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(1)Rev. BioLTrop., 47(3): 309c328, 1999. www.ucr.ac.cr.. www.ots.ac.cr. www.ots:d\1ke.edu. Bryopbytes of COCOS Island, Costa Rica:. --, -----------------. diversify, biogeQgraphy and ecolog==y ---�----�- --. Gregorio Daupbin. Albrecht von HalIer Institute of Plarit ScieIices, Dept. of Systematic Botany, University of Goettingen, Untere , KarSlJUele 2, 0-37073 Goettingen, Germany. Fax 49551392329. gdauphi@gwdg.de ,Rec¡:i:ved l-VI�1998. Corrected 16.IT-1999. Accépted 24-11-1999. Abstraet: A total of 98liverWort specíes (43 ge¡1era, 10 families), 54 moss species (33 g�eia, 17 families) and one species of hornwort háve been reported for Cocos Island (5°32''N, 87°ü4'W), Costa.iuaa' : Over 60% of the , bryophytes have a Neotropi�al or Pantropical distribution; about 10% are Caribbe�, �d lessthan 5% are endernic or subeIidernic. In comparison to the Galapagos Archipelag�,.Cocos Island harDors_a more typical. ,1f?pical bryoflora with fqliose hepátics(e. g. Lejel1lleaceát;, Lepid,oziaceae) constitutirtgthe bu� of div,efSity;. fewef tha1los!,! liverwortS,-aud 11;1,088 taxa as in the Archipelagó werefound. A richer hábitat variety Í¡1cludi,niwet and dry habltats, as well asits bigger área"seemto account f?r the higher llumberofbryophytespecies in ' Galapagos Archipelago. Mkre6 4y:q; s hi'Cocos ISland'are crirl1coloí,ls (46%); the remainingare epiphyll\>us·,, (25%), S colOUS (23%) o),: , ',' (124): Bryophyte occurre¡1cein eightplotS(10 x 10 m) witb. -2Ó qua¡j}�1;e � (30 x 30 cm) were recorded, at.�nt habiiitt&, and ¡Jtiij,Jd�s(Ó:-6ÓtJ� .Bryophyte distributiol). y.áthin the isl�d­ coincides lowlandfo;esf; (0-10(¡ l. Séc()ritlary !QWI d.Í9re t 200; ) �d �ontane.fdrest (tci 6 00 m). ", � Physanthólejeunea port(Jrice,mis (Hampe & GQjt.) Schu�t:'is,�portea¡as¡1ewto �ostaRica. '. � \Vith. ". ". ,. >. -. ,. � �. -,. � �� '�,. _ ,. ". �. _ . '.. ' ,1. r. -: ';.,." ;'"j' , O': ,, , Key wor&: Bryopñytes,Costa Rica;' Cocos Island" G\l!.apago� Arcpipeta�\>" ísland'1Jt? geo�l?hy,epology. ,. >:�' ". '. ,. \. ,. ';�:�.'. Cocos Island i,s an :4Is�lar territory of Cos- '. ta Rica"494 ,km SW,o{Cabo Blanco, Nicoya PeIlÍnsula, 5°32'N, 87°04'W, (Castillo et al. 1988). Declared National Park sÍnce 1970 and included in the Unescp World Heritage List since 1997 (Montoya 199,Oa, Qu�sada 1997), theisland has an area of 24krn2 (Alvarado; 19;8�, H{!rrera l,986), and a rnaxÍp)um altitude of .600 Ip. (Fi�ure 1). The me¡m annual tempe� rawre �tO'mi� of 25.5 °g,!J11A .the mean a)lnual rajnfall raDge� from 5000 IiUn (ChathamBay) to 709Q:Wtn (Cerro Yglesias); the 4tY er mqnth��ire.betW¡::eu Janu� andMarcq. Cocos Island,is ll, yoqp.g vQlpano originated on a hot s po� 011 tlreCo�9s Pa l te, ."o/hose age is calcula­ �dl;>etw�,n .1.2 ¡md 2 milJions y�ar8 (Alvara:' � 19�4jtas tillq et al. 1988). Geologica11y, tP� .. ". _/'. '��'t. f-'. -",<!,. /. i. .'. island is composed of iJ.U(aline :hasalts, PYFo­ clasticrocks, trachyteand hawantes (Castillo et al. 1988). TIré island surface show'�; an irregular morphology;. being almost inaccessible from the shore" except at Wafer and Chatham bays. There. !Í�\acentral ¡p¡d easte)¡fl ar�a ofundu}ate topography, between 2QO.¡p¡d 260 m elevati,on, and V--shapeQ v;;;tlJ.eY/l (�yarado,,1989). The soutl;n�ystsidé óf the islal)d ,has a !}lore pyndent area,,:with �ost v�rtical,w�s tptbe sea. The, flopr ;qf Cocos I$J;ap.¡! has {:>eenirr,�gJl�. larly:'stu!iied, �Gómez',197��;,About 1 �9.species of flowering pl¡p¡ts hav� beeQ rC(port�d" otwhich.)2; at;e endemic (M:Ql1t()y� 1990b), l;b:e fenw ¡md fern allies have be�n stu!iied in detail byL�s Diego GÓQlez (1975twh,0 recOl:ded 74; species , inciudin� ;$,� endeliljc qnes. TIre. same,.

(2) 310. REVISTA DE BIOLOGÍA TROPICAL. 87103'. �. Fíg.1. Topography ofCocos Island and sites mentioned in texto. author also reported 85 species of basidiomy­ cetes (cited by Montoya 1990b). A vegetation analysis was performed by Gómez (1975), who recognized two elevatio­ nal belts: 1) the Littoral Zone (O-50m), located along the coast and on valleys, with two habi­ tat types: the Annona glabra swamp and the fmn terrain with various species of flowering plants. 2) the Mountainous Zone dominated by the trees Saccoglottis holdridgei, Ocotea insu­ laris, and Euterpe macrospadix with an un­ derstory of Hypolitrum amplum. Further vege-. tation types are recognized by Montoya (1990b). The byological exploration of Cocos Island is closely related with that of Galapagos Archi­ pelago, since many scientific expeditions to the Galapagos also landed on Cocos (Weber 1966). As in The Galapagos, bryological collecting on Cocos was at fust done by non-specialists, i.e. vascular plant collectors who also collected br­ yophytes. Early collections date from more than 200 years, when Menzies with the Vancouver Expedition collected the endemic Pilotrichum.

(3) DAUPHIN: Bryophytes ofCocos Island rugifolium C. M., from logs originating from Cocos Island (Crosby 1969). Pittier (1899) c1ai­ med to have been the first to do botanical co­ llecting on Cocos during an expedition of the Costa Rican Government in 1898. He collected many vascular plants as well as sorne crypto­ gams, inc1uding Pilotnchum corrugatum Ren. . & Cardo (Crosby 1969) andlIygrolijeunea oee: lIata, described by Stephani (Stephani 1914). Robinson (1902 cited by Clark 1953) reported the occurrence of Macrolejeunea subsimplex (=M. cenna (Lehm. & Lindenb.) Gradst.) in a paper about the flora of Galapagos (cited by Clark 1953). In 1905 and 1906, Alban Stewart collected bryophytes which were identified by Wllliams (1924) who described the endemic Lepidopilum crassisetum Williams and Pilotn­ chum obtusatum (=P. corrugatum Ren. & Card.). John Thomas Howell made bryophyte collections along with the Templeton Croker Expedition in 1930. His mosses were treated by Bartram (1933), who recognized 25 species and Clark (1953) identified the liverworts. Howe (1934) described Anthoceros vegetans (=A. tu­ berculatus Lehm. & Lindenb.) from Cocos Is­ land based on collections from the 1930 Astor Expedition. Clark (1953) made the first significant contribution to the knowledge of the liver­ wort flora of Cocos. She newly described Ce­ ratolejeunea lobata and Lopholejeunea coco­ sensis (=L. eulopha (Tayl.) Schiffn.) and lis­ ted 26 liverworts and a homwort based on material coHected by HoweH and earlier re­ ports of Howe (1939), Stephani (1914) and Robinson (1901). In 1964 collecting was do­ ne by W. A. Weber, S. Itow and R. Schuster in the frame of the Galápagos Intemational Expedition (Weber 1993). Fosberg & K1awe (1966) reviewed the literature on the br" yophytes of Cocos Island and Usted 25 mos­ ses and 27 hepatics. In 1992 R. Luecking and A . Bemecker collected epiphyllous lichens and bryophytes and reported 99 lichen spe­ cíes (Luecking and Luecking 1995) and re­ corded 37 bryophytes (Bemecker in prep.). They were also the first to collect bryophytes. 311. in the upper elevations of the island, earlier collecting having been restricted to lower elevations near Chatham and Wafer bays, due to difficulties of access other areas. However, since the latter focused most1y on epiphylls, most of the bryophyte diversity remained unknown. The aimsof-this paperareto providea comprehensive overview of the bryophyte flora of Cocos Island, incorporating new and previous reports of aH bryophyte taxa, and to analyze of the origin of the flora. An analysis of the ecological distribution of the bryophy­ tes in the island is also given.. MATERIALS AND METHODS From January to March and in June 1994, bryophytes were collected hazardously on the island in the greatest possible variety of subs­ trates, the material was identified by the author and María Isabel Morales (liverworts), with help of specialists (see Acknowledgments). The samples where deposited in the Herbarium of the Universidad de Costa Rica (USJ) and in the Herbario Nacional de Costa Rica (CR), and selectively in herbaria of specialists. For the biogeographical analysis the area of distribu­ tion of each species was determined using flo­ ras, checklists and monographs. For many li­ verworts, data on geographical ranges were obtained from S. Rob Gradstein. For the study of the ecological distribu­ tion of the bryophytes, eight plots (10 x 10 m) were set at different altitudes and habitats from o to 600 m (fig. 1). In each plot, 20 qua­ drates (30 x 30 cm) where randomly set on different substrates, inc1uding humus, soil, rocks and trunks. Specíes occurrence in the squares was recorded as an estimate of the frequency of the species in the plots and areas. Similarities between areas were esta� blished through analysis of the bryophyte ve­ getation. The altitudinal distribution of single species was also tracked on the floristic sam­ pIes aH over the island..

