A survey of the literature and collections in the Monteverde Cloud Forest Preserve and adjacent pastureland yielded a total of 198 epiphytic bryophyte taxa (120 species of hepatics in 50 genera, 77 species of moss in 48 genera, and 1 hornwort): 178 in the primary forest, 63 in the secondary forest, and 84 in the pastureland

Gradstein, Stephan Robbert

Merwin, Mark C.

Nadkarni, Nalini M.

English

Hochschulschriftenserver - Universität Frankfurt am Main

63TROPICAL BRYOLOGY20 (2001)Epiphytic bryophytes of Monte VerdeTropical Bryology 20: 63-70, 2001Epiphytic bryophytes of Monteverde, Costa RicaMark C. Merwin1, S. Robbert Gradstein2, and Nalini M. Nadkarni1, 31The Evergreen State College, Olympia, Washington 98505 U.S.A.2Institute of Plant Sciences, University of Göttingen, Untere Karspüle 2, 37073 Göttingen,Germany3Author for correspondence: nadkarnn@elwha.evergreen.eduAbstract:  A survey of the literature and collections in the Monteverde Cloud Forest Preserve andadjacent pastureland yielded a total of 198 epiphytic bryophyte taxa (120 species of hepatics in 50genera, 77 species of moss in 48 genera, and 1 hornwort): 178 in the primary forest, 63 in thesecondary forest, and 84 in the pastureland.Tropical montane forests support atremendous abundance and diversity of epiphyticplants.  Historically, vascular epiphytes havereceived more attention from researchers thanepiphytic non-vascular plants (Nadkarni et al.2001), and bryophytes in the tropics are generallypoorly studied (Gradstein et al. 2001 ).  TheMonteverde Cloud Forest Reserve (MCFR) is oneof one of the most studied Neotropical montanecloud forests (Nadkarni and Wheelwright 2000),and the bryoflora of MCFR is fairly well known(Reed and Robinson 1971; Gradstein 2000;Morales 2000). Epiphytic bryophytes ofMonteverde have also been the focus of severalecological investigations concerning epiphyticbryophytes (Nadkarni 1984; Monge-Nájera1989; Nadkarni and Matelson 1989, 1992;Ingram and Nadkarni 1993; Matelson et al. 1993;Clark 1994; Sillett et al. 1995; Clark et al. 1998a,b; Gradstein 2000; Morales 2000; Nadkarni 2000;Nadkarni et al. 2000a,b; Gradstein et al. in press).In this paper, we bring together data thatconcern work on epiphytic bryophyte speciesdiversity across a gradient of human-induceddisturbance (i.e., primary forest, 35 year-oldsecondary forest, and pastureland).  First, a listof epiphytic bryophytes known to occur inprimary forest and secondary forest of the MCFRand adjacent pastureland was compiled from theliterature (Sillett et al. 1995; Gradstein et al. inpress).  We supplemented this with collectionsinvolving destructive sampling of trees in theprimary forest and secondary forest.  In primaryforest, species recorded as growing epiphyticallyon the trunk bases (0-1 m above the ground),shrubs, and treelets in the forest understory by64TROPICAL BRYOLOGY 20  (2001)Mervin et al.Gradstein et al. (in press) were excluded unlessthe taxa were also reported as growing <3 mabove the ground on tree trunks and/or onbranches in the canopy by Sillett et al. (1995) orMerwin and Nadkarni (unpubl. data).A total of 198 epiphytic bryophytespecies (120 hepatics, 77 mosses, 1 hornwort)have been recorded: 178 species in the primaryforest (111 hepatics, 66 mosses, and 1 hornwort),66 species in the secondary forest (32 hepatics,31 mosses), and 84 species in the pastureland (47hepatics, 37 mosses) (Table 1).  