Cryptogamie,Bryologie,2009,30 (1):207-215 ©2009Adac.Tous droits réservés

Noteworthyrange extensionsof two Aneura (, )species newforthe IberianPeninsula: Aneuramaxima (Schiffn.) Steph. and A.pseudopinguis (Herzog) Pócs

CecíliaSÉRGIO* &CésarGARCIA

Universidade de Lisboa, MuseuNacionaldeHistóriaNatural, Jardim Botânico/CentrodeBiologiaAmbiental, RuadaEscolaPolitécnica, 58,1250-102 Lisboa, Portugal

(Received 2008,accepted 31October2008)

Abstract –The recognition of Aneuramaxima (Schiffn.) Steph. in IberianPeninsulaand the discovery of Aneurapseudopinguis (Herzog) Pócsin Portuguesebryofloraaregiven. The identity of twomisunderstood and unknown Aneura from IberianPeninsulaisclarified based in herbariummaterial. The morpho-anatomicalobservationscarried out in herbarium samplesaresummarized in atable and illustrated. The geographicdistribution of the two speciesisoffered forthe IberianPeninsula. A.pseudopinguis isalsoreported newtothe WesternAfrica.

Aneura /Hepaticae/IberianPeninsula/Portugal/Spain /WesternAfrica

INTRODUCTION

During the identification of Aneura materialstored in LISU collections, twointeresting discoveriesweremade:the recognition of Aneuramaxima (Schiffn.) Steph. in the IberianPeninsulaand the discovery of Aneura pseudopinguis (Herzog) Pócsin the Portuguesebryoflora. Aneuramaxima was reported forthe first time in Europe during the 1990’sand its presencewastobe expected in the Iberianflora. Aneurapseudopinguis wasrestricted toCentral Africa and TropicalAmerica and its presenceinthe Iberianflorawasnotso predictable. Inthe IberianPeninsula, distribution dataof Aneura arescarcenot including in floristicworksasin Basqueregion (Infante,2000)and sofarthereis no taxonomicrevision. So,thispaperpresents the first taxonomicrevision based in selected herbariummaterial,withthe aim todetectthe presenceof Aneura maxima in the Iberianterritory. Finally,weintend togiveanaccuratecharacterization of the tree species, although thisdatawill becompleted withnewfield workinthe nearfuture.

*Correspondenceanreprints:[email protected] 208C.Sérgio &C.Garcia

MATERIAL AND METHODS

All herbariummaterialstored in LISU, COI and aselection of (L.) Dumort.specimensfrom Spain in BCB and MUB collectionswas studied using recenttaxonomiccriteriaforthisgenus.Toconfirmthe discovery of A.pseudopinguis ,certain collectionsatLISU and COI from SãoTomé e Principe and Angolawereconsidered and studied. So,the morpho-anatomical studieswerecarried out onlybased in herbariumsamples.Atotalofmorethan 50dried herbariumspecimenswerestudied (list of specimensannexed). Each specimen wasexamined,measured and some sectionsphotographed. From eachsample,normallythree thalli and atleast ten sectionswereobserved. Thallus sectionsweremade in the middle part of the thalli supported bythe largest and/orfertile of the specimen. Asoil bodiesin dried plants disappearin ashort period of time theywerenotconsidered in our observations.Weconsidered the specimen sexualcondition,howeverthe plants arefrequentlywithout sporophytes,sowehaveobserved onlyvegetative characters.

