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Distribution Patterns of Leucodon Species in Macaronesia, with Special Reference to the Canary Islands

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Distribution Patterns of Leucodon Species in Macaronesia, with Special Reference to the Canary Islands Cryptogamie,Bryologie,2009,30 (1):185-197 ©2009Adac.Tous droits réservés Distribution patternsof Leucodon speciesin Macaronesia, withspecialreferencetothe Canary Islands JuanaMaríaGONZÁLEZ-MANCEBOa*,JairoPATIÑOa ,OlafWERNERb , RosalinaMariade AlmeidaGABRIELc &RosaMaríaROSb a DepartamentodeBiologíaVegetal(Botánica),UniversidaddeLa Laguna, C/AstrofísicoFranciscoSánchezs/n,38071-La Laguna, Tenerife,IslasCanarias, Spain,[email protected]; [email protected] b DepartamentodeBiologíaVegetal(Botánica),UniversidaddeMurcia, Campus de Espinardo,30100-Murcia, Spain,[email protected]; [email protected] c CITAA -DepartamentodeCienciasAgrárias,Universidade dosAzores, P-9701-851 TerraChã-Angrado Heroísmo,Azores,Portugal,[email protected] (Received 10July2008,accepted 10November2008) Abstract –The genus Leucodon isrepresented in Macaronesiabythree species, L.sciuroides , L.canariensis and L.treleasei,the lattertwobeing endemictothisregion. An analysisof distribution,frequencyand habitats forall three speciesin thisregion shows singularpatternsforeachspecies,mainlyrelated tohabitatconservation and type of habitat. Leucodon canariensis isthe most restricted speciesasregardshabitatconditions,witha confirmed presenceonlyon Madeiraand the Canary Islands,although moreabundantin the latter. Leucodon treleasei exhibits the widest habitatamplitude,especiallyon Madeira, and itispresentin all three northernarchipelagos. Leucodon sciuroides isthe most tolerant speciestoaridity and ismainlyfound on Madeiraand the Canary Islands,although its occurrencewasconfirmed in all the Macaronesianarchipelagos. Leucodon /Leucodon sciuroides / Leucodon canariensis / Leucodon treleasei /Macaronesia /biogeography/conservation /habitatrequirements INTRODUCTION According toseveralauthors (e.g. Hansen &Sunding,1993),the MacaronesianRegion includesfour archipelagos,the Azores,Madeira, Canary Islandsand Cape Verde Islands.Nevertheless,the conceptof Macaronesiathat best appliestobryophytesmightbedifferent,atleast in the caseofmosses,since the Cape Verde Islandsshowhigheraffinitieswithtropicalmoss flora (Vanderpoorten etal .,2007; González-Mancebo etal .,2008b). Eitherin awider orstrictersense,Macaronesia, isone of the 25world biodiversity hotspots (Myers etal .,2000),and within the EuropeanMediterraneanregion,itisone of the most importantfloristicareas(Médail &Quézel,1997; Vanderpoorten &Long,2006). *Correspondenceand reprints :[email protected] 186J.M.González-Mancebo,J.Patiño,O.Werner,R.M.deA.Gabriel &R.M.Ros Macaronesiaisalsowell-knownforexhibiting ahigh degree of endemism, especiallyasfarasvascularplants areconcerned (e.g. Kim etal .,2008;Reyes- Betancort etal .,2008). Asregardsbryophytes,a10%rateofendemicity isnot high compared withotherisland groups.However,within the EuroAsiatic- MediterraneanRegion,Macaronesiaisespeciallyrichinendemicbryophytes (Bischler,2004),although endemicity isquitelowamong the Azoreanbryophytes (only2.05%,Gabriel etal .,2005). The genus Leucodon includes38species(Crosby etal .,1999) with analmost worldwide distribution in temperateand warmregions.Inthe Macaronesianregion,thisgenus includesthree species: Leucodon sciuroides (Hedw.) Schwägr., L.canariensis (Brid.) Schwägr.,and L.treleasei (Cardot)Paris, the last twobeing Macaronesianendemics. Leucodon sciuroides isawidespread speciesof the Palaearcticregion. InEurope,itrangesfrom Iceland and northern Scandinavia(Nyholm,1954-1960; Pippo,1982)tothe Mediterraneanregion, whereitisadominantepiphyteonthe coastalplains(Preston,1994). Ithasalso been recorded in NorthernAfrica (Ros etal .,1999) and East Africa (O’Shea, 2006)and in all the Macaronesianarchipelagos(Hedenäs,1992; Patiño Llorente &GonzálezMancebo,2005). Fordecades,identification problemsprevented toknowthe real distribution of thesethree species. Leucodon treleasei waseven considered asa synonymof L.canariensis byseveralauthors in the past (Corley etal .,1981; Dirkse etal .,1993). Thistaxonomicdifficulty wasovercome byHedenäs(1992), who clearlyestablished the morphologicaldifferencesbetween the twoendemic speciesand L.sciuroides .Thisauthorindicated thatthe occurrenceof L.canariensis in the Azoresisuncertain,and described the habitat,distribution and frequencyof thesespecieson the island of Madeira.However,these characteristicshavenotbeen studied in otherMacaronesianarchipelagos. The presentworkincludesadetailed revision of thesespecies,based on herbariumand freshmaterialfrom all the Macaronesianarchipelagos.Differences in habitat,distribution and frequencyof Leucodon specieswereanalysed in the Canary Islands,Madeira, the Azoresand the Cape Verde Islands,withspecial focus on the first archipelago,wherethe three