United States Department of Agriculture An Annotated List of the

Forest Service Flora of the Bisley Area, Southern Forest Luquillo Experimental Forest, Experiment Station

New Orleans, Puerto Rico Louisiana 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 Ex- perimental 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 comprise 60 percent of the aboveground biomass, 51 the species composition of various areas in Bis- percent of the basal area, 49 percent of the stem den- ley (Basnet 1990; Garcia Montiel 1991; Heaton and sity, and 57 percent of the importance value.' Details Letourneau 1989; Migenis and Ackerman, in press). of the ecology and history of the area have been pub- The most comprehensive species list of the Bisley lished elsewhere (Brown and others 1983; Garcia area was assembled by Perez (1988) a few years Montiel1991; Odum and Pigeon 1970; Scatena 1989). before Hurricane Hugo. Floristic differences between major habitats in the Bisley area have been noted. For example, abundance and diversity of introduced vascular decrease VEGETATION with distance from the roads, trails, and streams. In- troduced species are almost absent from the upper The forests of the Bisley area have been classified parts of the watersheds, which are the areas least af- as subtropical wet forests (Ewe1 and Whitmore 1973), fected by human disturbances. The abundance and lower montane rain forests (Beard 19441, and single- diversity of bryophytes also increase toward stream dominant forests (Richards 1966). The Bisley water- banks, a trend similar to the observed by Churchill (in sheds are covered by a secondary forest of the press) in Ecuador. tabonuco type ( 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 Iscatena, F.N.; Lugo A.E. 1n.d.l Natural disturbances and the the watersheds-D. excelsa, Sloanea berteriana vegetation in two subtropical wet steepland watersheds of Puerto Choisy, and Prestoea n~ontarza(R. Grah.) Nichols.- Rico. Illanuscript has been submitted to the "Journal of Ecology." Two growth forms dominate the bryophyte flora of the major growth forms are represented by 107 Bisley: cushions (e.g., Leucobryum, Octoblepharum, species of , 20 species of shrubs, 28 species of Syrrhopodon, and Calymperes) and mats (e.g., dicotyledonous vines, 86 species of herbs, and 43 Sematophyllum, Callicostella, Isopterygium, and species of ferns. Lejeunaceae) (Sastre de Jesus and Buck, in press). Ninety-three percent of the listed species are na- Pendent bryophytes are poorly represented. tives of Puerto Rico; only 22 species (all of them an- giosperms) are introduced species (table 3). Thirty of the 314 native species (mostly dicotyledones) are en- STATISTICAL SUMMARY demic to Puerto Rico; these species make up 9 percent endemism among all plant species, or 11 percent en- The bryophyte flora of the Bisley area contains 52 demism among the vascular plants. The largest species in 40 genera and 21 families, while the vas- families of flora in the Bisley area, those with 10 cular flora contains at least 284 species in 215 genera species or more, are listed in table 4. 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 Table 4.- The 9 largest families of flora (each had 10 or more Angiospermae 73 188 species) in the Bisley area of the Luquillo Experimental Dicot herbs 29 Forest, Puerto Rico, 1987 to 1992 Monocot herbs 57 Dicot vines 28 Family Number of species Woody shrubs" 20 Trees* 107 Bryophyta Lejeuneaceae Total 102 255 336 Pteridophyta "Categories based on Britton and Wilson (1923, 1925), Little and Polypodiaceae others (1974), Little and Wadsworth (1989), as well as the authors' Dicotyledones field experience. Lauraceae Leguminosae Melastomataceae Rubiaceae Compositae Monocotyledones Table 3.-Number of native, endemic, and introduced species Cyperaceae 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 Total 314 30 22 336 "Endemics are a subgroup of native species. ANNOTATED LIST Family: Bryaceae

Nomenclature follows van der Wijk and others Bryum truncorum Brid. On riparian rocks. Ar- (1959-69) for , Gradstein (1989) for hepatics, chegonia observed in May. Proctor (1989) for ferns, Ackerman and Del Castillo (in press) for orchids, and Liogier and Martorell Family: Rhizogoniaceae (1982) for most other species. The annotations vary somewhat, reflecting the dif- Pyrrhobryum spiniforme (Hedw.) Mitt. On tree ferent collaborators' comments. The absence of an an- trunks. Sporophytes produced in February. notation usually indicates that the species is uncom- mon or that it has not been observed after the original collection. Family: Bartramiaceae

