico and Central America. 

Trichilia pallida Sw. Entire area. 

Family: Malpighiaceae 

Byrsonima spicata (Cav.) HBK. Entire area. 

Heteropteris laurifolia (L.) A. Juss. Entire area. 
Family: Polygalaceae Family: Vitaceae 

Securidaca uirgata Sw. Common in entire area. 

Family: Euphorbiaceae 

Alchornea latifolia Sw. Entire area. 

Alchorneopsis floribunda (Benth.) Muell. Arg. Entire area. 

Croton poecilanthus Urban. Entire area. Endemic. 

Drypetes glauca Vah. Entire area. 

Phyllanthus urinaria L. Along roadsides. Introduced from Asia. 

Sapium laurocerasus Desf. Entire area. Endemic. 

Family: Anacardiaceae 

Comocladia glabra (Schultes) Spreng. Entire area. 

Magifera indica L. Along roadsides and in the 

watersheds. Introduced from Asia. 

Spondim mombin L. In the watersheds. 

Family Cyrillaceae 

Cyrilla racemiflora L. Reported by Perez (1988) but 

found only in Bisley 3. 

Cissus erosa L.C. Rich. Along roadsides. 

C. sicyoides L. Entire area. 

C. uerticillate (L.) Nicolson & Garvis. Along roadsides. 

Family: Elaeocarpaceae 

Sloanea berteriana Choisy. Entire area. 

Family: Malvaceae 

Pauonia fruticosa (Miller) Fawc. & Rendle. Entire 

area. 

Sida rhombifolia L. Along roadsides. 

Thespesia grandiflora DC. In the watersheds. Endemic. 

Urena lobata L. Along roadsides. 

Family: Bombacaceae 

Ochroma lagopus Sw. Several individuals recruited 

after Hurricane Hugo in Bisley 2 near the road. 

Quararibea turbinata (Sw.) Poir. In the watersheds. 

Family: Dilleniaceae 

Family: Stahyleaceae 

Pinzona coriacea Mart. & Zucc. In the watersheds. 

Turpinia occidentalis (SW.) G. Don. In the watersheds. 

Family: Sapindaceae 

Cupania americana L. In the watersheds. 

Matayba domingensis (DC.) Radlk. In the watersheds. 

Paullinia pinnata L. Entire area. 

Family: Sabiaceae 

Meliosma herbertii Rolfe. Entire area. 

Family: Balsaminaceae 

Impatiens wallerana H0ok.f. Along roadsides. Introduced from Zanzibar. 

Family: Ochnaceae 

Sauvagesia erecta L. Entire area. 

Family Marcgraviaceae 

Marcgravia rectiflora Triana & Planch. Entire area. 

M. sintenisii Urban. Endemic. 

Family: Theaceae 

Laplacea portoricensis (-Krug & Urban) Dyer. In the 

watersheds. 

Family: Guttiferae 

Calophyllum brasiliense Jacq. In the watersheds. 

Clusia clusioides (Griseb.) D'Arcy. In the watersheds. 

C. gundlachii Stahl. Entire area. Endemic. 

Rheediaportoricensis Urban. In the watersheds. Endemic. 
Family: Flacourtiaceae 

Casearia arborea (L.C. Rich.) Urban. Entire area. 

C. guianensis (Aubl.) Urban 

C. sylvestria Sw. 

Homalium racemosum Jacq. 

Laetia procera (Poepp. & Endl.) Eichl. Entire area. 

Family: Passifloraceae 

Passiflora edulis Sims. Along roadsides. Introduced 

from Brazil. 

P. rubra L. In the watersheds. 

Family: Begoniaceae 

Begonia decandra Pavon. In the watersheds. Endemic. 

Family: Lythraceae 

Cuphea carthagenensis (Jacq.) Macbride 

C. strigulosa HBK. 

Family: Rhizophoraceae 

Cassipourea guianensis Aubl. In the watersheds. 

Family: Combretaceae 

Buchenavia capitata (Vahl) Eichl. Inside and outside the watersheds. 

Family: Myrtaceae 

Calyptranthes pallens (Poir.) Griseb. Near water 

well field in lower end of Bisley 2. 