speciesoccur.Afrequencyanalysis wasalsoperformed,in ordertogeneratehypothesesabout the centreofthe dispersalrange foreachofthe endemicspecies,taking intoconsideration that theyoccur most abundantlyin the centreoftheirrange and thattheirabundance graduallydeclinestowardstheirgeographicallimits (e.g. Söderström,1989). MATERIAL AND METHODS Bibliographyreferences The recordsof Leucodon speciesfrom the four Macaronesian archipelagoshavebeen thoroughlycompiled foreachofthe four archipelagos. Forthe Azoresthe following referenceshavebeen used:Russel (1862),Cardot (1897,1905),Trelease(1897),Geheeb&Herzog (1910),Allorge &Allorge (1945, 1948,1952),Schwab (1981),Eggers (1982),Hübschmann (1974),Sjögren (1990, 1996,1997,2001,2003,2005),Fontinha&Sérgio (1995). ForMadeira: Eggers (1982),Hedenäs(1992),Sérgio etal .(1992),Sjögren (2001) and Kürschner etal . (2007a, b). Forthe Canary Islands:Pitard&Negri(1907),Bryhn (1908),Bines Leucodon in Macaronesia187 (1965),Düll (1980),During (1981),Losada-Lima etal .(1984),Schwab &Haustein (1984),Koppe &Düll (1986),Malme (1988),Zippel (1998),González-Mancebo et al. (1989,1996,2004,2008a). Forthe Cape Verde Islands:Potierde laVarde (1946),Frahm etal .(1996),Patiño Llorente&GonzálezMancebo(2005),O’Shea (2006). Withthesedata, apreliminary approachwasmade toestimatethe occurrenceand frequencyof the specieson eachisland. Onlyislandsmeasuring morethan50km2 wereconsidered,withthe single exception of Corvo(15 km2 , the Azores),duetoits high altitude (718 ma.s.l.). Thus,Selvagens,Deserta (Madeira),and the islets of Graciosa, Alegranzaand MontañaClara(Canary Islands)werenotconsidered. Plantmaterial The characters considered byHedenäs(1992)wereusefulfor establishing morphologicaldifferencesamong thesethree species,especiallyas regardscapsule shape and the sizeofthe middle-leafcellsin the secondary stems, aswell asthe cell shape of the cellssituated between the baseand middle leafat the leafmargin. DNA data(manuscriptin preparation) confirmed thatthe morphologicalcharacters used byHedenäsreliablyidentifythe three species. Plantmaterialfrom most of the areasreferred tointhe abovebibliographywas studied. Forthis,all classicallocalitieswerevisited,including thosewherethe recordsof the specieswereuncertain,eitherduetothe erroneous identification of herbariaspecimensorbecauseofdoubtconcerning habitataccording tothe main distribution signsof the species.Atotalof500 freshand 50herbarium specimensfrom the differentarchipelagoswereidentified,whichallowed to correcting the first bibliographicapproachonthe distribution and frequencyof the species. Habitatcharacteristics Forthe analysisof speciesdistribution,severalhabitatcharacteristics wereconsidered:altitudinalrange,type of habitatand habitatamplitude. Fivetypesof habitathavebeen distinguished:1. sun-exposed habitats (including cultivated and disturbed areasand alsonon-forestry natural vegetation);2.sheltered habitats outside the forest; 3.laurel forest (evergreen subtropicalforest);4. cloudlaurel forest (laurel forest withthe highest fog influence,mainlysituated along the mountain ridges);5. pine forest. Fivedifferentclassesof habitatamplitude havebeen considered: 1. unknown(old reports without precision);2.very restricted (presenceonlyin one ortwolocalities; maximumtwosquaresof 1km 2 );3.restricted disperse (restricted toone habitatwhereitisnotcommon);4. restricted locallyabundant (restricted toone habitatwherethe speciescancolonizedifferentmicrohabitats and reachahigh frequencyand/orbiomass);5. common (widelydistributed in severalhabitats and wide altitudinalrange). Dataanalysis Detrended correspondenceanalyses(DCA;Hill &Gauch,1980)have been used tostudyisland and speciescomposition forall the Macaronesian islands.Thesewereperformed withthe CANOCO package (TerBraak& 188 J.M.González-Mancebo,J.Patiño,O.Werner,R.M.deA.Gabriel &R.M.Ros Smilauer,2002),entering datacorresponding tofrequencyof the specieson each island. The frequencyforeachspecieson eachisland wasestimated counting the numberof 1km 2 cellswherethe specieshadbeen found,taking intoaccountour owncollectionsand alsoconfirmed recordsof otherauthors. RESULTS Dataof distribution,habitatand frequencyof Leucodon speciesin the MacaronesianRegion arepresented byeacharchipelago (Table 1). The Azores Leucodon canariensis hasbeen reported from SãoMiguel byTrelease (1897),Geheeb(1910),Allorge &Allorge (1945),and Hübschmann (1974). During our field workinthisarchipelago,only L.treleasei wasfound in the areas wherethe specieshadbeen recorded. The specimensreported bySchwab (1981) from Faialand SãoMiguel werelaterre-examined bythisauthorand re-identified as L.treleasei (Frahm, personalcommunication). The recordofSjögren (1990) from Graciosarefers to L.treleasei asitwasconsidered asynonymof L.canariensis .Consequently,the presenceofthisspeciesin the Azoresshould be considered uncertain,aswasalsopointed out byHedenäs(1992);forthisreason ithasbeen omitted in
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