Philonotis elongata (Dism.) Crum et Steere. On wet soil, rocks, rarely on logs; also on soil of uprooted Division: BRYOPHYTA trees. Class: MUSCI (Mosses) Family: Orthotrichaceae

Family: Fissidentaceae Groutiella apiculate (Hook.) Crum et Steere. On logs. Sporophytes observed in April. Fissidens inaequalis Mitt. Common in clay soil, mineralized clay. F. mollis Mitt. Family: Neckeraceae F. repandus Wils. ex Mitt. On riparian rocks. F. zollegeri Mont. In mineralized clay. Neckeropsis disticha (Hedw.) Kindb. On bases of tree trunks and logs. Sporophytes produced March to April. Family: Dicranaceae N. undulata (Hedw.) Reichdt. On bases of tree trunks, logs, and rocks. Dicranella perrottetii (Mont.) Mitt. On clay soil of uprooted trees. Leucoloma cruegerianum (C. Mull.) Jaeg. On tree Family: Callicostaceae trunks, rarely on logs. Callicosta evanescens (C. Mull.) Crosby. On logs in stream. Family: Leucobryaceae Callicostella depressa (Hedw.) Jaeg. Common on logs. Leucobrym martianum (Hornsch.) Hampe ex C. C. pallida (Hornsch.) hgstr. Mostly found on logs. Mull. On bases of tree trunks and logs. Sporophytes produced in February. Octoblepharum albidum Hedw. On bases of tree Crossomitrium patrisiae (Brid.) C. Mull. On logs in trunks and logs. shady areas. Cyclodictyon albicans (Hedw.) Broth. On logs. Family: Calymperaceae C. varians (Sull.) 0. Kuntze

Claymperes erosum C. Miill. On tree trunks and Family: Leucomiaceae logs. C. nicaraguense Ren. & Card. On logs. Leocomium strumosum (Hornsch.) Mitt. On logs and Syrrhopodon incompletus var. berteroanus (Brid.) rocks. Reese. On tree trunks and logs. S. lgulatus Mont. On logs. S. parasiticus var. parasiticus Florsch. On logs. 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. Sporo- phytes observed in February. Metzgeria uncigera Evans. On logs and rocks.

Family: Pallaviciniaceae

C1ass:HEPATICAE (hepatics) Pallavicinia lyellii (Hook.) S. Gray Order:

Family: Lepidoziaceae Division: PTERIDOPHYTA Bazzania schwanecheana (Hampe & Gott.) Trev. On (Ferns and fern-allies) logs.w Telaranea nematodes (Gott. ex Aust.) Howe. On logs and leaf litter. Family: Psilotaceae

Family: Geocalycaceae Psilotum nudum (L.) P. Beauv. Over bridge on Bis- ley Road at Bisley 3. Lophocolea bidentata (L.) Dum. On logs and rocks. Family: Lycopodlaceae Family: Lycopidium cernum L. Outside watersheds, along Plagiochila sp. roadsides.

Family: Lejeuneaceae Family: Selaginellaceae

Archilejeunea parviflora (Nees) Schiffn. On logs and Selaginella krugii Hieron. Entire area. Endemic. tree trunks. Ceratolejeunea patentissima (Hampe & Gott.) Family: Marattiaceae Evans. On logs- and tree trunks. C. valida Evans Cheilolejeunea rigidula (Nees ex Mont.) Schust. On Danaea elliptica J.E. Smith logs. D. nodosa (L.) J.E. Smith. Entire area. ~olurarynchophora 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. Trichomanes rigidum Sw. On high, shady slopes. M. bullata (Tayl.) Steph. Odorztolejeunea lunulata (Web.) Schiffn. Prionolejeunea innovata Evans Family: Cyatheaceae Division: EMBRYOPHSTA Class: ANGIOSPERMAE (flowering plants) Cyathea horrida (L.) J.E. Smith. Entire area. Subclass: DICOTYLEDONES C. arborea (L.)J.E. Smith. Entire area. C. borinquena (Maxon) Domin. Endemic. C. portoricensis Spreng. ex Kuhn. Entire area, ex- Family: Casuarinaceae cept sunny areas. Endomic. Casuarina equisetifolia J.R. & G. Forst. Along road- Family: Polypodiaceae sides. Introduced from Australia.

