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Sans Titre-1 Cryptogamie,Bryologie,2009,30 (1):207-215 ©2009Adac.Tous droits réservés Noteworthyrange extensionsof two Aneura (Jungermanniopsida, Metzgeriales)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 Aneura pinguis (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 plants 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
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