RESULTS

Aneuramaxima (Schiffn.) Steph. (Figs1-4) Aneuramaxima isaspecieswithanAsian-EasternNorthAmerican distribution,and itwasdiscovered in Europe Belgiumonlyin the 1990’sby Andriessen etal. (1995). Its knownrange hasbeen extended sincethen into differentEuropeancountries:France(Sotiaux &Sotiaux,1996),Finland (Frahm, 1997),Denmark(Thingsgaard,2002),Luxemburg(Werner,2003),Czech Republic(Ku ª era, 2004) and Poland (Buczkowska&Baczkiewicz,2006). This expansion reflects the increaseinthe taxonomicknowledge sincethe first report in Belgium. So, Aneuramaxima presentascattered,but quitewide distribution: wasfound outside NorthAmerica (Darigo,2004) from East AsiatoJapan (Furuki,1991). The Europeanrecordsarepredominantlyfrom Poland (Wachowiak etal., 2007)nevertheless itwasuntil recentlyonlyrecorded from Finland toFrance(Vanderpoorten etal., 2006). Therefore,one obvious reason for the fewknownlocalitiesof Aneuramaxima isthe rarity of fertile plants,asitisa dioicous species. Bearing in mind thatmost morphologicalcharacters arevariable in Aneura taxa, thisrenders the identification between the twoEuropeanspecies sometimesdifficult.The explanation forthe high variability of Aneurapinguis waspresented withthe prediction of crypticspeciation in fivedistantregions in Poland (Wachowiak etal., 2007). The plasticity of the speciesasanaquatic plantalsocontributestothisvariability,although thisstudyindicatesthatthe A.pinguis complexishighlydifferentiated atmolecularlevel. Recentlynew results of DNA analysisconfirmthe division of four crypticspeciesof Aneura pinguis in Europe,and alsoseparateitfrom Aneuramaxima (Baczkiewicz etal., 2008). Inrecentworks, A.maxima isconsidered “conservation dependent”to “nearthreatened” within the category “lowerrisk” bryophyte(Vanderpoorten etal., 2006)but itseemsnotvery infrequentin Atlanticareasof Europe. Two Aneura speciesnewtothe IberianPeninsula209

Figs1-13. 1-4. Aneurapinguis (L.) Dumort.; 5-8. Aneuramaxima (Schiffn.) Steph.; 9-13 . Aneura pseudopinguis (Herzog) Pócs.1,5and 9,cross sectionsof thallus; 2,6and 10,cross sectionsof wing margin;4,8and 13,cross sectionsof medianpart of thallus; 3,7,11 and 12,cross sections of wing (± 1/3). 210C.Sérgio &C.Garcia

Tothismoment Aneuramaxima seemsmuchmoreextended in Portugal thaninSpain asin atotalof14Spanishspecimensstudied only5correspond to thisspecies.All others are Aneurapinguis .Datasuggest ascattered distribution in Spain in relation toPortugalashavefound 13specimensof A.maxima to14 A.pinguis .Thismaybeduetothe absenceofsporedispersal,asthe sporophytes areabsent,makesinfrequent A.maxima . Ingeneral Aneuramaxima hasanAtlanticdistribution and arecent modeling distribution in Belgiumisnoteworthy(Vanderpoorten etal., 2006). HoweverThingsgaard(2002)hasincluded thisliverwort in aboreal-montane speciesassemblage.