Eugenia eggersii Kiaersk. Endemic. 

E. stahlii (Kiaersk.) Krug & Urban. Endemic. 

Myrcia deflexa (Poir). DC. In the watersheds. 

M. leptoclada DC. In the watersheds. 

M. splendens (Sw.) DC. In the watersheds. 

Syzgium jumbos (L.) Alst. Introduced from Asia. 

Family: Melastomataceae 

Calycogonium squamulosum Cogn. Endemic. 

Henriettea fascicularis (Sw.) Gdmez Maza 

Heterotrichum cyrnosum (Wendl.) Urban. Endemic. 

Mzconia laevigata (L.) DC. 

M. prasina (Sw.) DC. In the watersheds. 

M. racemosa (Aubl.) DC. Common in the entire area. 

M. serrulata (DC.) Naud. In the watersheds. 

M. tetrandra (Sw.) D. Don 

Nepsera aquatica (Aubl.) Naud. Entire area. 

Tetrazygia urbanii Cogn. In the watersheds and 

along roadsides. Endemic. 

Family: Onagraceae 

Ludwigia octovaluis (Jacq.) Raven. Along roadsides 

and on sunny ridgetops. 

Family: Araliaceae 

Dendropanax arboreus (L.) Decne & Planch. 

Didymopanax morototoni (Aubl.) Decne. & Planch. 

Entire area. 

Family: Myrsinaceae 

Parathesis crenulata (Vent.) Hook. f. 

Wallenia pendula (Urban) Mez. In the watersheds. 

Endemic. 

Family: Sapotaceae 

Chrysophyllum argenteum Jacq. Reported before 

Hurricane Hugo but not seen again. 

Chrysophyllum cainito L. In Bisley 1. 

Manilkara bidentata (A. DC.) A. Chev. Entire area. 

Micropholis chrysophylloides Pierre 

M. garciniifolia Pierre. Endemic. 

Family: Oleaceae 

Chionanthus domingensis Lam. 

Family: Apocynaceae 

Allanzanda cathartics L. Introduced from South 

America. 

Forsteronia portoricensis Woods. Endemic. 

Family: Convolvulaceae 

Ipomoea setifera Poir. 

I. tiliacea (Willd.) Choisy 

Family: Boraginaceae 

Cordia borinquensis Urban. Endemic. 

C. sulcata DC. In the watersheds. 

Tournefortia sp. In Bisley 2. 
Family: Verbenaceae Family: Compositae 

Lamtana camara L. Along roadsides. 

Stachytarpheta jamaicensis (L.) Vah. Along roadsides. 

Vitex divaricata Sw. 

Family: Solanaceae 

Cestrum macrophyllum Vent. Along roadsides. 

Solanum torvum Sw. Entire area; in sunny places. 

Family: Scrophulariaceae 

Bacopa stricta (Schrad.) Robins 

Family: Bignoniaceae 

Tabebuia heteropylla (DC.) Britton. Entire area. 

Aster subulatus Michx. Along roadsides. 

Bidens alba (L.) DC. var. radiata (Sch.-Bip.) Ballard. Along roadsides. 

Clibadium erosum (Sw.) DC. In the watersheds and 

along roadsides; in sunny, disturbed areas. 

Elephantopus mollis Kunth 

E. spicatus Juss. ex Aubl. 

Eupatorium odoratum L. 

Mikania cordifolia (L.f.1 Willd. In the watersheds. 

M. fragilis Urban. In watersheds. Endemic. 

Neurolaena lobata (L.) Cass. 

Rolandra fruticosa.(L.) Kuntze 

Synedrella nodiflora (L.) Gaertn. Along roadsides. 

Vernonia cinerea (L.) Less. 

Wedelia trilobata (L.) A.S. Hitchc. Along roadsides. 

Subclass: MONOCOTYLEDONES 

Family: Acanthaceae 

Family: Gramineae 

Odontonema strictum (Nees) Kuntze. Introduced 

from Central America. 

Ruellia coccinea (L.) Vahl. Entire area. 