(4) REVISTA DE BIOLOGÍA TROPICAL. 312. TABLE 1 Bryophyta and Anthocerotophyta of Cocos lsland. Costa Rica: Family. genera and species llumbers. Taxon (number of families). Species number. Number of genera. Hepaticae ( l O) Aneuraceae Cephaloziaceae Cephaloziellaceae Frullaniaceae Geocalycaceae Herbertaceae Lejeuneaceae Lepidoziaceae Plagiochilaceae Radulaceae Musci (17) Bartramiaceae Bryaceae Callicostaceae Calymperaceae Dicranaceae Fissidentaceae Hypnaceae Leucobryaceae Leucomiaceae Meteoriaceae Neckeraceae Orthorichaceae Pottiaceae Rhizogoniaceae Sematophyllaceae Thuidiaceae Anthocerotae (1) Anthocerotaceae. 43 2. 2 27 5. 33 1 1 6 2 2. 2.3. 6 3 1 63 9 6 4. 6 1.5 1 2.3 1.8 6 4 1.6 3. 10 8 3 8 3 5 1 2. 2 1. Total (28). 98 2. 54 3. 3 2. Sp.lgen. ratio. 1.7 4 1.5 8 2.5 1. 1 5. 1 6. 1.2. 76. 153. 2.0. RESULTS Floristics: the bryophyte species of Cocos Island are listed in Appendix 1. Families of he­ patics are according to Yano and Gradstein (1997), those of mosses according to Churchill & Linares ( 1995). For each species one or a few specimens and/or literature references are gíven. For a complete list of specímens and re­ ferences, see Dauphin ( 1995). Table 1 provides a summary of the br­ yophyte taxa reported for the island. A total of. 153 bryophyte species in 76 genera and 28 fa­ milies are listed. Physantholejeunea portori­ censis (Hampe & Gott.) Schust. is a new re­ cord for Costa Rica. Geographlcal distribution: the geographi­ cal distributíon of the bryophytes from Cocos Is­ land is shown in Table 2 and Fíg. 2. a.. Neotropical element: it ís the best repre­ sented in the bryoflora (55%). Among li­ verworts (50%), most of the elements belong to the families Lejeuneaceae,.

(5) 313. DAUPHIN: Bryophytes of Cocos Island. Plagiochilaceae, Lepidoziaceae and Ra­ dulaceae, and sorne of the species e. g. Plagioehila vineentina and Bazzania hookeri conform important elements of the bryophyte flora of premontane and montane forest also in continental areas. As for mosses (62%), typical lowland · elemént ate · gúúd represented-in-the is­ .. c;land by species like Syrrhopodon circi­ natus, Fissidens elegans and others (see d. altitudinal distribution). b. Pantropical element: being the second best represented (10%), contains most1y Lejeuneaceae (e. g. Lejeunea. [lava and Lopholejeunea nigrieans) and Araehniopsis diaeantha (Lepido­ ziaceae) among liverworts, and com­ mon species like Oetoblepharum albi­ dum, Philonotis uncinata, Calym- peres afzellii and Pyrrhobryum spiniforme among mosses. ·Caribbean element: better represented. among hepatics ( 14.3%) thanmosses ( 1.8%). Northemsouth-american element (4.5%): inc1udes species whose distributional ran­ ge is not necessarily well known (e. g. Le­ pidolejeunea omata and Symbiezidium dentatum).. TABLE 2. Biogeographical distribution of Cocos Island's Bryophytes Zone. % Musci (n=55) O 1.8 1 .8 O 5.4 O 62 1.8 1 2.7 22. AmphiatIantic (Aa) Caribbean (C) Central American (Ca) CosmopolitaIi (Co) Endemic (E) Sub-endemic (Es) Neotropical (Neo) Northem ( South America (Nsa) Pantropical (Pan) Unknown. 40. O. 1.9. 9.1. 1.3. 1.3. � () ()as ()o. 3.2. 1.3. w. U) w. 11.7 6.5 10.4 o <1) z. as U) z. c:. �. f.. ><. Fig. 2. Percentual geographical distribution oí Cocos Island's bryphytes (see appendix for regions).. g. e.. Endemic element: Pilotriehum rugifo­ lium C. M.; P, eorrugatum Ren. & Cardo and Lepidopilum erassisetum Williams are, so far, the onlyknownendemics for Cocos Island: the last two are commonly found in lowland areas of the island, and. h.. & Anthocerotae (n= l ) O O O O O O 100 O O O. Total % (n=1 54) 1.9 9.7 1 ..3 1 .3 3.2 1.3 54.0 4.5 10.4 12. eorrugatum ismore common at midd­ le and higher elevations of Cocos. Sub-endemic element (afier Pócs 1988): Calypogeia rhynehophylla (Herz.) Bisch1er and Stenorrhipis sp. nov. are plants known on1y for Cocos Island and specific loca­ tions. in the Guanacaste Cordillera, Costa Rica. Stenorrhipis sp. nov. has also been found in Panama (Dauphin et al. in press.). Cosmopolitan element: inc1udes 2 wides­ pread hepatics: Aneura pinguis (L.) Dum. and Lophoeolea bidentata (L.) Dum. The f1[st one found a very suitable habitat on boulders along protected ravines. Unknown: most1y material identified to genuslevel.. P.. 54.5. 60 20. % Hepaticae (n=98 ) 3.1 14.3 1 .0 2.0 2.0 2.0 50.0 6. 1 9.2 10.

(6) REVISTA D E BIOLOGÍA TROPICAL. 314. Ecology. Substrates: The isIand is mostly covered by forest, therefore more than 70% of the local bryophytes grow epíphytic: 46% on bark, 25% epyphyllous. Rocks provide another important substrate for bryophyte species (23%), most of them mosses (e. g. Fissidens and Callicostaceae) and taxa associated with riparían habits. Cocos Island does not show well developed colonies of terrestrial bryophy­ tes, even though 12% of them have been found growing on soB or humic soil. Only in higher elevation areas with exposed soil are fre­ quently seen species of Dicranella, but other­ wise the dense leaf cover on the understory of the surroundings do not allow terrestrial br­ yophytes to grow. On rotten logs a few well-. distributed moss taxa Iike Octoblepharum al­ bidum, Pyrrhobryum spiniforme, Isopterygium tenerum and CallicosteZZa spp. can be found (4.5%). General altitudinal distribution: Table 4 includes altitudinal distribution data from plots and individual samples from different areas of the island. Only tbree of the reported bryophyte species have been found in the whole elevation range of Cocos, Plagiochila vincentina, Pilotri­ chum corrugatum and Leucomium strumosum. Most of the bryophyte species (70%) are found under 250 m eIevation, corresponding to the 10wIand areas (Table 5). Altitudinal distributiou in plots: tabIe 6 summarizes the pIot result data. The bryophy-. TABLE 3. Comparisol! 01 the bryophte flora 01 different oceanic islands Is1and. Area (Km2). Cocos. 24. S aba & Saint 33 Eustatius 1 79 Juan Fernández 250 Seychelles Samoa. max altitude (m) 600. Hepatic genera. Moss Hepatic Moss species endemics genera. Gen/sp. ratio. References Dauphin in litt.. 33. 54. 6.35. 870. 45. 63. 2%. 1 .6moss 2.3 hep. 1 .4 moss. 1 65 1. 73. 1 29. 1 9%. 1 . 8moss. Robinson 1 975. 2 . 1 hep. 2.2hep.. Grolle 1 97 8 Gradstein & Weber 1982 Gradstein & Weber 1982 Matteri 1 986. 43. 98. 900 1 850. 38 ca. 60. 80 ca. 1 3 0. 50. 110. Ga1apagos. 7900. 1 700. Malvinas. 11718. 705. 2.3%. 15% 57. TABLE 4. Altitudinal distribution o[ Cocos lsland's Bryophtes, Costa Rica E1evation range. Group. AH (n=1 54) Mosses (n=55) Hepaticae (n=98) Lejeuneaceae (n=64). Moss Moss species endemics. <. 250m 71% 76% 67% 75%. 2:250 m 29% 24% 32% 23%. 1 02. 5 .9%. 2.2hep.. 141. 9 .2%. 2.5moss. Wiersma 1 984. te specíes number increases with altitude and reaches its maximum of 25 species/ 100 m2 in pIot 6 (380 m). The specíes number decrease with further elevation, due to an increasing do­ minance of Bazzania hookeri and Plagiochila vincentina, which cover a whole range of mi­ crohabitats and probably displacing almost all other epiphytes. Vegetation analysis based on bryophy­ tes:. 1. Lowland forest (plots 2-3). Plot 2 is an.