Aneurapseudopinguis (Herzog) Pócs (Figs5-8) The identification using Jones’scriteriaforAfricanflora(e.g. Wigginton, 2004a)and withour ownknowledge of Africanmaterial(SãoTomé Island) led tothe recognition of thesespecimensas A.pseudopinguis, aliverwort newto Portugaland the Europeanbryoflora.The thallus sectionsof Portugueseplants appeartohavevery fewmediancellsthickinthe thalli,featuresthatarevery distinctfrom bothspeciesof Aneura in Europe. Although gatheringsof this speciesaresomewhatinsufficientand some onlywithfewplants topermit observation of sexualfeatures(onlyfemale plants),the thallus sectionsaretotally similarto A.pseudopinguis from Africa. According toMeenks&Pócs(1985), Aneurapseudopinguis isclearly related to A.pinguis but the differencesbetween the twospeciesareaccepted as significantasvery fewmediancellsthickinthe thalli. However,some characters correspond tofertile plants,male lobesand antheridiumnumberand spore size. MorerecentlyPerold (2001) hasconsidered thatstudied plants from South Africa are Aneurapinguis, but forthisauthorisnotmeanttoimplythat A.pseudopinguis notoccurs in thisarea.Gradstein &daCosta(2003)consider alsothe twospeciesasindependenttaxa( A.pseudopinguis and A.pinguis ) . According totheseauthors,the first diagnosticdifferencesarerelated tothe structureofthe sterile thallus. Inreality the distribution mapofthisspeciespresented byGradstein etal. (1983)iscompared withothertropicalAfro-Americanelements,but this distribution mapdoesnotinclude WesternAfrica.However,atpresentwecan confirmthe presenceof A.pseudopinguis in Angolaand in SãoTomé Island (see list of studied material),aswell asextend the occurrenceofthisliverwort to southwesternpart of Europe (Fig. 15). According Wigginton (2004b)the African countriesof occurrenceof A.pseudopinguis are:Ivory Coast,Lesotho,Malawi, Rio Muni,SouthAfrica, Seychelles,Tanzania, Uganda, DemocraticRepublicof the Congo and Zimbabwe. Despitethe apparentdifficultin examining dry Aneura specimens,we haveobserved thatthe tree specieshavemorphologicaldifferencesthatsupport the maintenanceoftheirspecificstatus.Anupdated synthesisof the main diagnosticcharacters of the tree speciesisnowgiven,including illustrationsand dataon theirpossible ecologyand distribution. Aneurapinguis , A.maxima and A.pseudopinguis presentsimplicity on thallus organization withlittle anatomicdifferentiation,usuallywithsome distinction between wingsand midribornerve(the epithet Aneura indicatesthe absenceofanerve) but the thallus sectionsarequitedifferent(Table 1). Wehavefound astrong correlation between the featuresof sterile thalli and the dorsalsurfacecellsof the thallus in Aneurapseudopinguis .Inthisspecies Two Aneura speciesnewtothe IberianPeninsula211

Table 1. Differentialmorphologicalcharacters between Aneurapinguis , A.maxima and A.pseudopinguis (based in Iberianmaterial),fertility and distribution patternbased in herbariumspecimens(Figs1-13and 14-15).

Thallus characters AneurapinguisAneuramaximaAneurapseudopinguis

Thallus cross section Thallus nottranslucent, Thallus margins Thallus regularly (clearness,form) without anapparent translucent,withclear translucent,very thin,with centralmidrib, margin centralmidrib, plano- anincipientmidrib, plano-convextobiconvex convextoconcavo-convex concavo-convexorplane withwingsweakly withwingsweakly withwingsweaklyundulate recurved orplane. incurved. toflattened.

Numberof middle (9) 10-13(15) (8) 10-12(14) (5) 6-9 (10) cellsthick

Wings Becoming gradually Markedlythinnerto Becoming graduallythinner (formand number thinnertomargin;margin margin;margin acute tomargin;margin acute of bordercells) obtuseinsection toacute, skinnybordered by2-4 skinnybordered bordered by1-2single uni- single uni-stratoserow by2-4 single uni-stratose stratoserow,and wing and wing totally2,3to rowand wing totally2,3to with2,3to4stratus close 4stratosefrom the margin 4stratosefrom the margin tothe margin; tonearthe centralpart; tonearthe centralpart; wing less than10cells wing large morethan20 to wing imperceptible as wide. 25ormorecellswide. whole thallus isthin.

Epidermalcellsin Quadrangular, Quadrangular,iso- Rectangular,notiso- section isodiametricwithfirm diametricwithfirmnot diametric, withflexible walls,notcollapsed. collapsed walls. collapsed walls.

Fertility Sometimesfertile Neverfertile and spores Neverfertile and sporesnot in Portuguese developing capsulaand notobserved. Female observed. populations spores(near20%of plants apparentlymore Female plants apparently observed specimens). frequent. morefrequent.(InAfrican Female plants apparently specimensmale plants are morefrequent. morefrequent).