Adiantum latifolium Lam. Entire area. Family: Piperaceae A. pyramidale (L.) Willd. In the watersheds. Arachniodes chaerophylloides (Poir.) Proctor Lepianthes peltat um (L.) Rafinesque. Entire area. Asplenium auritum Sw. Peperomia spp. Several species in the watersheds. A. salicifolium L. Piper aduncum L. Along roadsides, in sunny areas. A. serratum L. In the watersheds. P. glabrescens (Miq.) C. DC. Entire area. Blechnum occidentale L. In the watersheds. P. hispidum Sw. Entire area. Bolbitis aliena (Sw.)Alston. Rare in the watersheds. P. jacquemontianum Kunth. Along roadsides, near B. nicotianifolia (Sw.) Alston. Very common in the the creek in Bisley 1. watersheds. Ctenitis subincisa (Willd.) Ching Dennstaedtia bipinnata (Cav.) Maxon. Entire area. Family: Chloranthaceae D. obtusifolia (Willd.) Moore. Entire area. Elaphoglossum sp. Entire area. Hedyosmum arborescens Sw. In the watersheds be- Hemidictym marginatum (L.) K. Presl. Inside and fore Hurricane Hugo. outside the watersheds; near creek beds. Hypolepis repens (L.) K. Presl. Common in sunny, Family: Ulmaceae disturbed areas. Lonchitis hirsuta L. Riparian. Trema micranthum (L.) Blume. Along roadsides; on Nephrolepis exaltata (L.) Schott high, disturbed slopes. Nephrolepis rivularis (Vahl) Mett. ex Krug. In the watersheds, along roadsides. Odontosoria aculeata (L.) J. Smith. Entire area, in Family: Moraceae drier areas. Pityrogramma calomelanos (L.) Link var. aureoflava Artocarpus altilis (S. Park.) Fosb. In drainages (Hooker) Weatherby ex Bailey along road. Introduced from islands of the South P. calomelanos (L.) Link var. calomelanos. Entire Pacific. area. Cecropia peltata L. Entire area. Polybotrya cervina (L.) Kaulf. Entire area, not com- Ficus citrifolia P. Miller. Inside and outside the mon. watersheds. Polypodium aureum L. P. crassifolium L. Upper slopes of watersheds. Family Urticaceae P. piloselloides L. In the watersheds, epiphytic. P. spp. Many epiphytic species. Pilea inaequalis (Juss. ex Poir.) Wedd. Entire area. Pteris altissima Poir. In the watersheds. P. krugii Urban. On upper slopes of Bisley 2. En- Tectaria trifoliata (L.) Cav. Entire area. demic. Thelypteris balbisii (Spreng.) Ching Pilea obtusata Liebm. T. deltoidea (Sw.) Proctor. Entire area. Urera baccifera L. Wedd. Entire area. T. reticulate (L.) Proctor. Sunny areas, roadsides. 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. Inga fagifolia (L.) Willd. Entire area. I. vera Willd. Entire area. Family: Magnoliaceae Mimosp pudica L. Along roadsides. Magnolia splendens Urban. In the watersheds, espe- cially on ridges. Endemic. Family: Leguminosae Subfamily: Caesalpiniodeae