Teliostachya alopecuroidea (Vahl) Nees. Along roadsides. 

Thunbergia alata Bojer. Introduced from Africa. 

Family: Rubiaceae 

Coffea arabica L. Introduced from Abyssinia. 

Faramea occidentalis (L.) A. Rich. 

Gonzalagunia spicata (Lam.) Gomez Maza. Entire 

area. 

Hamelia patens Jacq. 

Hemidiodia ocimifolia (Willd. ex R. & S.) K. Schum. 

Hillia parasitica Jacq. In the watersheds. 

Ixora ferrea (Jacq.) Benth. 

Palicourea crocea (Sw.) Roem. & Schult. var. crocea 

P. crocea var. riparia (Benth.) Griseb. In the 

watersheds. 

Psychotria berteriana DC. Common in entire area. 

P. brachiata Sw. Entire area. 

P. maleolens Urban. Endemic. 

Rodeletia portoricensis Krug & Urban. Endemic. 

Sabicea hirsuta HBK. Entire area. 

Family: Cucuritaceae 

Cayaponia racemosa (Miller) Cogn. In the watersheds. 

Andropogon bicornis L. 

A. leucostachyus HBK. 

Bambusa vulgaris Schrad. ex Wendl. Along roadsides. Introduced from Asia. 

Ichnanthus pallens (Sw.) Munro. Common in the 

watersheds. 

Olyra latifolia L. Fairly common after Hurricane 

Hugo; less common as canopy closes. 

Panicum trichoides Sw. 

Paspalum conjugatum Berg. In the watersheds. 

P. virgatum L. Along roadsides. 

Pharus latifolius L. In the watersheds. 

Family: Cyperaceae 

Eleocharis interstincta (Vahl) R. & S. Along roadsides. 

E. retroflexa (Poir.) Urban. In wet places along roadsides. 

Fimbristylis dichotoma (L.) Vahl. In wet places 

along roadsides. 

Fuirena umbellata Rottb. Along roadsides. 

Mariscus ligularis (L.) Urban. Entire area. 

Rhynochospora nervosa (Vahl) Boeck. ssp. ciliata 

(Vahl) T. Koyama 

R. stellata (Lam.) Griseb. Along roadsides. 

Scleria canescens Boeck. Entire area. Endemic. 

S. lithosperma (L.) Sw. 

S. pterota Presl. 
Family: Palmaceae Family: Zingiberaceae 

Prestoea montana (R. Grahm) Nichols. Entire area. 

Roystonea borinquena O.F. Cook. Along roadsides. 

Family: Araceae 

Aglaonema pictum (Roxb.) Kunth. Introduced from 

East Indies. 

Anthurium crenatum (L.) Kunth 

Dieffenbachia sp. May be an introduced species. 

Philodendron angustaturn Schott 

P. scandens C. Koch & H. Sello 

Xanthosoma atrovirens C. Koch & Bouche. Introduced from South America. 

Family: Commelinaceae 

Commelina diffusa Burm.f. In the watersheds and 

along roasides. 

Commelinopsis persicariifolia (DC.) M. Pichon. In 

the watersheds. 

Zebrina pendula Schniz. Mostly along roadsides. 

Family: Pontederiaceae 

Heteranthera reniformis Ruiz & Pavon. In mud, 

along roadsides. 

Family: Smilacaceae 

Smilax domingensis Willd. 

S. havanensis Jacq. Common in the watersheds. 

Family: Hypoxidaceae 

Hypoxis decumbens L. In tire tracks in the road. 

Family: Dioscoreaceae 

Dioscorea polygonoides Humb. & Bonpl. ex Willd. 

Entire area. 

Dioscorea sp. On upper slope. With spines and 

bulbs. 

Rajania cordata L. 

Family: Musaceae 

Heliconia sp. Entire area. 

Musa spp. In the watersheds. Introduced from 

India. 

Zingiber sp. Along the creeks and roadsides. Introduced from Asia. 

Family: Cannaceae 

Canna glauca Entire area. 

Family: Orchidaceae 

Cranichis muscosa Sw. Terrestrial. 

