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DNA-Based Identification of Herbertus Species on Gough Island, South

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DNA-Based Identification of Herbertus Species on Gough Island, South Cryptogamie,Bryologie,2010,31(1):67-74 ©2010Adac.Tous droits réservés DNA-based identification of Herbertus species on Gough Island,SouthAtlanticOcean Jochen HEINRICHSa*,Kathrin FELDBERGa ,Hans-PeterKREIERa ,Ji ÷ íVÁ~ A b a GeorgAugust University,Departmentof SystematicBotany,Albrechtvon Haller InstituteofPlantSciences,UntereKarspüle 2,37073 Göttingen,Germany. b CharlesUniversity,Departmentof Botany,Benátská2,12801Praha2, CzechRepublic. (Received 6November2009,accepted 3December2009) Abstract –Based on phylogeneticanalysesof nrITS sequencesand accompanying morphologicalinvestigations,severalaccessionsof Herbertus from the SouthAtlantic Gough Island wereidentified as Herbertus runcinatus or Herbertus sendtneri .The Gough Island accession of H.sendtneri incorporated in the molecularstudyisnested in atropical Americanclade of H.sendtneri ,and isplaced sistertoaBolivianaccession. Bothspeciesare newtoGough Island. Jungermanniales/Herbertaceae/ Herbertus /DNA taxonomy/Gough Island /nrITS INTRODUCTION Molecularstudieshaveimproved the taxonomyof liverworts atdifferent ranks,e.g.,families(Hentschel etal.,2006),genera(Wilson etal.,2007)orspecies (Feldberg etal.,2004). Thesestudiespointed tovast morphologicalhomoplasy of morphologicalcharacterstatesof liverworts and demonstrated the need of an integrativetaxonomythatconsiders evidencefrom bothmorphologyand molecules(Heinrichs etal.,2009). Herbertus S.Graywasonceregarded asanancientgenus (Schuster, 2000);however,recentdivergencetime estimates(Heinrichs etal.,2007)pointto aCenozoicorigin of thisgenus,and speciation events in the Neogene (Feldberg etal.,2007). Herbertus isone of the most difficultgenerain termsof morphologicalcircumscription of species(Feldberg etal.,2004;Feldberg& Heinrichs,2006)and maythereforebenefitfrom the introduction of molecular methods.The problemsin identifying Herbertus speciesarethe consequenceof limited morphologicalvariation in combination withmorphologicalvariation within populationsthatpossess largelysimilarsequences(Feldberg etal.,2004). Asingle representativeof Herbertus hasbeen recognized forthe subantarcticand temperateislandsof the easternSouthernHemispherebyGrolle (2002). Helists Herbertus capensis (Steph.) Sim asthe only Herbertus speciesin the region,ataxon thatisnowtreated asasynonymof H.dicranus (Taylor) *Correspondenceand reprints:[email protected] 68J.Heinrichs,K.Feldberg,H.P.Kreier&J.Vá≈ a Trevis.(Hodgetts,2008). Alternatively,Wace(1961) published H.oldfieldianus (Steph.) Rodwayin hisinvestigation of the vegetation of Gough Island. The latter identification wasmaintained in Wace&Dickson (1965) and Hänel etal. (2005). Morphologicalinvestigation of several Herbertus specimensfrom Gough Island revealed the presenceoftwomorphologicallydivergentphenotype groups. However,assignmentof thesegroupstospeciesproved problematicusing solely morphologicalevidence. Toovercome the problem of morphologicalambiguity (Feldberg etal.,2004),tworecentcollectionsfrom Gough Island wereused fora molecularphylogeneticstudy.Thisstudyallowed foranassignmentof the Gough Island accessionstospeciesclades. MATERIAL AND METHODS DNA-extraction,sequencing and phylogeneticanalyses Twomorphologicallydiverging specimensof Herbertus from Gough Island (Gremmen 2000-170 and 2000-904,PRC, duplicatesin GOET)wereused forthe molecularinvestigation. DNA extraction and sequencing of the nuclear ribosomalinternaltranscribed spacerregion (nrITS)followed the protocols described in Feldberg&Heinrichs(2006). The newsequencesweremanually inserted in the nrITS sequencealignmentof Feldberg etal. (2007)using BioEdit version 5.0.9 (Hall,1999). Asubsetof the nrITS sequencesfrom Feldberg etal. (2007)wasused forthe phylogeneticanalyses(Table 1). Table 1. Geographicorigins,vouchernumbers,and GenBank/EMBL accession numbers of the taxainvestigated. Ifnototherwiseindicated,vouchers areatGOET.Newsequencesin bold. Acc.-Numb. Taxon Origin Voucher nrITS Mastigophoradiclados Reunion SchneiderH05-01 AM259580 Triandrophyllumsubtrifidum Bolivia Churchill etal. 22800 AJ972455 H.aduncus BritishColumbia Schofield 86245 AM259582 H.aduncus subsp. hutchinsiae BritishIsles Rycroft01015 AJ783340 H.delavayi Bhutan Long 28864 AJ972450 H.delavayi BritishColombia Schofield 83644 AM259583 H.delavayi BritishIsles Rycroft030725A AJ783341 H.dicranus Ethiopia(1) Miehe &Miehe 2524 AJ783343 H.dicranus Ethiopia(2) Miehe &Miehe 2435 AM259584 H.dicranus Indonesia Gradstein 10243 AJ783344 H.dicranus Japan Kurita51 (HIRO) AJ972452 H.dicranus Madagascar Pócsetal. 