Family: Annonaceae Cassia aeschinomene DC. Along roadsides. Guatteria car-ibaea Urban. In the watersheds. Family: Leguminosae Subfamily: Papilionoideae Family: Lauraceae Andira inermis (W. Wright) HBK. Entire area. Aniba bracteata (Nees) Mez. Entire area. Desmodium adscendens (Sw.) DC. Beilschmiedia pendula (Sw.) Hemsl. In the water- - - Neorudolphia volubilies (Willd.) Britton. Entire sheds. area. Endemic. Licaria triandra (Sw.) Kostermans Ormosia krugii Urban. Entire area. Ocotea floribunda (Sw.) Mez. In the watersheds. Pterocarpus officinalis Jacq. Inside and outside the 0.giobosa (Aubl.) Schlecht. & Cham. watersheds. 0. LeucoxyZon (Sw.) Mez. Common in entire area. Pueraria phaseoloides (Roxb.) Benth. Along road- 0. membranacea (Sw.) Howard sides. Introduced from Asia. 0. moschata (Meissn.) Mez. In the watersheds. En- demic. 0.portoricensis Mez. In the watersheds. Endemic. Family: Rutaceae 0. sinienisii (Mez) Alain. In the watersheds. Persea americana Miller. Along roadsides. Intro- Citrus X paradisi Macfad. Along roadsides. Intro- duced from Mexico. duced from other islands of the West Indies. Ravenia urbanii Engler. In the watersheds, mostly on high ridges. Endemic. Family: Chrysobalanaceae Zanthoxylum rnartinicense (Lam.) DC. Entire area. HirteLLa rugosa Pers. Entire area. Endemic. H. triandra Sw.2 Observed in Bisley 1 before Hur- Family: ricane Hugo, but not found afterwards. Dacryodes excelsa Vahl. Entire area. Tetragastris balsamifera (Sw.) Kuntze. In the water- Family: Rosaceae sheds. Rubus rosifolius Smith. Along roadsides. Introduced from Southeast Asia. Family: Meliaceae

Guarea glabra Vahl. Entire area. Family: Connaraceae G. guidonia (L.) Sleumer. Entire area. Khaya nyasica Stapf. ex Baker. Entire area. Intro- Roz~reasurinamensis Miq. Entire area. duced from Africa. Swietenia macrophylla G. King. Entire area. Intro- duced from Mexico and Central America. Trichilia pallida Sw. Entire area.

Family: Malpighiaceae '~arcia,Diane. 1989. Personal communication (letter). On file with: International Institute of Tropical Forestry, U.S. Depart- Byrsonima spicata (Cav.) HBK. Entire area. ment of Agriculture, Forest Service, Rio Piedras, PR 00928. Heteropteris laurifolia (L.) A. Juss. Entire area. Family: Polygalaceae Family: Vitaceae

Securidaca uirgata Sw. Common in entire area. Cissus erosa L.C. Rich. Along roadsides. C. sicyoides L. Entire area. C. uerticillate (L.) Nicolson & Garvis. Along road- Family: Euphorbiaceae sides. Alchornea latifolia Sw. Entire area. Alchorneopsis floribunda (Benth.) Muell. Arg. En- Family: Elaeocarpaceae tire area. Croton poecilanthus Urban. Entire area. Endemic. Sloanea berteriana Choisy. Entire area. Drypetes glauca Vah. Entire area. Phyllanthus urinaria L. Along roadsides. Intro- Family: duced from Asia. Sapium laurocerasus Desf. Entire area. Endemic. Pauonia fruticosa (Miller) Fawc. & Rendle. Entire area. Family: Anacardiaceae Sida rhombifolia L. Along roadsides. Thespesia grandiflora DC. In the watersheds. En- Comocladia glabra (Schultes) Spreng. Entire area. demic. Magifera indica L. Along roadsides and in the Urena lobata L. Along roadsides. watersheds. Introduced from Asia. Spondim mombin L. In the watersheds. Family:

Family Cyrillaceae Ochroma lagopus Sw. Several individuals recruited after Hurricane Hugo in Bisley 2 near the road. Cyrilla racemiflora L. Reported by Perez (1988)but (Sw.) Poir. In the watersheds. found only in Bisley 3. Family: Dilleniaceae Family: Stahyleaceae Pinzona coriacea Mart. & Zucc. In the watersheds. Turpinia occidentalis (SW.) G. Don. In the water- sheds. Family: Ochnaceae

Family: Sapindaceae Sauvagesia erecta L. Entire area.