Cyclopogon cranichoides (Griseb.) Schltr. Terrestrial. 

Encyclia cochleata (L.) Dressler. Epiphytic or 

lithophytic. 

Epidendrum carpophorum Barbosa Rodrigues. 

Epiphytic. 

E. nocturnum Jacq. Epiphytic or occasionally 

lithophytic. 

E. ramosum Jacq. Epiphytic. 

E. tridens poeppig & Endlicher. Epiphytic or occasionally lithophytic. 

Erythrodes hirtella (Sw.) Fawc. & Rendle. Terrestrial. 

E. plantaginea (L.) Fawc. & Rendle. Terrestrial. 

Jacquiniella globosa (Jacq.) Schltr. Epiphytic. 

Liparis nervosa (Thumb.) Lindl. Terrestrial. 

Maxillaria coccinea (Jacq.) L. 0. Williams ex Hodge. 

Common epiphyte, especially on Guarea. 

Pleurothallis ruscifolia (Jacq.) R. Br. Common 

epiphyte, especially on Guarea. 

Polystachya concreta (Jacq.) Garay & Sweet. 

Epiphytic or lithophytic. 

P. foliosa (Hook.) Rchb.f. Epiphytic or lithophytic. 

Prescottia oligantha (Sw.) Lindl. Terrestrial. 

Scaphyglottis modestra (Rchb.f'.) Schltr. Epiphytic 

or lithophytic. 

Triphora latifolia G. Luer. Very rare terrestrial. 

Wullschlaegelia aphylla (Sw.) Rchb.f. Achlorphyllous, terrestrial. 

LITEUTURE CITED 

Ackerman, James D.; Del Castillo, Maria. [In press]. 

The orchids of Puerto Rico and the Virgin Islands: 

Las orquideas de Puerto Ric0.y Las Islas Virgenes. 

Rio Piedras, PR: Editorial de la Universidad de 

Puerto Rico. 

Basnet, Khadga. 1990. Studies of ecological and geological factors controlling the pattern of tabonuco 

forests in the Luquillo Experimental Forest, Puerto 

Rico. New Brunswick, NJ: Rutgers, The State 

University of New Jersey. 167 p. Ph.D. dissertation. 
Beard, J.S. 1944. Climax vegetation in tropical 

America. Ecology. 25(2): 127-158. 

Britton, N.L.; Wilson, Percy. 1923. Scientific survey of 

Porto Rico and the Virgin Islands. New York: New 

York Academy of Sciences. 626 p. Vol. 5. 

Britton, N.L.; Wilson, Percy. 1925. Scientific survey of 

Porto Rico and the Virgin Islands. New York: New 

York Academy of Sciences. 663 p. Vol. 6. 

Brown, Sandra; Lugo, Ariel E.; Silander, Susan; 

Liegel, Leon. 1983. Research history and opportunities in the Luquillo Experimental Forest. Gen. 

Tech. Rep. SO-44. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern 

Forest Experiment Station. 128 p. 

Churchill, Steven P. [In pressl. The mosses of Equatorian Amazonas. Aarhus, Denmark: University of 

Aarhus Botanical Institute AAU report. Vol. 24. 

Ewel, J.J.; Whitmore, J.L. 1973. The ecological life 

zones of Puerto Rico and the U.S. Virgin Islands. 

Res. Pap. ITF-18. Rio Piedras, PR: U.S. Department 

of Agriculture, Forest Service, Institute of Tropical 

Forestry. 72 p. 

Garcia Montiel, Diana. 1991. The effect of human activity on the structure and composition of a tropical 

forest in Puerto Rico. Rio Piedras, PR: University of 

Puerto Rico. 103 p. M.S. thesis. 

Gradstein, Stephan R. 1989. A key to the Hepaticae 

and Anthocerotae of Puerto Rico and the Virgin Islands. The Bryologist. 92: 329-348. 

Heaton, K.; Letourneau, A. 1989. Changes in forest 

structure and composition along a gradient from 

streams to ridges in a subtropical moist forest in 

Puerto Rico. Tropical Resource Institute report. 