(7) TABLE 5. Species occurrence in 10 x 10 m plots in an altitudinal gradient in Cocos Island, Costa Rica! Plot. Name. Elevation number. Wafer Bay. 10 m. Total species. 07. Bryophyte dominance (taxa occurence rate in 30 x 30 cm quadrates in the plots). Vascular Plants. Fissidens neglectus 0.85; Callicostella depressa 0.70; Taxithelium planum 0.4; Stictoejeunea squamata 0.30 Symbiezidium transversale 0.75, Radula macostachya 0.65; Plaiochila rutilans, Calymperes aizellii, Fissidens neglectus 0.55. Hibiscus titiaceus on old seéondary growth Selaginella sp. on a rocky (basalt) sollo Annona glabra on a bare sandy soll and densely coveredwith epihptes, esp. Guzmania sanguinea.. 2. Wafer B ay "Annona swamp". 3. Genio River. 20 m. 14. Radula macrostachya 0.45; Frullania mucronata 0.20; Symbiezidium transversale 0.5; S. barbiflorumO.15; Cyclolejeunea convexistipa 0.15.. Saccoholdridgei, Clusia rosea, Miconia spp, Hypotitrum amplum; undsturberd forest on river side. Densely covered with epiphytes e .. g. Guzmania sanguínea, Marcgravia,waferi and Tillandsia sp.. 4. "Gallinero". 110m. 12. Callicostella depressa 0.40; Lepdopilum crassisetum 0.60; Taxithelium planum 0.30; Plagiochila rutilans, Fissidens angustifolius 0.15.. Ocotea insularis, Cecropia pittierii o n a rocky soil with Coffea arabica in the understory.. 5. Mirador. 250m. 12. Calymperes aizellii 0.45; Octoblepharum pulvinatum 0.35; Plaggiochila rutilans 0.10. Saccoglottis holdridgei, with tree fems and Hypolitrum amplum and Melastomataceae on a hurnic soil. G. sanguinea, M. waferi, peperomia sp., Tillandsia s Orchidpceae, Polypodium subgen. goniophlebium, Eaphoglossum sp., Oleandra sp., Hymenophillaceae.. 6. Cerro Pelón ridge. 380m. 25. Syrrhopodon ly copodioides 0.60; Bazzania hookerii 0.60; Plaiochila vincentina 0.50; Octoblepharum pulvinatum 0,32.. S. holdridgei, other taxa similar to previous plot.. 7. Cerro Pelón ridge. 500 m. 22. Bazzania hookerii 0.70; Octoblepharm pulvinatum, Plagiochia vincentina, Cephalozia crassifolia 0.50.. S. holdridgei, other taia similar to previos plots.. 8. Cerro Y glesias. 600 m. 18. Plagiochila vincentina 0.60; Bazzania hookeri 0.50; Cephalozia crassifolia 0.40. Om. 12. 1 Sorne 52 species, i.e. 34% of the total species number reported were found in the plots.. ' S. holdridgei, Euterpe macrospadix, and terresmal and tree-fems; Peperomia, Marcravia and Orchidaceae as epiphytes..

(8) 316. REVISTA DE BIOLOGÍA TROPICAL. Annona glabra swamp that may become inun­ dated, and has no terrestrial bryophytes and with a low canopy. Plot 3 belongs to the most common and extensive Cocos tropical forest. Saccoglottis holdridgei is the dominant tree, and on the ground grows abundant Hypolitrum amplum. The epiphyte flora is rich with Guz­ mania sanguinea, Marcgravia waferi and Ti­ llandsia sp. Radula macrostachya and Symbie­ zidium transversale are the bryophytes most frequently found here on lower parts of the trunks (Table 5), higher branches show a diffe­ rent bryophyte component, where Frullania mucronata, F. riojaneirensis and sun bryophy­ tes are found. 2. Secondary lowland forest (plots 1 & 4): located on rocky and exposed soll, ofien also on high pendent terrains, this forest type may reach up to 200-250 m elevation. Sorne of the­ se areas where probably exploited agricultural at a time, plot 4 shows saplings of coffee plants . that may have been planted somewhere near before. The vascular flora elements are Hibis­ cus tiliaceus at the sea level and Ocotea insu­ laris with Cecropia pittieri which make up an open canopy on hins. Mosses domínate typi­ caHy this areas, which provide aboye all rocks and the hard dry barks of the Ocotea and Ce­ cropia, which are frequently covered by Lepi­ dopilum crassisetum; Callicostella depressa and Taxithelium planum are growing on logs. Aneura pinguis, Fissidens neglectus and Calli­ costella depressa grow on rocks. 3. Montane forest (plots 5-8): the domi­ nant tree in this forest type is Saccoglottis hol­ dridgei, accompanied in higher areas by Euter­ pe macrospadix. Found over 200 up to 600 m, this forest is typified by an increasing abun­ dance of corticolous hepatics, especially Baz­ zania hookeri and Plagiochila vincentina, that first grow conspicuously on the horizontal branches of trees and then cover down the trunk and are also important part of the mantle. This is particularly evident in the upper part of Cerro yglesias where an overwhelmíng mass of bryophytes covers aH trees and saplings. In a more detailed scale, sorne areas show diffe­ rent dominance pattems, for instance, Cerro. Escondido (Figure 1), where Herbertus diver­ gens is covering all trunks and branches at about 450 m. Protected areas as creeks also provide shelter for other bryophyte associations. Parti­ cular bryophyte associations: 4.a. Pasture areas: on the eastem side of Chatham bay the­ re are. sorne open areas that bear sorne particu­ lar associations similar to those described for secondary lowland forests. Typical of these ha­ bitats are Sematophyllaceae and Callicosta­ ceae. 4.b. Rivers and waterfalls: provide a substrate and habitat for different bryophyte species and associations, which vary depen­ ding on substrate and light availability. For ins­ tance waterfalls that are richly covered with Philonotis uncinata, and give a metallic glance as the drops reflect the sunlight.. DISCUSSION Floristics. As usual in tropical lowland rain forests (Richards 1983), liverworts ove­ rrun mosses in numbers of species and genera, but moss farnilies are more numerous. Data for comparison with other island areas are provi­ ded by Table 3. A different situation was found in the Galapagos bryophyte flora, where the number of hepatics is almost the same as that of mosses (Gradstein & Weber 1982). This may be due to the greater diversity of habitats, including dry and wet ones, in the Galapagos which has also a larger area than Cocos Island. A similar number of mosses is found in the West Indian islands Saba and Saint Eustatius (Wiersma 1984), territory of somewhat similar geographical and climatic conditions. In Juan Femández and Malvinas, the higher species numbers of mosses can be explained by their larger territory and more temperate climate, that seems to favor the growth of other moss taxa. The most species rich moss farnilies in Cocos, Callicostaceae, Calymperaceae and Fissidentaceae are also farnilies which attain the highest diversity in tropical areas (Richards 1983). Families like Pottiaceae and Bryaceae appear poorly represented in the list. This may.

(9) DAU,f1HIN; Bryophytes of Cocos Island. ·. ·. bedue to undercollecting of exposed areas of the island, e. g. Cabo Dampier, vertical risks around the island. The liverwort diversity in Cocos Island shows the typical pattern of tropical lowland forest, where Lejeuneaceae is by far the most dominant. family in terms of species number (Tablel). This domiriance paUerriis also typi­ cal for tropical continental and island areas (e. g. Grolle 1978, Gradstein 1995, Dauphin et al. in press). There is no poor representation of important mainland groups of other tropic al Jungermaniinae as seen in Galapagos (Grads­ tein & Weber 1982). In comparison to Galapa­ gos, Cocos Island shows more tropical compo­ nents (see biogeography), as stated by Grads­ tein & Weber (1982). Thallose hepatic taxa (except Aneura and Riccardia) are absent in this listo A similar situation was found in the Seychelles (Groll e 1978). This may be due 10 the few natural exposed areas in t�eisland; AI­ so a. wetter climate does notfavor the. growth of thallose taxa, as it happens in the dryet Ga- . lapagos Archipelago (Gradstein & Weber. 1982). Ilryophyte speciesnumbers in oceanÍc 1S­ lands seem 10 increase with terri!ory size (Table 3). . Another importantelementi#ll!encing br­ yophytediversity is the presence ofdifferentaltitudinalbelts, what is reflected in a greatermicrohabitat \diversity and. niches for more bryophyte taxa. This latter factor .Q1a};' be more important .t:han.· the long"range dispersal capa­ city of·the different taxa to eJ(plaín the occu­ trence of a taxon in an oceanic isIand. Geogra�hical distribution. Most of. the bryophyte taxa in Tropical America havea wi.. qe distríbl1tion. Thy dominant Neotropical ele­ .Q1ent . may cOntain more species now known only frOillSOrtlerestricted areas. In the same way ·thePantiÓpical.eIement may . show more eh�ments,what can beput in evidence only th­ rough worldtreatments of many t�a, which areusually lacking. Concerning the Caribbean element, Gradstein & We\Jer (1982) explained the generic resemblance of. Puerto Rico and Cocos Islandsin terms c1imatic factors. The latter authors also found a higher affínity. ·. 317. among Galapagos and West Indian lUosses. Abetter knowledge of the Northernsouth­ amerícan element component of the flora may result in putting these taxa together with the Central American or Caribbean elements. Be­ cause of the situation of Cocos before the South Americat) Coast, and its even further southem origin on the Galapagos hotspot (Castillo et al. 1988), it is. to expect sorne affi­ nity of both areas. Endemic elements: the status of the two hepatic species reported as endemic is uncer­ tain: Ceratolejeunea lobata Clark may falI in­ to synonymy of a wide distributed tropical ta­ xon in that genus, as commented bythe author (Clark 1953); the generic and specific identity of Hygrolejeunea ocellata Steph. must be checked on the type1. This low bryophyte en­ demism rate (3%) compared to that of flowe­ ring plants(9.3% fide Montoya 1990b) i8 normal in oceanic islands (Gradstein & Weber 1982, Pócs 1997). A higher endemism of mos­ seS is also see n in oceanic islands, as the case of Cuba, where 12% of moss endemicsis shown (Pócs 1988). In respect toGalapagos, the cauSeS of lower endelUism in Cocos may be eaused by the homogeneity of the area (Fout" .nier 1966), that do notprovideareas.so suit�­ ble for species diversification as the dry, expo­ sed and older areas of Ga.lapagos Archipelago (Gradsteih & Weber 1982). In the Sub-ende­ mie element (afterPócs 1988), belongspecies like Calypogeia rhynchophylla andSumorrhi­ pis sp. nov., which have restricteddistributioI1 ranges.Even though the first taxon has been mentioned lo be endangered (Gradstein 1992), this speciesgrows abunpantlyon Iogsin the upperelevations of the island (Dauphin 1995). The species conformingthe Cosmopolitan ele­ mentoccur in non-disturbed .habitats. The ab­ senee of other eosmQpolitan taxa like Bryum argenteum Hedw.· can beexpl�éd by the ab­ sence of roeks in the higher e1evatíons where jt. .. Mizutani (1970) names this plant Pycnolejeunea cocosana Mizut. According 10 Xiao-Lan He (Pers comm. 1998), the type does no! be10ng into that gem:js. Its identity still has to be t<.stablishe4.. ..