Most of themosses of Monteverde are widespread species,but hepatics include several uncommon ones notpreviously recorded from Costa Rica (Gradsteinet al., 1994), including Adelanthus carabyensis,Bazzania affinis, Calypogeia crenulata (=Mnioloma crenulata), Colura ulei, Frullanialaxiflora, Lophocolea connata, Marchesiniarobusta, Plagiochila deflexirama, P.rudischusteri, Prionolejeunea schlimiana,Radula antillana, R. tenera, and Syzygiellapectiniformis. The Monteverde Cloud Forest isalso one of the few localities of the rare CostaRican endemic liverwort Calypogeiarhynchophylla (= Mnioloma rhynchophylla),known otherwise from only two localities on themailand and from Cocos Island (Dauphin 1999).The rare endemic Nowellia reedii Robins.,described from Monteverde (exact localityunknown) and not recorded anywhere else, wasnot found during this study.Study AreaResearch was carried out in the MonteverdeCloud Forest Reserve (MCFR) (10°18’N, 84°48'W, elevation ca. 1500 m), in the Cordillera deTilarán, Costa Rica.  The forest of MCFR isclassified as tropical lower montane wet forestin the Holdridge Life Zone System (sensuHoldrige 1967) and further described as LeewardCloud Forest by Lawton and Dryer (1980).Average annual rainfall is 2,000 - 2,500 mm y-1,with an additional 20% contributed from mist(Clark 1994).  There are three seasons: wet-mistyseason (November - January), dry season(February- April), and wet season (May -October).  Detailed descriptions of the climate,geology, and vegetation of Monteverde are inNadkarni and Wheelwright (2000).  Fieldworkwas conducted in permanent study plots locatedin the Research Area of MCFR and adjacentpastureland.Acknowledgments: We thank Gregorio Dauphin,Ingo Holz, Zach Magombo, Maria Morales, MichellePrice, and Ronald Pursell for assisting withidentifications.  Special thanks to Bruce Allen, SteveChurchill, and Elena Reiner Drehwald for makingnumerous identifications. Rigo Solano, JeremyDonaghue, Sylvia Englund, Leah Lessen, RobertLücking, Gretchen Vos and Steve Yanoviak assistedwith the fieldwork.  The first author thanks the staff ofthe herbaria of the University of Göttingen (GOET),Missouri Botanical Garden (MO), and Universidád deCosta Rica (USJ) for assistance during research visits.Vouchers were deposited in USJ. Research wassupported by National Science Foundation grants (BIR9975110, DEB 9974035, INT 9981531, DEB9615341).ReferencesClark, K. L. 1994. The role of epiphytic bryophytesin the net accumulation and cycling of nitrogen ina tropical montane cloud forest. Ph.D. dissertation,University of Florida, Gainesville, Florida.Clark, K. L., N. M. Nadkarni, and H. L. Gholz.1998a. Growth, net production, litterdecomposition, and net nitrogen accumulation byepiphytic bryophytes in a tropical montane forest.Biotropica 30:12-23.Clark, K. L., N. M. Nadkarni, D. Schaefer, and H.L. Gholz. 1998b. Atmospheric deposition and netretention of ions by the canopy in a tropicalmontane forest, Monteverde, Costa Rica.  Journalof Tropical Ecology 14:27-45.Dauphin, G. 1999. Bryophytes of Cocos Island, CostaRica: diversity, biogeography and ecology. RevistaBiologica Tropicál 47:309-328.Holdridge, L. 1967. Life zone ecology. TropicalScience Center, San José, Costa Rica.Gradstein, S. R. 2000. Bryophytes. In: N. M. Nadkar-ni & N. T. Wheelwright (eds.) Monteverde: eco-logy and conservation of a tropical montane fo-rest, pp. 78-79. Oxford University Press, NewYork, New York.Gradstein, S. R., S. P. Churchill, N. Salazar Allen.2001.  A Guide to the Bryophytes of TropicalAmerica (Memoirs of the New York BotanicalGarden, Vol. 86).  New York Botanical Garden,Bronx, New York.Gradstein, S. R., D. Griffin III, M. I. Morales, and65TROPICAL BRYOLOGY20 (2001)Epiphytic bryophytes of Monte VerdeN. M. Nadkarni (in press).  Diversity and habitatdifferentiation of mosses and liverworts in thecloud forest of Monteverde, Costa Rica. Caldasia.Gradstein, S. R., A. Lücking, M. I. Morales & G.Dauphin. 1994. Additions to the hepatic flora ofCosta Rica. Lindbergia 19:73-86.Ingram, S. W., and N. M. Nadkarni. 1993.Composition and distribution of epiphytic organicmatter in a Neotropical cloud forest, Costa Rica.Biotropica 25:370-383.Lawton, R. O., and V. Dryer. 1980.  