Cupania americana L. In the watersheds. Family Marcgraviaceae Matayba domingensis (DC.) Radlk. In the water- sheds. Marcgravia rectiflora Triana & Planch. Entire area. Paullinia pinnata L. Entire area. M. sintenisii Urban. Endemic.

Family: Sabiaceae Family: Theaceae

Meliosma herbertii Rolfe. Entire area. Laplacea portoricensis (-Krug & Urban) Dyer. In the watersheds. Family: Balsaminaceae Family: Guttiferae Impatiens wallerana H0ok.f. Along roadsides. Intro- duced from Zanzibar. Calophyllum brasiliense Jacq. In the watersheds. clusioides (Griseb.) D'Arcy. In the water- sheds. C. gundlachii Stahl. Entire area. Endemic. Rheediaportoricensis Urban. In the watersheds. En- demic. Family: Flacourtiaceae M. tetrandra (Sw.) D. Don Nepsera aquatica (Aubl.) Naud. Entire area. Casearia arborea (L.C. Rich.) Urban. Entire area. Tetrazygia urbanii Cogn. In the watersheds and C. guianensis (Aubl.) Urban along roadsides. Endemic. C. sylvestria Sw. Homalium racemosum Jacq. Family: Onagraceae Laetia procera (Poepp. & Endl.) Eichl. Entire area. Ludwigia octovaluis (Jacq.) Raven. Along roadsides Family: Passifloraceae and on sunny ridgetops.

Passiflora edulis Sims. Along roadsides. Introduced Family: Araliaceae from Brazil. P. rubra L. In the watersheds. Dendropanax arboreus (L.) Decne & Planch. Didymopanax morototoni (Aubl.) Decne. & Planch. Family: Begoniaceae Entire area.

Begonia decandra Pavon. In the watersheds. En- Family: Myrsinaceae demic. Parathesis crenulata (Vent.) Hook. f. Family: Lythraceae Wallenia pendula (Urban) Mez. In the watersheds. Endemic. Cuphea carthagenensis (Jacq.) Macbride C. strigulosa HBK. Family: Sapotaceae

Family: Rhizophoraceae Chrysophyllum argenteum Jacq. Reported before Hurricane Hugo but not seen again. Cassipourea guianensis Aubl. In the watersheds. Chrysophyllum cainito L. In Bisley 1. Manilkara bidentata (A. DC.) A. Chev. Entire area. Micropholis chrysophylloides Pierre Family: Combretaceae M. garciniifolia Pierre. Endemic. Buchenavia capitata (Vahl) Eichl. Inside and out- side the watersheds. Family: Oleaceae

Chionanthus domingensis Lam. Family: Myrtaceae

Calyptranthes pallens (Poir.) Griseb. Near water Family: Apocynaceae well field in lower end of Bisley 2. Eugenia eggersii Kiaersk. Endemic. Allanzanda cathartics L. Introduced from South E. stahlii (Kiaersk.) Krug & Urban. Endemic. America. Myrcia deflexa (Poir). DC. In the watersheds. Forsteronia portoricensis Woods. Endemic. M. leptoclada DC. In the watersheds. M. splendens (Sw.) DC. In the watersheds. Family: Convolvulaceae Syzgium jumbos (L.) Alst. Introduced from Asia. Ipomoea setifera Poir. Family: Melastomataceae I. tiliacea (Willd.) Choisy

Calycogonium squamulosum Cogn. Endemic. Family: Boraginaceae Henriettea fascicularis (Sw.) Gdmez Maza Heterotrichum cyrnosum (Wendl.) Urban. Endemic. Cordia borinquensis Urban. Endemic. Mzconia laevigata (L.) DC. C. sulcata DC. In the watersheds. M. prasina (Sw.) DC. In the watersheds. Tournefortia sp. In Bisley 2. M. racemosa (Aubl.) DC. Common in the entire area. M. serrulata (DC.) Naud. In the watersheds. Family: Verbenaceae Family: Compositae