(10) 318. REVISTA DE BIOLOGÍA TROPICAL. could eventually grow. In general, the geographical distribution of bryophyte taxa in Cocos Island agree with that of other areas, especially with Neotropical ones with wmch most of species are shared. If the specíes/genus ratio is used as a measure of the degree of isolation (Gradstein & Weber 1982, Pócs 1997), Cocos shows similar figures as islands with comparable species numbers (2.3 hepatícs,l.6 for mosses, Table 3). Low en­ demism is a common feature for spore-produ­ cing orgarusms as fems, lichens and bryophy­ tes, groups that also show low endemism in Cocos Island and elsewhere (Gómez 1975, Luecking & Luecking 1995). The lack of mo­ re detailed data on distribution of the different taxa do not allow to make more precise analy­ ses of biogeographical relationship between different areas. Vegetation analysis based on bryopby­. l. Lowland forest (plots 2 & 3). Hen'era & Gómez ( 1993) call this as a Tropical, Tropical very wet with one or two dry months biotic zo­ neo These authors show similar areas in conti­ nental Costa Rica mainly in the plain of Osa Península on the Pacific coast and the Northem part of the San Carlos plain. Tms forest shows the typical structure of a Lowland Tropical fo­ rest, with a vertical distribution of the species according to their light and moisture require­ ments (Comelissen & Steege 1989, Frahm & Gradstein 199 1). Secondary lowland forest (plots 1& 4): the structure of these areas is si­ milar to secondary growth areas in continent, with the particularity of showing abundant en­ demic species. Montane forest: such a forest type can be compared with the "Lower Montane Forest" described from El Darién, Panama by Grads­ tein and Salazar Allen ( 1992). Montane forests are normally found between 1800 and 2400 m in continental areas, but due to the excessive moisture in this place, the altitudinal belts are lower, what has been caBed "Massenerhebung effect" (Fralun & Gradstein 199 1). This i8 par­ ticularly evident in the upper part of Cerro yglesias where an overwhelming mass of br­ yophytes covers all trees and saplings. Cerro tes:. yglesias. forest can be compared with that from El Darién, between 1 100 and 1200 m aboye sea level, as described by Gradstein and Salazar Allen ( 199 1). Herrera & Gómez ( 1993) provide two biotic zone subdivisions for Cocos Island's higher elevations: Subtropical, tropi­ cal with one to two dry months and without dry months. This latter subdivision corresponds to the summit of Cerro yglesias. These authors base this subdivision on c1imatic factors (more rain), but the differences seen on bryophyte ve­ getation, do not support this division, for a continuous species association and dominance pattern is seen in the whole mountain area. The latter results provide bryophytical evidence f01" Grubb's observation, who pointed out that me­ langes of f10wering plant species of the diffe­ rent forest belts may be found where the altitu­ dinal limits for the formation types are lowered (cited by Frahm & Gradstein 1991). Further collecting and survey work of the bryophyte flora on Cocos island is needed to provide informatíon on many more different bryophyte communities there, also as source of informatíon on bryophyte conservatíon.. AKNOWLEDGEMENTS The present work could not have been do­ ne wíthout the help of the staff of the Isla del Coco National Park, especially Joaquín Alvara­ do who gave his approval and logístic help and Wilfrido Conejo who helped me reach with ski­ Hed boat otherwise inaccessible places. Many group specialists have kindly helped with the detennination of specimens, namely (alphabetical order) B. H. Allen (Dicranaceae), A. Bernecker (Bazzania), M. Crosby (Pilotrichum), S. R. Gradstein (Plagiochila and general hepati­ cae), D. Griffin m (Bartramiaceae), M. I. Morales (general hepaticae), R. Pw'seU (Fissidens), N. Sa­ lazar Allen ( Octoblepharum) and K. Yamada (Ra­ dula). A. Bemecker & R. Luecking provided the species list from their manuscript 011 epiphyll bryophytes and M. Montoya his manuscript on Cocos' vascular plant flora. L. Fournier kindly.

(11) DAUPfIlN: B ryophytes of Coco� Island. helped;with references.. 1 thanJ<: also Luis Die­ go Gómez, S. Rob Gradstein imd María Isabel Morales for their valuable c omments on diffe­ renLphases of this work.. RESUMEN. 319. Bartram, E. B. 1933 : Mo'ss es of the Templeton Crocker Ex­ pedition, collected by John Thomas Howell; and lists of mosses known from the Galápagos Islands and from Cocos Island. Proc. Cal. Acad. ScL 21: 75-86. Bartram, E. B. 1949. Mosses of Guatemala. Fieldiana Bo­ tany 25: i-v, 1�442.. �.. c er, 1%4:te geme D7�UfU71ejeunelrSreph;� Se comunica la presencia de 15 3 � specit:$ de briófi tas �is hl Aménque Centrale et Méndionale. Rev. Bryol. Lly antocerotófitas en 1 a !sIa de C ocos ·(5032'N, 870Q4'W) , chéno1 . 35'. 95-137 . Costa Rica. Estas son 98 especies de hepáticas en 43 géneros y 10 fanrilias; 54 especies de musgos en 33 géneros y Bischler, H., C. E. B. Bonner & H. A. Miller. 1963. Studies in 17 familias Y una especie de antocerotófita. Más del 60% Lejeuneaceae VI. The genus Microlejeunea Steph. in de las briófitas tiene amplia distribución en el Neotrópico, Central and SouthAmerica. Nova Hedwigia 5: 359-411. 10.4% distribución pantropical, 9.7% pertenecen al ele­ .mtlnto Caribe, 1 .3% al elemento Centroamericano , 4.5% al Castillo , P., R Baliza, p. Vanko, E. Malav�si, J. sai-qúero Nor-sudamericano, 3.2% son endémicos , 1.3% 'subendémi­ & E. Femández. 1988. Anomalously young volcanoes cos y 1.3% subcosmopolitas. En comparación con Galápa­ on old hot spot traces: 1. Geology and petrology of Co­ gos, la Is� de Cocos muestra una brioflora más tropical, en cos Islánd. Geol. Soco America Bul. 100: 1400-1414. su rnayorp8rte COmpuesta por hepáticas foliosas (e. g. Le­ jeuneácea6, Lepidoziaceae) y una menor rep�sentación de hepática$ talosas y musgos. La mayor variedad de hábitats .'Churchill, S. P. & E. L. Linares. 1995. Prodromus Bryolo­ tanto húinedós como secos y expuestos y el area mayor del giae Novo-Granatensis, Introduc�i!5n a la . flora de .J\rcWpié!ltgo :expliCán el rnayor número de. especies de musgos de Colombia.. fustiWto � :qenci as NatlJra­ .' .brfófl.iá§:¡;Ill<lOntradas en Galápagos. U11 abáli sik ,del uso de les-Museo de. His.toria Natural Wblioteca "José Jeró­ sUstratos revela que la rnayoip� de lasbrlófltas (46%) nimo Triána", 12, Bogl;>tá. 926 p. ;utilizan corteza como sustrato, 25 % s�n epífilas, 23% epi­ ;líRCas y 12%N�íTesttes· Se registró ;laocUt;ren:�ia de:btiófi­ Clark, L. 1�53; Some Htlpat;icaefrom the, Galápago�,'Co­ 't$e� p'¡u-oefás de lOx 10 m con cuadiículliS de30x30 cm co� ándo�er Pacific Coast islands. Proc . Cal. Acad. en: ((listintos hábitáts y elevacionés en la Isla.;<ta:;dilitribu­ SóL, 27: 593�624. cióíh:ie: bgófltlts en la Isla ;¡:oincide con bO$que de bajura (OC100 ni); bosque seqJ111dlolÍ:i de bájura (0-2OQ m) y bos­ Cornelissen,J;H. C. & lJ¡ Ter Steegé. 1989;J:>i¡¡tributio lII!d que 'm6Iltanq í200-600 m).:La excesivá h�e<la,d del am- ' .. bry()phytes anct lichens in.. dtY e<:üloror:pf. bieniep�ióGa:que el cambio altitlldiÍl.al dp.!J'1I cBmp osición evergteen fore,�í ot'Guyana J. 'fróp/'Ecol. 5: 131-150. de especies en la isla se vea acortado y sé presenten unida­ des .. montaiíosas que se e�e!1triu:1 a 1Illj.yór áltitud en areás Cn¡J¡¡by, M. R 1969. A re\lis�bn of the trqpicalIDOSS genus continehtliIé$ (HeCto ''M:ássenerhe11ífi¡g''). Phys�nthiJ(ejeu- ' , . J>i(fltrichlim. Bryolpgist 72: 275...343. nea ¡'órtoricenSis (RamPe & Gott) Séhl,Tst. .es un n�vo re:­ gistropw;a Costa Rica. .Crum, H. 1994. Various fa,milies and genera treatments. InA.. �. eP!Jl�c. J, Sharp, H; CnwJ.8¡;P. &¡kel (e¡ls.). TheMoss'flora or . Mexieo. Mem. New '(brk Bot. Gard. 69: 1-1113.. REFERBNCES?. AlÍ�lll.¡g}H. '1994: Dicrijn¡¡céae. mB. ¡:f¡AIleP· (ed.) Moss ' flPi!l of Central Ameiiéa. Mo?ógrapbs Syst. Bot. .. ,, ;;Mis souri Bot. Gard.,49: 81-168. '<;;�. ,( , ' , ;, : ' AlÍeH;,'B¡H;'&; M. R, Ci6sí¡y, i 986. Revisionofthe genus .. Squamklium (Múscí: MeteoriaCéae), J.'ltlltto¡:i Bot. . ' .' " Lab. 61: 423.-476.. ' A1varado l., G. E. 1984. Aspedas petro16gic¡)s�geoló@.cos de los volcanes y urildades lawc as del ' eenózoic¡:¡ su­ perior de Costa Rica. Tesis de LiceÍlclahrra. Eséuela. Centroamericana de Geología; Universidad de Costa ". Rica, &1!IlJosé.. �. ,. ,. C¡:um, l:l. & W. R Buck. 1994. Thuidiaceae. m A.J. Sharp, . a, CfQIIl & P. Eckel (eds;). TheMoss.Flora of Mexi, co. Mem, New York B ot. Gard .. 69: 873:886,. " '. ,',: " ,' ",' .,J " , Da,uphin,G.. 1995 . . Briófj.tos de la !slá de Cocos: Diversidad . . y'Ecolpgfa.Tesis de Licencia,tura.;Escueh de. Biolo-. gía: Universidad de Co sta .Rica, SI!Il Pedro .. ' ' ( Dauphin, O,,' S. R. ÓradsU; , A. Bemecker-Lu�lting & M. 1. Morales. 1998. Additions to .the H�patic flora of Costa RicaII ;t..in4bergia 43:. 74-80. f. k. , " ,', ;'\.;". ,; -' " .' � -�, ¡; ',' Delgadillo , C., B. Belló & A. Cárdenas. 1924,J-atmoss, a cat¡p.o gue of neotropical Mosses. Morió�íl�hs Syst. Bot. Misso�Bot. Gard. 56: 1-191. ". ¡. ,. \. :.