The vegetationof the Monteverde Cloud Forest Reserve. Brenesia8:101-116.Matelson, T. J., N. M. Nadkarni, and J. T. Longino.1993. Survivorship of fallen epiphytes in aneotropical cloud forest, Monteverde, Costa Rica.Ecology 74:265-269.Monge-Nájera, J. 1989. The relationship ofepiphyllous liverworts with leaf characteristics andlight in Monteverde, Costa Rica. Cryptogamie,Bryologie, Lichénologie 10:345-352.Morales, M. 2000. Plants growing on living leaves.In N. M. Nadkarni and N. T. Wheelwright (Eds.).Monteverde:  ecology and conservation of atropical cloud forest, pp. 80-81. Oxford UniversityPress, New York, New York.Nadkarni, N. M. 1984. Epiphyte biomass and nutrientcapital of a neotropical elfin forest. Biotropica16:249-256.Nadkarni, N. M. 2000. Colonization of strippedbranch surfaces by epiphytes in a lower montanecloud forest, Monteverde, Costa Rica. Biotropica32:358-363.Nadkarni, N. M., and T. J. Matelson. 1989. Birduse of epiphyte resources in Neotropical trees.Condor 91:891-907.Nadkarni, N. M. & T. Matelson. 1992.  Biomassand nutrient dynamics of epiphyte litterfall in aneotropical montane forest, Costa Rica.  Biotropica24:24-30.Nadkarni, N. M. and N. T. Wheelwright (eds.).2000. Ecology and natural history of a tropicalmontane cloud forest, Monteverde, Costa Rica.Oxford University Press, New York, New York.Nadkarni, N. M., A. R. Cobb, and R. Solano. 2000a.Interception and retention of macroscopicbryophyte fragments by branch substrates in atropical cloud forest: an experimental anddemographic approach. Oecologia 122:60-65.Nadkarni, N. M., R. O. Lawton, K. L. Clark,  T. J.Matelson,  and D. A. Schaefer. 2000b.Ecosystem ecology and forest dynamics. In N. M.Nadkarni and N. T. Wheelwright (eds.).Monteverde:  ecology and conservation of atropical cloud forest, pp. 303-350. OxfordUniversity Press, New York, New York.Nadkarni, N.M., M.C. Merwin, and J. Nieder. 2001.Forest canopies, plant diversity. In S. Levin (Ed.).Encyclopedia of Biodiversity. Vol. 3. pp. 27-40.Academic Press, San Diego, California.Reed, C. F., and H. Robinson. 1971. Bryophytes ofMonteverde, Costa Rica. Phytologia 21:6-21.Sillett, S. C., S. R. Gradstein, and D. Griffin III.1995. Bryophyte diversity of Ficus tree crownsfrom cloud forest and pasture in Costa Rica.Bryologist 98:251-260.66TROPICAL BRYOLOGY 20  (2001)Mervin et al.TABLE 1.  The epiphytic bryophyte taxa found in primary and secondary forest in MonteverdeCloud Forest Reserve and adjacent pastureland following Sillett et al. (1995), Gradstein et al. (inpress), and M. Merwin and N. Nadkarni (unpubl.). 1 = Primary forest, 2 = Secondary forest, 3 =Pastureland.1 2               3HEPATICAEAdelanthus carabayensis (Mont.) Grolle xAdelanthus decipiens (Hook.) Mitt. xAdelanthus pittieri (Steph.) Grolle xAmphilejeunea reflexistipula (Lehm. & Lindenb.) Gradst. x xAnoplolejeunea conferta (Meissn.) Schiffn. x xBazzania affinis (Lindenb. & Gott.) Trevis. xBazzania denticulata (Lindenb. & Gott.) Steph. xBazzania gracilis (Hampe & Gott.) Steph. x xBazzania hookeri (Lindenb.) Trevis. xBazzania longa (Nees) Trevis. xBazzania longistipula (Lindenb.) Trevis. x xBazzania stolonifera (Sw.) Trevis. xBlepharolejeunea saccata (Steph.) van Slag. & Kruijt xBrachiolejeunea laxifolia (Tayl.) Schiffn. xBryopteris filicina (Sw.) Nees x x xCephalozia crassifolia (Lindenb. & Gott.) Fulford x x xCeratolejeunea cornuta (Lindenb.) Steph. x x xCeratolejeunea filaria (Tayl. ex Lehm.) Steph. x x xCeratolejeunea patentissima (Hampe & Gott.) Evans xCheilolejeunea adnata (Kunze) Grolle x xCheilolejeunea inflexa (Lehm.) Grolle x xCheilolejeunea rigidula (Mont.) Schust. x x xCheilolejeunea trifaria (Reinw. et al.) Mizut. xColura ulei Jov.-Ast. xCyclolejeunea convexistipa (Lehm. & Lindenb.) Evans xCyclolejeunea luteola (Spruce) Grolle xCyclolejeunea peruviana (Lehm. & Lindenb.) Evans xCyrtolejeunea holostipa (Spruce) Evans xDicranolejeunea axillaris (Nees & Mont.) Schiffn. xDiplasiolejeunea alata Jov.