Lamtana camara L. Along roadsides. Aster subulatus Michx. Along roadsides. Stachytarpheta jamaicensis (L.) Vah. Along road- Bidens alba (L.) DC. var. radiata (Sch.-Bip.) Bal- sides. lard. Along roadsides. divaricata Sw. Clibadium erosum (Sw.) DC. In the watersheds and along roadsides; in sunny, disturbed areas. Elephantopus mollis Kunth Family: Solanaceae E. spicatus Juss. ex Aubl. Eupatorium odoratum L. Cestrum macrophyllum Vent. Along roadsides. Mikania cordifolia (L.f.1 Willd. In the watersheds. Solanum torvum Sw. Entire area; in sunny places. M. fragilis Urban. In watersheds. Endemic. Neurolaena lobata (L.) Cass. Family: Scrophulariaceae Rolandra fruticosa.(L.) Kuntze Synedrella nodiflora (L.) Gaertn. Along roadsides. Bacopa stricta (Schrad.) Robins Vernonia cinerea (L.) Less. Wedelia trilobata (L.) A.S. Hitchc. Along roadsides. Family: Bignoniaceae

Tabebuia heteropylla (DC.) Britton. Entire area. Subclass: MONOCOTYLEDONES

Family: Acanthaceae Family: Gramineae Odontonema strictum (Nees) Kuntze. Introduced from Central America. Andropogon bicornis L. Ruellia coccinea (L.) Vahl. Entire area. A. leucostachyus HBK. Teliostachya alopecuroidea (Vahl) Nees. Along road- Bambusa vulgaris Schrad. ex Wendl. Along road- sides. sides. Introduced from Asia. Thunbergia alata Bojer. Introduced from Africa. Ichnanthus pallens (Sw.) Munro. Common in the watersheds. Olyra latifolia L. Fairly common after Hurricane Family: Rubiaceae Hugo; less common as canopy closes. Coffea arabica L. Introduced from Abyssinia. Panicum trichoides Sw. Paspalum conjugatum Berg. In the watersheds. Faramea occidentalis (L.) A. Rich. Gonzalagunia spicata (Lam.) Gomez Maza. Entire P. virgatum L. Along roadsides. area. Pharus latifolius L. In the watersheds. Hamelia patens Jacq. Hemidiodia ocimifolia (Willd. ex R. & S.) K. Schum. Family: Cyperaceae Hillia parasitica Jacq. In the watersheds. Ixora ferrea (Jacq.) Benth. Eleocharis interstincta (Vahl) R. & S. Along road- Palicourea crocea (Sw.) Roem. & Schult. var. crocea sides. P. crocea var. riparia (Benth.) Griseb. In the E. retroflexa (Poir.) Urban. In wet places along road- watersheds. sides. Psychotria berteriana DC. Common in entire area. Fimbristylis dichotoma (L.) Vahl. In wet places P. brachiata Sw. Entire area. along roadsides. P. maleolens Urban. Endemic. Fuirena umbellata Rottb. Along roadsides. Rodeletia portoricensis Krug & Urban. Endemic. Mariscus ligularis (L.) Urban. Entire area. Sabicea hirsuta HBK. Entire area. Rhynochospora nervosa (Vahl) Boeck. ssp. ciliata (Vahl) T. Koyama R. stellata (Lam.) Griseb. Along roadsides. Family: Cucuritaceae Scleria canescens Boeck. Entire area. Endemic. Cayaponia racemosa (Miller) Cogn. In the water- S. lithosperma (L.) Sw. sheds. S. pterota Presl. Family: Palmaceae Family: Zingiberaceae

Prestoea montana (R. Grahm) Nichols. Entire area. Zingiber sp. Along the creeks and roadsides. Intro- Roystonea borinquena O.F. Cook. Along roadsides. duced from Asia.