(12) 320. REVISTA DE BIOLOGÍA TROPICAL. Fosberg, F. R & w. 1. Klawe. 1 966. The Galápagos Inter­ national Scientific . Preliminary list of plants from Cocos Island, p. 1 87- 1 89. In R S . B owman (ed.). The International Scientific Proj ect, Berkeley & Los Angeles. Fournier, L. A. 1 966. Botany of Cocos Island, Costa Rica, p. 1 83- 1 86. In R S. B owman (ed.) . The Internatíonal Scientific Project, Berkeley & Los Angeles. Frahm, J. P. & S . R. Gradstein. 1 99 1 . An altitudinal zona­ (ion of rain forests using bryophytes . J. Biogeo­ graphy 1 8 : 669-678 . Fulford, M. H. 1 945. Studies o n American Hepaticae IV. Ceralolejeunea. Brittonia 5: 368-403 .. Gradstein, S. R, A. Luecking, M. I. Morales & G. Daup­ hin. 1 994. Additions to the hepatic flora of Costa Ri­ ca. Lindbergia 19: 73-86. Gradstein, S. R. & N . Salazar Allen. 1 992. Blyophyte di­ versity along an altitudinal gradient in Darién Natio­ nal Park, Panamá. Trop. Bryo1. 5: 6 1 -7 1 . Gradstein, S . R & w. A . Weber. 1 982. Bryogeography of the Galápagos Islands. J. Hattori Bol. Lab. 52: 1 271 52. Griffin III , D. 1 994. Bartramiaceae. In A. J. Sharp, H. Crum & P. Eckel (eds.). The Moss Flora of Mexico. Mem. New York B ot. Gard. 69: 537-574.. Fulford, M. H. 1 962. Leafy Hepaticae of Latin America. Part 1. Mem. New York Bot. Gard. 1 1 : 1 - 1 72.. Grolle, R 1 956. Revision der Clasmatoeolea Arten. Rev. Bryo!. Liche' nol. 25: 288-303 .. Fulford, M . H. 1 966. Leafy Hepaticae of Latin Ametica. Part n. Mem. New York Bot. Gard. 1 1 : 1 73-276.. GroBe, R 1978. Die Lebermoose der Seychellen. Wiss. Zeitschr. Friedrich-Schiller Univ. Jena, Math-Nat. R 27: 7- 1 7 .. Fulford, M. H. 1 968. Leafy Hepaticae of Latín Arnerica. Part III . Mem. New York Bot. Gard. 1 1 : 277-392. Fulford, M. H. 1 976. Leafy Hepaticae of Latin Arnerica. Part IV. Mem. New York Bol. Gard. 1 1 : 393-535.. Haarbrink, P. 1 9 8 1 . Studies o n Colombian Cryptogams XI. High Andean Species of Frullania sllbg. Chonante­ lia (Hepaticae). Lindbergia 7: 47-57 .. Gómez, L. D. 1975. Contribuciones a la pteridología cos­ tarricense. VII. Pteridófitos de la Isla de Cocos. Bre­ nesia 6: 33-48.. Haessel de Menéndez, G . A. 1 990. L a s especies d e Ant­ hoce ros y Folioeeros (Anthocerotophyta) de Amé­ rica del Norte, Sur y Central: la ornamentación de sus esporas taxonomía. Candollea 45 : 20 1 -220.. Gómez, L. D . 1 986. Vegetación de Costa Rica, vol. ii. In L. D Gómez (ed.). Vegetación y Clima de Costa Ri­ ca. EUNED, San Jose ' .. He, X. L. 1 996. Type studies on Pyel10lejeunea (Lejeunea­ ceae: Hepaticae), n. Ann. Bol. Fennici 3 3 : 5 1 -58.. Gradstein, S . R 1 992. Threatened bryophytes o f the neo­ tropical rain forest: a status reporto Trop. Blyol. 6: 83-93. Gradstein, S . R 1 994. Lejeuneaceae: Ptychantheae, Bra­ chiolejeuneae. Flora Neotropica Monograph 62. New York Botanica] Garden, New York. 2 1 6 p. Gradstein, S . R. 1 995. Diversity of hepaticae and Antho­ cerotae in montane forests oí' the tropical Andes, p . 3 2 1 -334. I n S . P. Churchill el al. (eds . ) . Biodiver­ sity and Conservation of Neotropical Montane Fo­ rests. The New York Botanical Garden, New York. Gradstein, S. R. 1 997. A guide to the bryophytes of Tropi­ cal America 1. Liverworts and Hornworts. Brussels. Gradstein, S. R & W. H. A. Hekking. 1 979. A catalogue of the Hepaticae of Colombia. J. Hattori Bol. Lab. 45 : 1 97-230.. HeITera, W. 1 986. Clima de Costa Rica, vo1.2. In L. D. Gó­ mez (ed.). Vegetación y Clima de Costa Rica. EU­ NED, S an José. Herrera, W. & L. D. GÓmez. 1 993. Mapa de Unidades Bió­ ticas de Costa Rica, escala 1: 685 000. The Nature Conservancy el al. , Costa Rica. Hertlein, L. G. 1 963. Contribution to the biogeography of Cocos Island, including a bibliography. ProC. Cal. Acad. Sci. 32: 1 29-289. Howe, M. A. 1 934. The Hepaticae (chiefly Riecia and Ant­ hocerotae) of the Galápagos Islands and !he coast Is­ lands of Central America and Mexico. Pmc. Ca!. Acad. Sci. 2 1 : 1 99-2 10. Ireland, R R 1 992. The moss genus Isoptel}'gium (Hypna­ ceae) in Latín America. Trop. Bryo!' 6: 1 1 1 - 1 32. Jovet-Ast, S. 1953. Le genre Calura. Rev. Bryo!. Liche­ ' no!' 22: 206-3 1 2..