New Haven, CN: Yale University School of Forestry 

and Environmental Studies. 40 p. 

Liogier, Henri Alain; Martorell, Luis F. 1982. Flora of 

Puerto Rico and adjacent islands: a systematic 

synopsis. Rio Piedras, PR: Editorial de la Universidad de Puerto Rico. 342 p. 

Little, Elbert L., Jr.; Wadsworth, Frank H. 1989. 

Common trees of Puerto Rico and the Virgin Islands. 2nd printing. Reprinted privately by the 

authors from Agric. Handb. 249 [1964]. Washington, DC: U.S. Department of Agriculture. 556 p. 

Little, Elbert L., Jr.; Woodbury, Roy 0. 1976. Trees of 

the Caribbean National Forest, Puerto Rico. Res. 

Pap. ITF-20. Rio Piedras, PR: U.S. Department of 

Agriculture, Forest Service, Institute of Tropical 

Forestry. 27 p. 

Little, Elbert L., Jr.; Woodbury, Roy 0.; Wadsworth, 

Frank H. 1974. Trees of Puerto Rico and the Virgin 

Islands. Second volume. Agric. Handb. 449. 

Washington, DC: U.S. Department of Agriculture. 

1,024 p. 

Lugo, A.E.; Scatena, F.N. [In pressl. Ecosystem-level 

properties of the Luquillo Experimental Forest with 

emphasis on the tabonuco forest. In: Lugo, A.E.; 

Lowe, C., eds. A century of tropical forestry research: results from the first half, plans for the 

second. New York: Elsevier. 

Migenis, Luis E.; Ackerman, James D. [In press]. Orchid-phorochyte relationships in a forest watershed 

in Puerto Rico. Journal of Tropical Ecology. 

Odum, Howard T.; Pigeon, Robert F., eds. 1970. A 

tropical rain forest: a study of irradiation and ecology at El Verde, Puerto Rico. Springfield, VA: 

National Technical Information Service (U.S. 

Atomic Energy Commission): [Chapters paginated 

separately]. 

Perez, Ivette E. 1988. Field guide to the trees of Bisley 

watersheds, Luquillo Experimental Forest, Puerto 

Rico. Rio Piedras, PR: U.S. Department of Agriculture, Forest Service, Institute of Tropical Forestry. 

89 p. 

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

Virgin Islands. Memoirs of the New York Botanical 

Garden. Bronx; NY: The New York Botanical Garden. 389 p. Vol. 53. 

Richards, P.W. 1966. The tropical rain forest: an 

ecological study. Cambridge MA: Cambridge University Press. 450 p. 

Sastre de Jesus, Ines; Buck, William R. [In press]. Annotated checklist of mosses of Puerto Rico. Caribbean Journal of Science. 

Scatena, F.N., Larsen, M.C. 1991. Physical aspects of 

Hurricane Hugo in Puerto Rico. Biotropica. 23(4a): 

317-323. 

Scatena, Frederick N. 1989. An introduction to the 

physiography and history of the Bisley Experimental Watersheds in the Luquillo Mountains of Puerto 

Rico. Gen. Tech. Rep. SO-72. New Orleans, LA: U.S. 

Department of Agriculture, Forest Service, 

Southern Forest Experiment Station. 22 p. 

van der Wijk, R.; Margadant, W.D.; Florschutz, P.A. 

1959-69. Index Muscorum; Regnum Vegetabile. 

Utrecht, Netherlands: Association of Plant 

Taxonomy. 5 vol. 
Chinea, Jesus Danilo; Beymer Ren6e J.; Rivera, Carlos, Sastre de 

Jeses, Ines; Scatena, F.N. 1993. An annotated list of the flora of 

the Bisley area, Luquillo Experimental Forest, Puerto Rico, 1987 

to 1992. Gen. Tech. Rep. SO-94. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment 

Station. 12 p. 

Known species of plants, including bryophytes and ferns, are listed 

for the area of the Bisley experimental watershed area, a subtropical wet forest in the Luquillo Mountains of northeastern Puerto 

Rico. 

Keywords: Subtropical wet forest, tabonuco forest, watershed. 

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