Distribution Itseemstobewidespread Itappears tobespreadin Mainlyin lowland, in the Iberian in mountain zonesin mountain areasin the ascending toatleast 800 in Peninsula Iberianterritory Northernhalf part of the centralWesternpart of ascending to3300 m,but Peninsularanging from Portugal. Notin downto20 minAtlantic 200 to2300 m. Mediterraneanenclaves. areas.

(numberof (Portugal14,Spain 9) (Portugal13,Spain 5) (Portugal12,Spain 0) specimensstudied)

theyare,in section,rectangularand notisodiametricwithflexible walls,whereas in A.maxima and A.pinguis arequadrangular,isodiametricand withfirmwalls, notcollapsed (Figs1-13). The morpho-anatomicalobservationscarried out in herbariumIberian samplesaresummarized and illustrated in table 1and figures1-13. 212C.Sérgio &C.Garcia

Figs14-15. Iberiandistribution of: 14. Aneuramaxima (Schiffn.) Steph.; 15. Aneurapseudo- pinguis (Herzog) Pócs.

DISCUSSION

Asexplained above,the morphologicaldifferencesamong the three speciesarerelativelyeasy toobserveand itisnotcrucialtohavefertile plants to distinguisheachone. Howeverthisscheme formorpho-anatomicalcharacters based onlyin herbariummaterialneedstobesupported byfreshand if possible byfertile plants.The relationshipswithothertaxaincluded within the synonymes of A.maxima need toberecognized. Itisthe example of Aneurapellioides (Horik.) Inouefrom Japan,included bySchuster(1992)inthe same complexof A.maxima. Two Aneura speciesnewtothe IberianPeninsula213

Weconsiderthatthe taxaarrangementof Aneurapinguiss.lat .requires aworld-wide study,notonlywithmolecularstudiesusing Europeanpopulations (Baczkiewicz etal., 2008),but togetherwithAfricanand Americanmaterial,to identifygeneticsimilarity and its phylogeography. The Iberianpopulationsof Aneurapseudopinguis areconsiderably disjunctfrom thoseinAfrica and SouthAmerica.Thus,theyareapparently geographicallyisolated,but thismaymerelyreflectour poorknowledge of the hepaticfloraof NorthernAfrica and/oralsothe Mediterraneanregion. Onthe otherhand,apossible explanation forthe currentdistribution of A.pseudopinguis in Portugalsuggests thatthe populationssurvived in the areaasrefugiaduring the glacialepisodes.Ineffect,numerous disjuncttermophyticspeciesoccur in habitats thatwould likelyhavebeen available in suchperiglacialrefugia. Infact,the localitieswhere Aneurapseudopinguis wasfound correspond toareaswithastrong Atlanticinfluence,withmoderatetemperaturesand alarge amountof humidity. Wecanpointout some bryophyteswithsimilardistribution in Portugal suchas Dumortierahirsuta (Sw.) Nees, Lejeuneaeckloniana Lindenb., Asterella africana (Mont.) A.Evans.and Racomitriumlamprocarpum (Müll.Hal.) A.Jaeger. So Aneurapseudopinguis ,recognized asaEuropeantaxon,needstobeincluded in analysestoconfirmthe similarity between differenttaxaof the complex A.pinguis and tosupport the possible Iberianvegetation histories.