Family: Araceae Family: Cannaceae

Aglaonema pictum (Roxb.) Kunth. Introduced from Canna glauca Entire area. East Indies. Anthurium crenatum (L.) Kunth Family: Orchidaceae Dieffenbachia sp. May be an introduced species. Philodendron angustaturn Schott muscosa Sw. Terrestrial. P. scandens C. Koch & H. Sello Cyclopogon cranichoides (Griseb.) Schltr. Ter- Xanthosoma atrovirens C. Koch & Bouche. Intro- restrial. duced from South America. Encyclia cochleata (L.) Dressler. Epiphytic or lithophytic. Family: Commelinaceae Epidendrum carpophorum Barbosa Rodrigues. Epiphytic. Commelina diffusa Burm.f. In the watersheds and E. nocturnum Jacq. Epiphytic or occasionally along roasides. lithophytic. Commelinopsis persicariifolia (DC.) M. Pichon. In E. ramosum Jacq. Epiphytic. the watersheds. E. tridens poeppig & Endlicher. Epiphytic or oc- Zebrina pendula Schniz. Mostly along roadsides. casionally lithophytic. Erythrodes hirtella (Sw.) Fawc. & Rendle. Ter- restrial. Family: Pontederiaceae E. plantaginea (L.) Fawc. & Rendle. Terrestrial. Jacquiniella globosa (Jacq.) Schltr. Epiphytic. Heteranthera reniformis Ruiz & Pavon. In mud, Liparis nervosa (Thumb.) Lindl. Terrestrial. along roadsides. Maxillaria coccinea (Jacq.) L. 0. Williams ex Hodge. Common epiphyte, especially on Guarea. Family: Smilacaceae Pleurothallis ruscifolia (Jacq.) R. Br. Common epiphyte, especially on Guarea. Smilax domingensis Willd. Polystachya concreta (Jacq.) Garay & Sweet. S. havanensis Jacq. Common in the watersheds. Epiphytic or lithophytic. P. foliosa (Hook.) Rchb.f. Epiphytic or lithophytic. Prescottia oligantha (Sw.) Lindl. Terrestrial. Family: Scaphyglottis modestra (Rchb.f'.) Schltr. Epiphytic or lithophytic. decumbens L. In tire tracks in the road. Triphora latifolia G. Luer. Very rare terrestrial. Wullschlaegelia aphylla (Sw.) Rchb.f. Achlorphyl- Family: Dioscoreaceae lous, terrestrial.