(13) 321. DAUPHIN: Bryophytes ófCocos Island. Lichénd,. R. & A. Lücking. 1995. Fo1iicolous lichens and . bryophytes from Cocos Island, Costa Rica, a taxono­ mical and ecogeographical study. 1. Lichens. Herzo­ gia 1 1 : 143- 1 74.. Magill , R. 1 994a. Acroporium Mitt: In A. J. Sharp, H. Crum & P. Eckel (eds.). The Moss Flora of Mexico. Mem. New York: Bot. Gard. 69. 989-993.. Re�se, W. l). 1994 . Chlymperaceae. In B. H. Allen (ed.). . Moss · Flora of Central Anlerica. Monographs ' Syst. Bot. Missouri Bot. Gard. 49: 1 92-225. Richards, P. W. 1 98 3 . 'fhe ecology of tropiCal rain forest . bryophytes, p. 1233-1270. In R;; M. Schiister (ed.). New Manual of Bi"yology. The Hattori Botanical La- . boratory, Japan. 1295 p.. ,. <�. � �---��---_._;--��. - �'- " �'-'. Magill, R. 1 994b. Sematophyllum Mitt. In A. J. Sharp, H. Crum & P. Eckel (eds.). The Moss Flora of Mexico. Mem. New York Bot. Gard. 09. 993- IQOO.. Maneri, C. M. 1 986. Los Musci (Bryophyta) de las Islas Malvinas, su ha'bitat y distribución. Nova Hedwigia 43: 1 59-89. Mizutani, M . 1 970: Lejeuneaceae, subfamilies Lejeuneoi­ deae and CololejeuÍleoideae from S¡ibah (North Bor­ neo). J. Hattori Bot. Lab 33: 225-265. Montoya, M. 1990a. Plan de manej�, Parque Nacional Is­ la del Coco. Docmnento preparado para el �istema de Parques . y. ��etVa!i ' Marinas (SIPAREMA.) � Co�ta Rica, San JOSé. .. . Mí>ntoy�M; 199Ób, L�JF1o�a y Vegetacjón � la Islá <k1 ' Cde�, .C�ta Rica.. llscuelide Biol�g¡f!.\Universi . dád ��:�?:s4iruca, Sa/l:.los�.. v· ' •. M . ora1�s i :;, fd : 1. i99f:Ús 'bepátic� C��l1����. Para COs�;,Wéa: 1'rop. BryQJ, !h �$;57: :;: ' d( :. PiippO': S� 1986,. A monograph o( ��ig�pFa iepi�kJ/�jeu� ríéa ru:td Luteolejeunea (t�jeune'ab�ae" Hepaticae): '. . Arm. Bo� Ferlruei 132: .1-69.. Pittier, H. R 1 899. A-puntanJ.ieiItOs prelliÍlll1¡s¡re sobre la ISla de COCos, posesiÓn costarri�nse enel Océano Pacífico. Memoria de la Secretaría � FomentO,. S¡m �(jsé. PóCs, 1Y 1 984. Pre�ent khowledge on Aphanolejeunea Evans. J. Hattori Bot Lab. 55: 307-3 1 3 . ,. ,ji;. �. ,, '. :. ,. ,. PóCs, T. 1988, Biogeography of the Cuban bryophyte flo. ra. Tax011 37: 6r5�621. P6cs, T. 1997. The distribution and origin of the fo1iico " , lous btyophyta in the Indian Ocean islands. Abstfac' ta Bot¡inica 2 1:) 1 23� 1 34. PurSell, . JI. A. 1 994. Fissidentaceae. In B. H. All t<A (e<i:). Mos.s Flora of Central America. Monógraphs Syst. Iiot. Mi ss6� Bot. Gard. 49: 40-80. �. r,. ' ,' ,. ". ,;. Quesaqa Chanto, B . ' 1'997. Isla del Coco declarada patrimo-. . ni� mundia1. "La Republica" 5.Xll. 1 997: 4A.. RooiIri>on;-H:-i:9T5:-1'he-Mosses-tlf-JuatrFema.'.ndez.Js­ lands. Smithsonian Contributions to Botany No.27.. Salazar Allen, N. 1 994. Octoblepharum. In B . H. Allen (ed.). Moss Flora of Central America. Monographs Syst Bot. Missouri Bot. Gard 49: 1 82- 1 89 . Schuster, R. M . Í982. The Hepaticae and Anthocerotae of North America, East of the Hundreth . Meridian. Vo1.. IV. Columbia University Press, New York. 133 1 p . Schuster, R . M: 1 992. The Hepaticae and Anthocerotae of North America, East of the Hundreth Meri¡lian. Vol. v; Fielll Museum of Natural History, Chicago. . 9;37 p . Smjth, D; K. 1994. Neclf:eraceae. In A. J. sl:Íarp, H. Cmm & P. Éc��l (eds.). The �oss lfIora'of Mexico , M;em. New York Bot. Gard. 69: 741 �765. ,. ,. \. ". '. ,. , ', >. �. Stephani , F. 19,j4.<�pecie� HepatiC ¡"¡V0i ; \T: ;565�� 6(Í. Complement to g�e� de l' Ne��er �oissi�, va.. �n�. '. Sf0tler, R'. 1E. 1969;;Th� geb us �rullania(�ú'bg. Frulla. ni� iÍ1 't,atih Anie ca. N�Ya Hed� iií a 18:. 397.. 55�.. �. ". M'. .. . Vttt., D; 6. 1994. érthotticháceae. InA. J; Sharp, F{: C� & P; Eckel (eds.). The Moss Flóta .ot'Me.ii.�. , Mem. . New Yo.rk Bot. Gard. 69: 590-656. ,.. Weber, \". A. 1966. Lichenology and bryologyi:rl the Ga�. . l\ípagos Islánds, with check1i�ts of the ÚcHerls and Bryophytes thus fa!: reporte!!. In �. i. B?winab. (ea.). The Galapagos. orCiillfornia .'Press; Ber' . University ' .. keley.. Web�r, W. A. 1 993. Addit,i.ons to the Galápagos..ru:td Cocos . Islands lichen ánd bryophyte. iIoras�" . �lYo19gist 96: .. 431 434.. Wiersma, ¡p. 1984. lI4()ss Flora and the vege1:<l;tio n Opt. Eustatius CWestIndies). Proceedings C 87: 337�364.. Williams, R S. 1 924. Galápagos and Cocos IsIand Jf1osses ' collected by Mban Stewart in i9(15�1906, BryolQg¡st . .. . 27: 37-44..

(14) 322. REVISTA DE BIOLOGÍA TROPICAL. Yamada, K. 1988. The genus Radula from Cuba. J. Hatto­ ri Bot. Lab. 65 : 379-390. Yano, O. & S . R. Gradstein. 1 997. Genera of Hepatics. Systematisch-Geobotanisches Institut Universitaet Goettingen, Germany. 29 p..

(15) APPENDIX 1. Cocos Island's Bryophyte species genera andfamilies, Specific sex, substrate and geographical distribution Class 1. Family. Genus. Species. Sub2. Sex3. Distr4. Record referenee/ distr. referenee Dauphin 1 1 22/ Hiissel de Menénd�z 1 990 Dauphin 1 096/ Seliuster 1 992 Dauphin 966, 984, , 1 11 , 1 145 Dauphin 963/ Fulford 1 968. Dauphin 965, 982,i 1 2 1 2/ Gradstein et al. . 1 994 Dauphin 965, 1 1 59/ Gradstein et al. 1 994. Dauphin 1024/ D apphin et al 1 99 8 Morales 1 99 1/ Gr�dstein pers. COmIDo Morales 1991/ Stotler 1 969 Fosberg & Klawe :1 966, Morales 1991/ Stotler 1 969. . Dauphin 947, 952 6/ Stotler 1 969. Dauphin 944, Motales 1 99 1 / Haarbrink 1 9 8 1 Dauphin 1 14 1 / Fu;\ford 1 976. Dauphin 967/ Fulford 1 976. Dauphin 1 2 1 0/ GJ!adstein, pers. COmIDo Dauphin 1 146/ Fqlford 1 963. Bemeeker & Lüe\ang in prep Bemecker & Lüeldng in prep Bemecker & Lüekmg in prep Dauphin A-8/ Pó¡;s 1 9 84. Morales 1 99 1 , Weber 1 9937/ Gradstein 1 994.. A H H H H. Anthocerotaeeae Aneuraeeae Aneuraceae Calypogeiaceae Calypogeiaceae. Anthoceros Aneura Riccardia Calypogeia Calypogeia. tuberculatus Lehm. & Lindenb. pinguis (L.) Dum. sp. cyclostipa (Spruee) Steph rhynchophylla (Herz.) Bisehler. r r s lb b,h. X Di x Di Di. Neo Co X Nsa Es. H H H H H. Cephaloziaceae Cephaloziellaeeae Jubulaceae Jubulaceae Jubulaeeae. Cephalozia Stenorrhipis Frullania5 Frullania Frullania. h r b. Di Au Au Di Di. Neo Es Neo Neo Neo. H H H H H H H H H H H. Jubulaceae Jubulaceae Geoealyeaceae Geoealyeaeeae Geocalyéaeeae Herbertaceae Lejeuneaeeae Lejeuneaceae Lejeuneaeeae Lejeuneaeea Lejeuneaeea. Frullania Frullania Leptoscyphus Leptoscyphus Lophocolea Herbertur Aphanolejeunea Aphanolejeunea Aphanolejeunea Aphanolejeunea Archilejeunea. crassifolia (Lindenb. & Gott.) Fulford sp nov. caulisequa (Nees) Nees ecuadoriensis Sreph. intumescens (Lehm. & Lindenb) Lindenb. & Lindenb. mucronata Lehm. & Lindenb. riojaneirensis (Raddi) Aongstr. gibbossus (Tayl.) Mitt. ovatus (Spruee) Grolle cf. bidentata (L.) Dum divergens (Steph. ) Herz. angustissia Steph. camilli (Lehm.) R.M. Sehust. costariensis Bemecker sicaefolia (Gott.) A. Evans fuscescens (Hampe ex Lehm.) Fulford. Di Au Di Di Au Di X X X X Di. Nsa Neo Neo C Co Neo X X X Neo Nsa. x. x. lb lb b lb b b e. e e b,e x. Class: A=&thocerotae, H=Hepatics, M=Mosses. 2 3. Substrate types: b=bark, e=leaves (epiphylls), h=humus, Lb=living bark, r=rocks, rb= rotten bark, s=soil, x=unknown. Sex distribution: Au=autoicous, Di=dioicous, Mo=monoicous, x=unknow.. 5. Pa=Pantropical, x=unknown. Clark ( 1 953) included also Frullania cucuata Lindnb. & Gott. and F. gymnotis Nes & Mon!.. .. 4 Distribution types: Aa=amphiatlantic, C=caribbean, Ca=central aerican, Co=cosÍnopoitan, E=endemic, Es=subnemic, Neo=neotroical, Nesa=NE outh Americi., Na=Northem South America,. 6 7. Clark 1953, Fosberg & Klawe 1 966 as F. cocosenis Steph. As. A. juliformis (Nees) Grads!..