-Ast. xDiplasiolejeunea cavifolia Steph. xDiplasiolejeunea johnsonii Evans xDiplasiolejeunea pellucida (Meissn. ex Spreng.) Schiffn. xDrepanolejeunea cf. bidens Steph. x xDrepanolejeunea inchoata (Meissn.) Evans xDrepanolejeunea lichenicola (Spruce) Steph. xEchinocolea dilatata (Evans) Schust. xFrullania apiculata (Reinw. et al.) Nees x x xFrullania arecae (Spreng.) Gott. x x67TROPICAL BRYOLOGY20 (2001)Epiphytic bryophytes of Monte VerdeFrullania brasiliensis Raddi x x xFrullania convoluta Lindenb. & Hampe x xFrullania kunzei Lehm. & Lindenb. xFrullania laxiflora Spruce xFrullania riojanerirensis (Raddi) Ångstr. x xHarpalejeunea cf. stricta (Lindenb. & Gott.) Steph. xHarpalejeunea cinchonae (Nees) Schiffn. x xHerbertus divergens (Steph.) Herz. x xHerbertus juniperoideus (Sw.) Grolle x x xHerbertus pensilis (Tayl.) Spruce xJamesoniella rubricaulis (Nees) Grolle xJubula bogotensis Gott. xKurzia capillaris (Sw.) Grolle xLejeunea cerina (Lehm. & Lindenb.) Gott.et al. xLejeunea cf. caespitosa Lindenb. xLejeunea cf. filipes Spruce xLejeunea controversa Gott. x xLejeunea flava (Sw.) Nees x xLejeunea laetevirens Nees & Mont. x xLejeunea phyllobola Nees & Mont. xLepidolejeunea involuta (Gott.) Grolle x xLepidozia armata Steph. xLepidozia cupressina (Sw.) Lindenb. x x xLepidozia muenchiana Steph. x xLepidozia squarrosa Steph. xLeptoscyphus porphyrius (Nees) Grolle x xLeucolejeunea xanthocarpa (Lehm. & Lindenb.) Evans x xLophocolea muricata (Lehm.) Nees xLophocolea trapezoidea Mont. xLopholejeunea subfusca (Nees) Schiffn. xMarchesinia brachiata (Sw.) Schiffn. x x xMarchesinia robusta (Mitt.) Schiffn. xMetzgeria albinea Spruce x x xMetzgeria aurantiaca Steph. xMetzgeria decipiens (Mass.) Schiffn. x x xMetzgeria leptoneura Spruce x x xMetzgeria liebmanniana Lindenb. & Gott. xMicrolejeunea acutifolia Steph. xMicrolejeunea bullata (Tayl.) Evans x xNeurolejeunea breutelii (Gott.) Evans x x xOdontolejeunea lunulata (Web.) Schiffn. xOdontoschisma longiflorum (Tayl.) Steph. xOmphalanthus filiformis (Sw.) Nees x x xOmphalanthus grandistipulus Steph. xOmphalanthus ovalis (Lindenb. & Gottsche) Gradst. x x xPallavicinia lyelli Hook. xPlagiochila adiantoides (Sw.) Lindenb. x x x68TROPICAL BRYOLOGY 20  (2001)Mervin et al.Plagiochila aerea Tayl. x x xPlagiochila bidens Gott. xPlagiochila cristata (Sw.) Lindenb. xPlagiochila deflexirama Tayl. xPlagiochila diversifolia Lindenb. & Gott. x xPlagiochila gymnocalycina (Lehm. & Lindenb.) Lindenb. x x xPlagiochila laxa Lehm. & Lindenb. xPlagiochila micropteryx Gott. x xPlagiochila miqueliana Lehm. & Lindenb. xPlagiochila patula (Sw.) Lindenb. xPlagiochila raddiana Lindenb. x x xPlagiochila stolonifera Lindenb. & Gott. x xPlagiochila stricta Lindenb. xPlagiochila subplana Lindenb. xPlagiochila tenuis Lindenb. xPorella swartziana (Web.) Trevis. xRadula antilleana Castle xRadula frondescens Steph. xRadula gottscheana Tayl. x xRadula javanica Gott. x xRadula tenera Mitt. ex Steph. x xRiccardia fucoidea (Sw.) Schiffn. xScapania portoricensis Hampe & Gott. xSymbiezidium barbiflorum (Lindenb. & Gott.) Evans x xSymbiezidium transversale (Sw.) Trevis var. hookerianum    (Gott. et al.) Gradst. & van Beek xSymphyogyna brasiliensis Nees xSymphyogyna brogniartii Mont. xSyzygiella pectiniformis Spruce xTaxilejeunea pterigonia (Lehm. & Lindenb.) Schiffn. xTelaranea nematodes (Aust.) Howe xTrichocolea flaccida (Spruce) Jack & Steph. xTrichocolea tomentosa (Sw.) Gott. x x xTylimanthus cf. approximatus (Lindenb.) Besch. xTylimanthus laxus Spruce xANTHOCEROTAEDendroceros crispus (Sw.) Nees xMUSCIAcroporium estrellae (Müll. Hal.) Buck & Schäf.