Selection of studied specimens(IberianPeninsulaand WesternAfrica) Aneuramaxima (Schiffn.) Steph. Portugal BeiraAlta :S.PedrodoSul,Rio Sul,2km ajusantedaVila, 08.1949, E.Mendes1209 , LISU221079;SerradaEstrela, estradaparaoPoçodoInferno,1020 m,29TPE2572, 26.07.1995, C.Sérgio &J.Jansen 10134 ,LISU176458;SerradaEstrela, Fontes.,1430, 29TPE1470,29.06.1996, J.Jansen 96-074 ,LISU177915;SerradaEstrela, Manteigas,entrea SenhoradosVerdeseosCovais,Quedade águajuntodapontedoAceiro,1050m, 29TPE2371,02.07.2000, C.Garcia ,LISU180996.–BeiraLitoral :Coimbra, Corrente, 01.03.1968, J.Ormonde 431,COI10000005;EntreÁguedaeCaramulo,pr.rio do Águeda, 22.02.1967, J.Ormonde 438 ,COI10000004. – Minho:Arcosde Valdevez,Rio Vez, 29TNG4541,16.07.2004, A.Albuquerque&Rodriguez-González ,LISU147533; Serrade CastroLaboreiro,07.1949, E.Mendes1134 ,LISU221063; Serrado Gerês,Albergaria, 700 m,NG72,1982, C.Sérgio 4046 ,LISU153693; Serrado Gerês,ParqueTude de Sousa, 350m,1948, I.Tavares ,LISU147533; Serrado Gerês,próximo daBouça daMó,600 m, 1949, C.Tavares ,LISU147535;Serrado Gerês,RibeiradaLage,estradaparaLeonte, 750m,29TNG72,1984, C.Sérgio &R.Schumacker5330,LISU153695. Spain Jaén:Sierrade Cazorla, BarrancodelaCanal(Quesada),1600 m,WG0382,04.1996, R.M.Ros&R.Monreal ,MUB14378. – Salamanca:EntreelMaíllo yMonsagro,29TQE38- 39,1984, Elías ,BCB19635. – Zaragoza :Sierradel Moncayo,ParquedeAgramonte,1100 m, 30TWM92,02.06.1982 ,R.M.Ros ,MUB6257.–Álava :Eguileta, puertodeAzáceta, 800 m, 30TWN3937,14.01.1986, P.Hera s,MUB1686.

Aneurapinguis (L.) Dumort. Portugal BeiraAlta :SerradaEstrela, CasaldoRei,400-450m,29TPE0663,25.01.1995, C.Sérgio, M.Brugués&R.M.Cros, LISU180107; SerradaEstrela, descidado Vale do Zêzerepara Manteigas,1300m,29TPE2373,25.07.1995, C.Sérgio &J.Jansen 10113 ,LISU176435;Serra daEstrela, FontedaMariaSamarra, spring with Chrysosplenium ,860 m,29TPE1273, 214 C.Sérgio &C.Garcia

27.07.1995, J.Jansen 95-032,LISU177366; SerradaEstrela, Rapa, 600-650m,29TPE4093, 01.03.2000, C.Garcia&C.Sérgio,LISU181621;SerradaEstrela, from S.Romão,just after PontedeJugais,spring with Chrysosplenium ,550m,29TPE0971,27.07.1995, J.Jansen 95- 033,LISU177373.–BeiraLitoral :Matade Foja, 06.1880, Moller129, COI10000010; Ribeira Fria, entreAlbergaria-a-VelhaeS.JoãodeLoure,23.04.1965, A.Fernandes,J.Paiva& Cardoso55 ,COI10000008. – Minho:Serrado Gerês,a1kmdeAlbergaria, estradaparaa GeiraRomana, 650m,29TNG72,1984, C.Sérgio &R.Schumacker5392 ,LISU153694; Serrado Gerês,entreAlbergariaeBouça daMó,650.m,04.1949, E.Mendes401 , LISU221070; Serrado Gerês,FontedoRio do Forno,04.1949, E.Mendes540and 532, LISU221065and LISU2210766.–Trás-os-MonteseAltoDouro :Serrade Montesinho, Bragança, Vale daCoroa, 04.1953, E.Mendes&C.Romariz1414,LISU2210769. Spain Alicante :Benifallim,SerradelsPlans,Barrancde Clots,750m,YH2482,25.02.1993, M.J.Cano &R.M.Ros ,MUB16982&MUB5900; Callosad'en Sarrià, Fonts del riuAlgar, Fontdel Moro,350m,YH58,08.02.1983, Brugués,Cros,Sérgio &Sim-Sim 21.1, LISU149834. – Asturias :Ballota, 29TQJ12,1982 ,C.Casas ,BCB34268. – Cantabria : Lamasón,Ventade Fresnedo,30TUN79,200 m,1988, Aedo BCB45139;Camaleño,Vegade Tarna, 30TUN56,1600 m,1994, Casas,Cros&Brugués ,BCB40977.–Granada :Sierra Nevada, nacimientoRío Veleta, 30SVG60, Cros,Brugués&Sérgio,BCB52501. – León: Ancaresde León,Valle del Cuiña, PH84,1100 m,1984, R.M.Cros&Lloret ,BCB19800.– Lérida :Vall d'Aran,Salardú,Vall de Ruda, 1850m,31TCH32,05.09.1988, C.Sérgio etal. 6491,LISU155032.–Tarragona :Serrade Prades,Alforja, 31TCF36,1952, C.Casas BCB18325;ElsReguers,Barrancde laCaramella, 290-480m,BF61,05.06.1984,Casas , Sérgio,Brugués&Cros 21.1,LISU150404.