(16) H H H H H H.. Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae .Lejeuneaceae. Ceratolejeunea Ceratolejeunea Ceratolejaunea Ceratolejeunea Ceratolejeunea Cheilolejeunea. cubensis (Mont.) Schiffn. lobata Clark maritima (Spruce) Steph. plumula (Spruce) Steph. ruginosa Steph. adnata (Kunze) Grolle. b x b,e e b,e e. Au X Au Au Au Di. Neo E Neo Neo C Neo. H H H H. Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae. Cheilolejeunea Cheilolejeunea Cololejeunea Cololejeunea. decurviloba (Steph. ) X. L. He rigidula (Mont.) Schust. cadiocarpa (Mont.) Steph. submarginata P. Tix.. b,e b,e e e. Di Di Mo Mo. Neo Aa Neo Neo. H H H. Lejeuneaceae Lejeuneaceae Lejeuneaceae. Colura Colura Cycloejeunea. clavigera Gott. ex Jovet-Ast tortifolia (Mont.) Steph. accedens (Gott.) A. Evans. b e e. Di Di Mo. C Neo Neo. H. Lejeuneaceae. Cycloejeunea. lb,e. Di. Neo. H H H H. Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae. Cycloejeunea Cycloejeunea Cystolejeunea Diplasiolejeunea. convexistipa (Lehm. & Lindenb.) A. Evans luteola (Spruce) Grolle peruviana Lehm. & Lindenb. lineata & Lehm. & Lindenb.) Herz. pellucida (Meissn.) Schiffn.. e lb,e lb,e e. Di? Di Di Di. Neo Neo C Neo. H H H H H H H H H H H. Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae. Drepanolejeunea Drepanolejeunea Drepanolejeunea Echinocolea Hygrolejeunea Lejeunea Lejeunea Lejeunea Lejeunea Lejeunea Lejeunea. crucianella (Tayl.) A. Evans inchoata (Meissn.) A. Evans lichenichola (Spruce) Steph. dilatata (A. Evans) Schust. ocellata Steph. cf. glaucescens Gott. cladogyna A. Evans flava (Sw.) Nees laetevirens Nees & Mont. ex Schiffn. phyllobola Nees & Mont. setiloba Spruce. h,e e b r x lb x e e b x. Di? Di? Di? Mo X Mo Mo Mo Di Au Mo?. Neo Neo Neo C , Ca E Pan neo Pan Neo Neo Neo. 8 9. Dauphin 1 078, 1 1 06/ Fulford 1 945 . Clark 19538 Dauphin 971/ Fulford 1 945 Bemecker & Lücking in prel Fulford 1 945. Dauphin 1 055, 1 076/ Fulford 1 945 Lücking 92-284 (USJ), Morales 1991/Schuster 1 980, Gradstein, pers cornm. Dauphin 1 068/ He 1 996. Dauphin 1 057/ Schuster 1 980 Bemecker & Lücking in prep.l Schuster 1 980 Bemecker & Lücking in prep/ Gradstein, pers . . Cornm. Dauphin 1 1 42/ pócs . 1 988. Bemecker & Lücking in prep.l Jovet-Ast 1953 Bemecker 92-328, Bemecker & Lücking in prep. Dauphin 926, 979/ Gradstein, pers. Cornm. Lücking 92-3 1 7 (USJ) Gradstein et al. 1 994 Dauphin 969/ Gradstein, pers. Cornm. Dauphin 972/ Gradestein, pers. Cornm. Bemecker & Lücking in prep.l Gradestein, pers. Cornm. Dauphin 965/ Bischler 1 964 Bemecker & Lücking in prep./Bischler 1 964. Dauphin A-7/ Bischler 1 964. Dauphin 1 003/ Gradestein et al. 1 994. Stephani 1 9 14, Clark 1 95319 Dauphin 1 046/ Schuster 1 980. Morales 1 9 9 1 / Schuster 1 980. Morales 1991/ Schuster 1 980. Bemecker & Lücking in prep.l Schuster 1 980. Dauphin 1 056/ Pocs 1988 10 Morales 1991/ Schuster 1 9 80.. After lhe aulhor it could be a ynonim of C. maritima (Spruce) Steph., known only from lhe original description. The type is not available at lhe GIS Herbarium.. The genus Hygrolejeunea is consiered a synonim of Lejeunea (e. g. Yano & Gradstein 1 997). Because of lhe ocellli, this laxon does not tit in lhe present concept of lhe genus Lejeunea (Gradstein 1 997).. Mizutani 1971 placed it as Pycnolejeunea cocosensis Mizul. After He (pers. Corom.) it oes not belong in lhat genus. Only recorded from Cocos Island. 10 As Rectolejeunea phyllobo/a (Nees & Monl.) A. Evans..

(17) H H H H. Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae. Lejeunea Lejeunea Lepidolejeunea Lepidolejeunea. sp. 1 sp. 2 involuta (Gottsche) Grone omata (Robins. ) R.M.Schust.. r e e b,e. X X Di Di. X X Neo Nsa, Ca. H. Lejeuneaceae. Leptolejeunea. elliptica (Lehm. & Lindenb. ) Schiffn.. e. Di, Au. Neo. H H. Lejeuneaceae Lejeuneaceae. Leptolejeunea Lopholejeunea. radicosa (Nees & Mont.) Grone eulopha (Tayl.) Schiffn.. e r, Ib. Di Au. C , Ca Pan. H. H H. Lejeuneaceae Lejeuneaceae Lejeuneaceae. Lopholejeunea Lopholejeunea Macrolejeunea. nigricans (Lindenb.) Schiffn. subfusca (Nees) Schiffn. cerina (Lehm. & Lindenb. ) Gradst.. x r x. Au, Di Au Di. Pan Pan Neo. H. Lejeuneaceae. Microlejeunea. acutifolia Steph.. e. Di. C. H H. Lejeuneaceae Lejeuneaceae. Microlejeunea Microlejeunea. bullata Tayl. epiphylla Bischi.. e e. Di Di. Neo C. H. Lejeuneaceae. Odontolejeunea. lunulata (F. Weber) Schiffn.. e. Mo. Aa. H. Lejeuneaceae. Omphalanthus. filiformis (Sw.) Nees. e. Di. Neo. H H H H H H H H H H H. Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae. Physantholejeunea portoricensis (Hame & Gott. ) Schust. Pictolejeunea picta (Grone ex Steph.) Grone Prionolejeunea cf. innovata A. Evans Prionolejeunea denticulata (Web. ) Schiffn. Prionolejeunea sp. sp. Pycnolejeunea Rectolejeunea bert�roana (Gottsche) A. Evans Rectolejeunea cf. emarginuliflora(Gott.) A. Evans Schiffneriolejeunea polycarpa (Nees) Gradst. ? Stictolejeunea squamata (Willd. ex Web. ) Schiffn. Symbiezidium14 barbiflorum (J,-indenb. & Gott) A. Evans. Lb lb h x e X b,e b b r b. ? Mo X Di X X Di Di Au, Di Di Mo. Ca Neo X Neo X X Neo C Pan Neo Neo. 1 1 C1ark 1 953 as L. cocosensis sp. nov. 12 C1ark 1953 as L. muelleriana (Gott.) 8 chiffn . 13 New to Costa Rica. 14 C1ark 1 953 reported other species now put under synonymy by Gradstein 1 994.. Dauphin s.n., Chatham. Bemecker & Lücking in prep. Dauphin 9921 Gradstein, pers. Comm. Dauphin A-71 Piippo 1 986, Gradstein et al. , 1 994) Bemecker & Lücking in prep.! Gradstein, pers. Comm. Bemecker & Lücking in prep./ Bischler 1 969 Dauphin 978, 1 1 39, Gradstein et al. 1 9941 Gradstein 1 9941 1 Gradstein 1 9941 2 Dauphin s.n.! Gradstiin 1 994. Morales 1 99 1 / Grone 1 956. I. Bemecker & Lücking in prepl Bischler et al. 1 96? Dauphin 1 1 871 Gradlltein & Weber 1 982. Bemecker & Lückin� in prep/. Gradstein, pers. Comm. Bemecker & Lücking in prep/. Gradstein 1 994. Bemecker & Lücking in prepl Gradstein & Webr 1 9 82. Dauphin 943a!31 Dauphin 1 099/ Grad�tein et al. 1 994. Dauphin 965 Dauphin A-71 Gradstein et al . 1 994. Dauphin 976. Weber 1 993 Dauphin 1 1431 Sch*ter 1 980. Dauphin 11 071 Gra�stein, pers. Comm. Dauphin A-51 Grads,tein 1 994. Dauphin 1 00 1 , 1 02 lt Gradstein 1 994. Dauphin 9 1 8, Mor¡¡f s 1 99 1/ Gradstein 1 994..