-Verw. x xAcroporium pungens (Hedw.) Broth. x xActinodontium standleyi Bartr. x x xAmphidium tortuosum (Hornsch.) Robins. xBrachymenium sp. x x69TROPICAL BRYOLOGY20 (2001)Epiphytic bryophytes of Monte VerdeBryohumbertia filifolia (Hornsch.) J.-P. Frahm xBryum capillare Hedw. xCampylium praegracile (Mitt.) Broth. xCampylopus arctocarpus (Hornsch.) Mitt. x xCampylopus densicoma (Müll. Hal.) Paris xCampylopus flexuosus (Hedw.) Brid. xCampylopus nivalis (Brid.) Brid. x x xCampylopus savannarum (Müll. Hal.) Mitt. xCaribaeohypnum polypterum (Mitt.) Ando & Higuchi x xDaltonia gracilis Mitt. x xDaltonia longifolia Tayl. x xEctropothecium leptochaeton (Schwägr.) Buck. xFissidens lagenarius Mitt. var. lagenarius xGroutiella apiculata (Hook.) Crum & Steere xGroutiella chimborazensis (Spruce ex Mitt.) Florsch. x xHerzogiella cylindricarpa (Card.) Iwats. xHolomitrium arboreum Mitt. x xHolomitrium pulchellum Mitt. xHypnella diversifolia (Mitt.) Jaeg. xHypnella pallescens Herz. xIsodrepanium lentulum (Wils.) Britt. x x xLeiomela bartramioides (Hook.) Par. xLepidopilum falcatulum Müll. Hal. x xLepidopilum muelleri (Hampe) Spruce xLepidopilum scabrisetum (Schwägr.) Steere x xLeptotheca boliviana Herzog x xLeucobryum antillarum Besch. x x xLeucobryum giganteum Müll. Hal. xLeucoloma cruegerianum (Müll. Hal.) Jaeger x x xLeucoloma serrulatum Brid. x x xMacromitrium cf. tonduzii  Ren. & Card. xMacromitrium cirrosum (Hedw.) Brid. x xMacromitrium parvirete Bartr. x xMacromitrium podocarpi Müll. Hal. x x xMacromitrium richardii Schwägr. xMeteoridium remotifolium (Müll. Hal.) Manuel x x xMeteorium illecebrum Sull. xMittenothamnium lehmanni (Besch.) Card. xMittenothamnium reptans (Hedw.) Card. x xOrthodontium pellucens (Hook.) B.S.G. xOrthostichella pentasticha (Brid.) Buck x xPalamocladium leskoides (Hook.) Britt. x xPapillaria deppei (Hornsch. ex Müll. Hal.) Jaeg. xPapillaria imponderosa (Tayl.) Broth. xPhyllogonium fulgens (Hedw.) Brid. x x xPhyllogonium viscosum (P. Beauv.) Mitt. x x xPilotrichella flexilis (Hedw.) Ångstr. x x70TROPICAL BRYOLOGY 20  (2001)Mervin et al.Porotrichodendron superbum (Tayl.) Broth. xPorotrichum cf. guatemalense Bartr. x xPorotrichum korthalsianum (Dozy & Molk.) Mitt. x x xPorotrichum longirostre (Hook.) Mitt. xPrionodon densus (Hedw.) Müll. Hal. x x xPrionodon fuscolutescens Hampe xPterobryon densum Hornsch. x xPyrrhobryum spiniforme (Hedw.) Mitt. x xRhegmatodon polycarpus (Griff.) Mitt. x xRhizogonium lindigii (Hampe) Mitt. xRhynchostegium serrulatum (Hedw.) Jaeg. xSchlotheimia rugifolia (Hook.) Schwägr. x xSematophyllum subsimplex (Hedw.) Mitt. x xSquamidium isocladum (Ren. & Card.) Broth. xSquamidium leucotrichum (Tayl.) Broth. xSquamidium livens (Schwägr.) Broth. x x xSquamidium nigricans (Hook.) Broth. xSyrrhopodon gaudichaudi Mont. x xSyrrhopodon incompletus Schwägr. x x xSyrrhopodon lycopodioides (Sw. ex Brid.) Müll. Hal. x xSyrrhopodon prolifer Schwägr. x x xThuidium delicatulum (Hedw.) B.S.G. x xZelometeorium allionii Manuel xZelometeorium patulum (Hedw.) Manuel xZygodon liebmanii Schimp. x         SynonymsCampylium hispidulum (Brid.) Mitt. = Campylium praeglaciale (Mitt.) Broth.Frullania exilis Tayl. = Frullania apiculata (Reinw. et al.) NeesFrullania neesii Lindenb. = Frullania kunzei Lehm. & Lindenb.Macromitrium mamillosum Bartr. = Macromitrium cirrosum (Hedw.) Brid.Macromitrium portoricense Williams = Macromitrium podocarpi Müll.Plagiochila acanthoda Lindenb. & Gott. = Plagiochila stricta Lindenb.Plagiochila bursata (Desv.) Lindenb. = Plagiochila aerea Tayl.Plagiochila guilleminiana Nees & Mont. = Plagiochila raddiana Lindenb.Porotrichum mutabile Hampe = Porotrichum longirostre (Hook.) Mitt.Prionodon luteovirens (Tayl.) Mitt. = Priondon densus (Hedw.) Müll.Radula macrostachya Lindenb. & Gott. = Radula javanica Gott.Schlotheima jamesonii (Arnott) Brid. = Schlotheimia rugifolia (Hook.) Schwägr.Thuidium antillarum Besch. = Thuidium tomentosum Besch..         