Aneurapseudopinguis (Schiffn.) Steph. Portugal BeiraAlta :SerradaEstrela, Senhorado Desterro,juntoaoAlva, 750m,1952, C.Tavares , LISU147534;SerradaEstrela.Senhorado Desterro,próximo daPontedaGaia, no vale do Alva, ±800 m,29TPE1072,08.1954, C.Tavares ,LISU183386.–BeiraLitoral :Entre PessegueirodoVougaeSeverdo Vouga, 02.12.1967, C.Sérgio 418,COI10000006; Severdo Vouga, SilvaEscura, 29.12.1969, C.Sérgio 879 ,COI10000003; SerradaLousã, Candal, 01.1950, E.Mendes1107,LISU221064;Zombaria, pr.Coimbra, 1.01.1879, Henriques37 , COI10000009(aff. Aneurapseudopinguis (Schiffn.) Steph.). – DouroLitoral :Próximo de Cinfães,RibeiradasBichas,próximo de Baião,09.1945, F.Resende 1495,LISU221072; Serrade Montemuro,próximo de Alvarenga, 400 m,07.1949, E.Mendes1200,LISU221071. – Estremadura :Azenhasdo Mar,arribasjuntoàpraia, 24.01.1999, C.Garcia ,LISU225896.– Minho:Serrado Gerês,10.1879, M.Ferreira ,COI10000011;Serrado Gerês,entre AlbergariaeBouça daMó,04.1949, E . Mendes515,LISU221068;Serrado Gerês,Pontede Bacelo,04.1949, E.Mendes372,LISU221067. SãoTomé ePríncipe NovoDestino,analtitude about 2500 ft.,Welwitsch267,LISU;NovaMoka, 900 m,1885, A.Moller 57,COI;PicadaparaaLagoaAmélia, 1400 m,1975, A.Sérgio 94,LISU;estrada paraBombaim,juntoàestradaparaapicadade Zampalma, 700 m,1975, A.Sérgio 36, LISU;Descidado PicodeSãoTomé,caminho paraPontaFigo,1109m.5.07.2007, C.Garcia Lev.79,LISU;Descidado PicodeSãoTomé,caminho paraPontaFigo,802 m,5.07.2007, C.Garcia Lev.83,LISU. Angola Huila, 1887, F.Newton,COI. Acknowledgements. WearegratefultoM.Bruguésand R.MCrosaswell as J.Guerraforthe loanofspecimensatBCB and MUB.SpecialthanksareduetoJ.Ku ª era, who confirmssome Portuguesematerialof Aneuramaxima .JoãoVilhenaisthanked forhis helpfulcomments on the Englishmanuscript.Weextend our gratitude toAnaBelén Barrón forthe illustrationsof thallus sections.Thisstudywasin part financed bythe Project“Floraicartografiade Briofits”PT2008-S0213of the Institut d’EstudisCatalans, the projectPOCTI/AFR/58699/2004and the grant(SFRH/BPD/22304/2005) of FCT (FundaçãoparaaCiênciaeaTecnologia). Two Aneura speciesnewtothe IberianPeninsula215

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