(18) H H H H H H h H H H H H H H H H H H H H H H H M M M M M. Lejeuneaceae Lejeuneaceae Lejeuneaceae Lejeuneaceae Lepidoziaceae Lepidoziaceae Lepidoziaceae Lepidoziaceae Lepidoziaceae Lepidoziaceae Lepidoziaceae Lepidoziaceae Lepidoziaceae Plagiochilaceae Plagiochilaceae Plagiochilaceae Plagiochilaceae Plagiochilaceae Plagiochilaceae Radulaceae Radulaceae Radulaceae Radulaceae Bartramiaceae Bartramiaceae Bartramiaceae Bryaceae Callicostaceae. Symbiezidium Symbiezidiwn Taxilejeunea Taxilejeunea Arachniopsis Bazzania Bazzania Bazzania Bazzania Bazzania Kurzia Micropterygium Zoopsidella Plagiochila Plagiochila Plagiochila Plagiochila Plagiochila Plagiochila Radula Radula Radula Radula Philonotis Philonotis Philonotis Bryum subg. Bryum Pilotrichum. dentatum Herz. transversale (Sw.) Trev. pterigonia (Lehm & Lindenb.) Schiffn. sp. diacantha (Mont.) Howe affinis (Lindenb. ) Trev. cubensis (Gott.) Pagán cuneistipula (Gott. & Lindenb. ) Trev. hookeri (Lindenb.) Trev. stolonifera (Swarz Trev. capillaris (Swarz) Grolle carinatum Steph. antillana (Steph.) Schust. bursata (Desv.) Lindenb. guilleminiana Nee & Mont. gymnocalycina (L. & L.) Mont. martiana Nees rutilans Lindenb. vincentina Lindenb. afinis Lindenb. & Gott. flaccida Lindenb. macrostachya Lindenb. & Gott. pusilla Spruce e/ongata (Dism.) Crum & Steere sphaericarpa (Hedw.) Brid. uncinata (Schwaegr.) Broth. sp. corrugatum Ren. & Cardo. lb b,e x b H b b b b b s b b x x b lb x h X e b,lb r r r r r h,r,lb,. Au, Di Mo, Di Mo X X Di Di Di Di Di Di Di Di Di DI Di Di Di Di Di Di Di Di Di Di Di Di? Di?. M M. Callicostaceae Callicostaceae. Pilotrichum Callicostella. rugifolium C. M . depressa (Sw. ) Jaeg.. lb rb. Di? Au?. M M. Callicostaceae Callicostaceae. Callicostella Callicostella. oerstediana (C. M. ) Jaeg. sp.. rb r. Au, Syn X. 1 5 Also as P. andersonii Aognst. 16 Fosberg & KIaw 1 9 66 as P. gracillima Aongstr; Weber 1 993.. 1 7 Williams 1 924 (as Pilotichum obtusatum sp. nov.), Fosberg &v KIawe 1 966, Crosby 1 969 (as P. corrugatum Ren. & Card.).. Dauphin 952/ Gradstein 1 994. Dauphin 9 1 9/ Gradstein 1 994. Morales 1991/ Gradstein & Weber 1 982. Dauphin 956, 1025. Dauphin A-5, 96Opp/ Gradstein et al. 1 994. Pan Dauphin 1 063/ Fulford 1 962. Ca, Nsa Dauphin 1 1 90/ Fulford 1 962. C Neo Dauphin A-6/ Fulford 1 962. Dauphin 970, 974/ Fulford 1 962. Neo Dauphin 1063, 1 1 05/ Gradstein, pers. Comm. C Dauphin 1 05 9/ Gradstein et al. 1 994. Neo Dauphin 96 1 / Fulford 1 966. C , Nea Dauphin 960/ Gradstein et al. 1 994 C, Ca Neo Morales 1991/ Gradstein, pers. Comm. Neo Morales 1 99 1 15/ Gradstein, pers. Comm. Neo Dauphin 987, Gradstein, pers. Comm. Neo Clark 1 953/ Gradstein, pers. Comm. Neo Dauphin 9 14, 925/ Gradstein, pers. Comm. Neo Dauphin 1 104/ Gradstein, pers. Comm. X Morales 1 99 1 . Aa Dauphin 1 2 3 1 / Schuster 1 980. Neo Dauphin 9 1 6 , 923/ Yamada 1 9 8 8 . Nsa Dauphin 1 0 1 3/ Gradstein, pers. Comm. Neo Dauphin 1 008, 1 1 93/ Griffin 1 994. Neo Dauphin 1 1 34/ Griffm 1 994. Pan Dauphin 1023, 1 1 23 16/ Griffm 1 994. X Dauphin 1 060. E Dáuphin 973, 997, 1 027, 928, 943 , 1 028, 11/ Crosby b 1 969 E Dauphin 943/ Crosby 1 969. C, Ca, Nsa Dauphin 937pp, 1 0 1 6pp, 1 125/Delgadillo et al. 1 995. Ca Dauphin 1 0 1 6pp/ Delgadillo et al. 1 995 . X Dauphin 1 040, 1 004, 1 007, 10 1 9 . Nsa Neo Neo.

(19) M M M M. Callicostaceae Callicostaceae Callicostaceae Callicostaceae. Crossomitrium Crossomitrium Hookeriopsis Hypnella. epiphyllum (Mitt.) C M. patrísiae (Brid.) CM. sp. pallescells (Hook.) Jaeg.. M M M M M M. Callicostaceae Callicostaceae CaJymperaceae Calymperaceae Calymperaceae Calymperaceae. Lepidopilum Thamniopsis Calymperes Calymperes Syrrhopodon Syrrhopodon. M M. Calymperaceae Calymperaceae. Syrrhopodon Syrrhopodon. M M M M M M M M M. Calymperaceae Dicranaceae Dicranaceae Dicranaceae Fissidentaceae Fissidentaceae Fissidentaceae Fissidentaceae Fissidentaceae Fissidentaceae Fissidentaceae Fissidentaceae Hypnaceae Hypnaceae Hypnaceae Leucobryaceae Leucobryaceae Leucobryaceae Leucobryaceae. Syrrlwpodon Campylopus Dicranella Dicanella Fissidens Fissidens Fissidens Fissidens Fissidens Fissidens Fissidens Fissidens Ctenidium Isopte/ygium Vesicularia Leucob/ywn Octoblepharum Octoblepharum Octoblepharum. crassisetum Willims cruegeriana (C M.) Buck afzeliii Sw. erosum C M. circinatus (Brid.9 Milt. incompletus Schwaegr. var berteroanus (Brid.) Reese lycopodioides (Brid.) CM. prolijer Schwaegr. vaL scaber (Mite) Reese rigidus Hook & Grev. savannarum (C M.) Mitt. harrisii (CM.) Broth. hilariana (Mont.) Mitt. angustifolilts Sull. elegans Brid. intramarginatus (Hampe) Jaeg. leptophyllus Mont. microcladus Twait & Mitt. neglectus Cmm pel/ucidus Hornsch. sp. sp. tenerum (Sw.) Mitt. vesicularis (Schwaegr.) Broth. martianum (Hornsch.) Hampe ex CM. albidwn Hedw. cocuiense Mitt pulvinatum (Doz. & Molk.) Mitt.. NI M M M M M M M M M. 18 Fosberg & KJawe J 966 as C. oerstedianum C.M.. Bartram 1 949 as C. donnellíi Aust 20 As F. garberi S. & L. j9. Di Di X X. Neo Neo X Ca, C, Nsa. b b b, lb,r b. Au? X Di, V Di Di Di. E X Pan Pan Neo Neo. Weber 1 993 1 8! DelgadilJo et aL 1 995. Dauphin 929! Delgadillo et al. 1 995. Dauphin 1095, 1 098. Fosberg & Klawe 1 966, Weber 1 993! Delgadillo et aL 1 995. Dauphin 994! Williams 1 924. Weber 1993. Dauphin s.n. 1 9! Reese 1 994. Weber 1 993! Reese 1994. Dauphin 9 1 5 , 1 1 88! Reese 1 994. William 1 924, Webef' l 993! Reese 1 994.. lb, b rb. Di Di. Neo Neo. Dauphin A-9, 1 2 1 1 1 Reese 1 994. Dauphin 10611 Reese 1 994.. b h. Di Di Di Di Au Au Au X X X Au X X Au Au Au Au Au Au. Neo Neo C, Ca Neo Neo Neo Neo Neo X Neo Pan X X Neo Neo Neo Pan Neo Neo. Dauphin 1 1 001 Reese 1 994. Dauphin 1 2231 AlIen 1 994. Dauphin 991! Allen 1 994. Dauphin 988, 1 1 201 Allen 1 994. Dauphin A-4! Purse1l 1 994. Dauphin A-4! Pursell 1 994. Dauphin 1 07 1 1 PurseJl 1 994. Dauphin 1 1 1 01 Pursell 1 994. Weber 1 99320 Dauphin 1 0091 Pursell 1 994. Dauphin 1065! Pursell 1 994. Dauphin 1 1 54. Dauphin 1 043. Dauphin 1 047, l OS l ! Ireland 1 ')92. Dauphin 1 1 26! Cm 1 994g. Dauphin s.n., 1 236/ Allen 1 994. Dauphin 1 0481 Salazar Allen 1 994. Dauphin 9621 Sal azar AlIen 1 994. Dauphin 933, 1 062/Salazar AlIen 1 994.. e lb, e x lb, b, r. r r. X. h h, rb r s� r rb b b.

(20) M M M M M. M. Leucobryaceae Leucomiaceae Meteoriaceae Meteoriaceae Neckeraceae Orthotrichaceae. Oetoblepharum Leueomium Squamidium Zelometeorium Neekeropsis Groutiella. M M M M M M M M M. Pottiaceae Rhizogoniaceae Sematophyllaceae Sematophyllaceae Sematophyllaceae Sematophyllaceae Sematophyllaceae Sematophyllaceae Thuidiacea. Hyophila Pyrrhobryum Aeroporium Glossadelphus Sematophyllum Sematophyllum Taxithelium Triehosteleum Cyrto-hypnum. stramineum Mitt. strumosum (Homsch. ) Mitt. isocladum (Ren. & Card.) Broth. patulum (Hedw.) Manuel undulata (Hedw.) Reichardt. mueronifolia (Hook. & Grev.) Crum & Steere sp. spiniforme (Hedw.) Mitt. pungens (Sw.) Broth. laevifolius (Mitt.) Bartr. galipense (C. M.) Mitt. subpinnatum (Brid.) Britt. planum (Brid.) Mitt. ef fluviale (Mitt.) Jaeg. involvens (Hedw.) Buck. lb x. lb lb, h lb b r rb lb, b r x. x b, lb, r rb x. Au Syn, Au Di Di? Syn Au X. Neo Pan Neo Neo Neo Neo X. Dauphin 1 0 8 1 / Salazar Allen 1 994. Williams 1 9242 1 / Crum 1 994. Dauphin 935/ Alen & Crosby 1 986. Dauphin 934, 938, 22/ Crum 1 994. Dauphin 9 1 7/ Smith 1 994. Dauphin 946, 1 1 94/ Vitt 1 99423 Dauphin 1 0 1 7 , 1 192.. Di Au Au Au Au Au Au Au. Pan Neo Neo Neo Neo Neo Ca, Nsa Neo. Dauphin 94 1 / Crum 1 994b. Dauphin 970, 975/ Magil 1 994. Dauphin 1 0 1 424/ Crum 1 994. Gómez 4568 (CR, MO)/ Magill 1 994. Weber 1 993. Dauphin 922, 1 04 1 , 932/ Crum 1 994. Dauphin 937pp./ Crum 1 994 Weber 1 99325•. 21 Fosberg & Klawe 1 966 as L. cuspidatifolium (C. M.) Mitt. 22 Fosberg & Klawe 1 966 as Meteoriopsis patula (Hedw.) Br o th.; Weber 1 993. 23 =Groutiella apiculata (Hook.) Crum & Steere. 24 WIlliams 1 924 as Hookeriopsis cocoensis sp. DOV; Bartram 1 949 as G. cocoensis (Williams) Bartra), Fosberg & Klawe 1 966 also as G. longisetus Bartr., Weber 1 993 as Taxiphyllum laevifolium (Mitl) W. R. Buck 25 Fosberg & Klawe 1 966 as Thuidium involvens Hedw..

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