A Guide to the Bryophytes of Tropical

Additions to the hepatic flora of Costa Rica. Lindbergia

Atmospheric deposition and net retention of ions by the canopy in a tropical montane forest, Monteverde, Costa Rica.

Biomass and nutrient dynamics of epiphyte litterfall in a neotropical montane forest, Costa Rica.

Bird use of epiphyte resources in Neotropical trees.

Bryophyte diversity of Ficus tree crowns from cloud forest and pasture in Costa Rica.

Bryophytes of Cocos Island, Costa Rica: diversity, biogeography and ecology.

Colonization of stripped branch surfaces by epiphytes in a lower montane cloud forest, Monteverde, Costa Rica.

Composition and distribution of epiphytic organic matter in a Neotropical cloud forest, Costa Rica.

Ecology and natural history of a tropical montane cloud

Ecosystem ecology and forest dynamics.

Epiphyte biomass and nutrient capital of a neotropical elfin forest.

Epiphytic bryophytes of Monte Verde

Forest canopies, plant diversity. In

Growth, net production, litter decomposition, and net nitrogen accumulation by epiphytic bryophytes in a tropical montane forest.

Interception and retention of macroscopic bryophyte fragments by branch substrates in a tropical cloud forest: an experimental and demographic approach.

Life zone ecology. Tropical Science Center,

Monteverde: ecology and conservation of a tropical montane forest,

Plants growing on living leaves. In

Survivorship of fallen epiphytes in a neotropical cloud forest, Monteverde, Costa Rica.

The epiphytic bryophyte taxa found in primary and secondary forest in Monteverde Cloud Forest Reserve and adjacent pastureland following Sillett

The relationship of epiphyllous liverworts with leaf characteristics and light in Monteverde,

The role of epiphytic bryophytes in the net accumulation and cycling of nitrogen in a tropical montane cloud forest.

The vegetation of the Monteverde Cloud Forest Reserve.

Epiphytic bryophytes of Monteverde, Costa Rica

Epiphytic bryophytes of Monteverde, Costa Rica

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