Dioscorea polygonoides Humb. & Bonpl. ex Willd. LITEUTURE CITED Entire area. Dioscorea sp. On upper slope. With spines and Ackerman, James D.; Del Castillo, Maria. [In press]. bulbs. The orchids of Puerto Rico and the Virgin Islands: Rajania cordata L. Las orquideas de Puerto Ric0.y Las Islas Virgenes. Rio Piedras, PR: Editorial de la Universidad de Family: Musaceae Puerto Rico. Basnet, Khadga. 1990. Studies of ecological and geo- Heliconia sp. Entire area. logical factors controlling the pattern of tabonuco Musa spp. In the watersheds. Introduced from forests in the Luquillo Experimental Forest, Puerto India. Rico. New Brunswick, NJ: Rutgers, The State University of New Jersey. 167 p. Ph.D. dissertation. Beard, J.S. 1944. Climax vegetation in tropical Little, Elbert L., Jr.; Woodbury, Roy 0.; Wadsworth, America. Ecology. 25(2): 127-158. Frank H. 1974. Trees of Puerto Rico and the Virgin Britton, N.L.; Wilson, Percy. 1923. Scientific survey of Islands. Second volume. Agric. Handb. 449. Porto Rico and the Virgin Islands. New York: New Washington, DC: U.S. Department of Agriculture. York Academy of Sciences. 626 p. Vol. 5. 1,024 p. Britton, N.L.; Wilson, Percy. 1925. Scientific survey of Lugo, A.E.; Scatena, F.N. [In pressl. Ecosystem-level Porto Rico and the Virgin Islands. New York: New properties of the Luquillo Experimental Forest with York Academy of Sciences. 663 p. Vol. 6. emphasis on the tabonuco forest. In: Lugo, A.E.; Brown, Sandra; Lugo, Ariel E.; Silander, Susan; Lowe, C., eds. A century of tropical forestry re- Liegel, Leon. 1983. Research history and oppor- search: results from the first half, plans for the tunities in the Luquillo Experimental Forest. Gen. second. New York: Elsevier. Tech. Rep. SO-44. New Orleans, LA: U.S. Depart- Migenis, Luis E.; Ackerman, James D. [In press]. Or- ment of Agriculture, Forest Service, Southern chid-phorochyte relationships in a forest watershed Forest Experiment Station. 128 p. in Puerto Rico. Journal of Tropical Ecology. Churchill, Steven P. [In pressl. The mosses of Equa- Odum, Howard T.; Pigeon, Robert F., eds. 1970. A torian Amazonas. Aarhus, Denmark: University of tropical rain forest: a study of irradiation and ecol- Aarhus Botanical Institute AAU report. Vol. 24. ogy at El Verde, Puerto Rico. Springfield, VA: Ewel, J.J.; Whitmore, J.L. 1973. The ecological life National Technical Information Service (U.S. zones of Puerto Rico and the U.S. Virgin Islands. Atomic Energy Commission): [Chapters paginated Res. Pap. ITF-18. Rio Piedras, PR: U.S. Department separately]. of Agriculture, Forest Service, Institute of Tropical Perez, Ivette E. 1988. Field guide to the trees of Bisley Forestry. 72 p. watersheds, Luquillo Experimental Forest, Puerto Garcia Montiel, Diana. 1991. The effect of human ac- Rico. Rio Piedras, PR: U.S. Department of Agricul- tivity on the structure and composition of a tropical ture, Forest Service, Institute of Tropical Forestry. forest in Puerto Rico. Rio Piedras, PR: University of 89 p. Puerto Rico. 103 p. M.S. thesis. Proctor, George R. 1989. Ferns of Puerto Rico and the Gradstein, Stephan R. 1989. A key to the Hepaticae Virgin Islands. Memoirs of the New York Botanical and Anthocerotae of Puerto Rico and the Virgin Is- Garden. Bronx; NY: The New York Botanical Gar- lands. The Bryologist. 92: 329-348. den. 389 p. Vol. 53. Heaton, K.; Letourneau, A. 1989. Changes in forest Richards, P.W. 1966. The tropical rain forest: an structure and composition along a gradient from ecological study. Cambridge MA: Cambridge Uni- streams to ridges in a subtropical moist forest in versity Press. 450 p. Puerto Rico. Tropical Resource Institute report. Sastre de Jesus, Ines; Buck, William R. [In press]. An- New Haven, CN: Yale University School of Forestry notated checklist of mosses of Puerto Rico. Carib- and Environmental Studies. 40 p. bean Journal of Science. Liogier, Henri Alain; Martorell, Luis F. 1982. Flora of Scatena, F.N., Larsen, M.C. 1991. Physical aspects of Puerto Rico and adjacent islands: a systematic Hurricane Hugo in Puerto Rico. Biotropica. 23(4a): synopsis. Rio Piedras, PR: Editorial de la Univer- 317-323. sidad de Puerto Rico. 342 p. Scatena, Frederick N. 1989. An introduction to the Little, Elbert L., Jr.; Wadsworth, Frank H. 1989. physiography and history of the Bisley Experimen- Common trees of Puerto Rico and the Virgin Is- tal Watersheds in the Luquillo Mountains of Puerto lands. 2nd printing. Reprinted privately by the Rico. Gen. Tech. Rep. SO-72. New Orleans, LA: U.S. authors from Agric. Handb. 249 [1964]. Wash- Department of Agriculture, Forest Service, ington, DC: U.S. Department of Agriculture. 556 p. Southern Forest Experiment Station. 22 p. Little, Elbert L., Jr.; Woodbury, Roy 0. 1976. Trees of van der Wijk, R.; Margadant, W.D.; Florschutz, P.A. the Caribbean National Forest, Puerto Rico. Res. 1959-69. Index Muscorum; Regnum Vegetabile. Pap. ITF-20. Rio Piedras, PR: U.S. Department of Utrecht, Netherlands: Association of Plant Agriculture, Forest Service, Institute of Tropical . 5 vol. Forestry. 27 p. 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. Depart- ment 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 subtropi- cal wet forest in the Luquillo Mountains of northeastern Puerto Rico.

Keywords: Subtropical wet forest, tabonuco forest, watershed.

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