90116/O AJ783346 DNA-based identification of Herbertus specieson Gough Island 69 Table 1. Geographicorigins,vouchernumbers,and GenBank/EMBL accession numbers of the taxainvestigated. Ifnototherwiseindicated,vouchers areatGOET.Newsequencesin bold. Acc.-Numb. Taxon Origin Voucher nrITS H.dicranus Réunion SchneiderH05-02 AM259587 H.dicranus Rwanda SolgaRWA-1117 AM259589 H.dicranus Thailand Schäfer-Verwimp &Verwimp AJ972451 23850 H.juniperoideus subsp .pensilis Bolivia Heinrichsetal. 3998 AM113607 H.juniperoideus subsp .acanthelius Bolivia Heinrichsetal. 4000 AM113600 H.juniperoideus subsp .acanthelius Ecuador Gradstein &Mandl 10144 AJ783339 H.juniperoideus subsp .bivitattus FrenchGuiana HolzFG-00-201 AM113598 H.juniperoideus s . str. Ecuador Schäfer-Verwimp 23346 AM113605 H.juniperoideus Madagascar Pócs9444/D(EGR) AM259592 H.oldfieldianus NewZealand (1) Glenny9584 AM259594 H.oldfieldianus NewZealand (2) Engel 18886 AM259595 H.runcinatus Chile (1) Rycroft031123-1120 AM259596 H.runcinatus Chile (2) Rycroft031202-1125 AM259597 H.runcinatus Gough Island Gremmen 2000-170 FN582125 H.sendtneri Austria Heinrichs4377 AJ783347 H.sendtneri Azores RG2402 AJ783349 H.sendtneri Bhutan Miehe 00-380-01 AM259598 H.sendtneri Bolivia(1) Churchill etal. 22797 AM259599 H.sendtneri Bolivia(2) Heinrichsetal. 4002 AJ783351 H.sendtneri Bolivia(3) Heinrichsetal. 4004 AJ783352 H.sendtneri CostaRica Holz&Schäfer-Verwimp AJ783354 CR 00-77 H.sendtneri Ecuador Heinrichsetal. 4418 AM113611 H.sendtneri Gough Island Gremmen 2000-904 FN582126 H.sendtneri Malaysia Gradstein 10353 AJ972453 H.sendtneri Panama Dauphin etal. 1576 AJ783355 H.stramineus BritishIsles Rycroft00039 AJ783356 Phylogenetictreeswereinferred using maximumlikelihood (ML)criteria implemented in PAUP* 4.0b10(Swofford,2003). Modeltest 3.06 (Posadaand Crandall,1998) wasemployed todetermine the nucleotide substitution model 70 J.Heinrichs,K.Feldberg,H.P.Kreier&J.Vá≈ a withthe smallest numberof parameters thatbest fits the data.The hierarchical likelihood ratio tests selected the TrNmodel (Tamura&Nei,1993)withgamma shape parameterforamong-sitevariation. Thismodel wasimplemented in the ML analysis.The analysiswascarried out asaheuristicsearchwith3random- addition sequencereplicates.Clade support wasassessed bynon-parametric bootstrapping with200 replicates(Felsenstein,1985). Bootstrappercentage values (BSP)above70 wereregarded asgood support (Hillisand Bull,1983). Morphologicalinvestigation Twomorphologicalgroupswereidentified in the Herbertus collections from Gough Island using standardlightmicroscopy.Based on the identification of the twoDNA vouchers representing thesegroups,the collectionsfrom Gough Island wereassigned tospecies. RESULTS The ML analysisyielded asingle optimaltopologywithln= –4072,0731 (Fig. 1). Arobust clade withtwoaccessionsof H.runcinatus (Taylor)Trevis.and the Gough Island specimen Gremmen 2000-170 isplaced sistertothe rest of Herbertus.Aclade with Herbertus oldfieldianus from NewZealand and accessions of H.juniperoideus (Sw.) Grolle from tropicalAmerica and Madagascaris resolved in awell supported sisterrelationship withaclade made upby H. sendtneri (Nees)Lindb., H.aduncus (Dicks.) S.Gray, H.stramineus (Dumort.) Trevis., H.delavayi Steph. and H.dicranus.The well supported H.sendtneri clade comprisesaccessionsfrom Asia, Europe including Macaronesia, aswell asCentral and tropicalSouthAmerica.The Gough Island accession Gremmen 2000-904is nested in aNeotropicalsubclade of H.sendtneri and placed sistertoanaccession from the BolivianAndes.Based on the outcome of the molecularinvestigation, otherrepresentativesof the twomorphologicalgroupswereassigned toeither H. runcinatus or H.sendtneri : Herbertus specimensexamined (from Gough Island) Herbertus runcinatus (Taylor)Trevis.: S-slope of SouthPeak,loosegroupsin bryophyte matin heathland on mountain slope,530 m,15 Sep 1999 coll. N.J.M.Gremmen 99-168 (PRC);Sof Tafelkop,undergrowthinfernbush,slightlysheltered,very damp,300 m,7Jun 2000 coll. N.J.M.Gremmen 2000-0170 (GOET, PRC);top of SouthPeak,on shallow,wet organicsoil,slightlysheltered byboulders,800 m,16Sep 2000 coll. N.J.M.Gremmen 2000- 823 (PRC). Herbertus sendtneri (Nees)Lindb.: Main Glen,Michael’sRidge,1150ft.,21Jan1956,coll. N.M.Wace579 ,det.S.W.Arnell as H.oldfieldianus (cf. Wace,1961:353; BM, UPS); Tafelkop,growing in small numbers in large hepaticmats on very wetpeat,500 m,14 Sep 1999 coll. N.J.M.Gremmen 99-138 (PRC);Tafelkop,extensivemats in bryophytedominated mireonvery wetpeat,500 m,14 Sep 1999 coll. N.J.M.Gremmen 99-141 (PRC);N-side of Tafelkop,large mats on the lowerpart of rockoutcropssurrounded bymire,500 m,15 Sep 1999 coll. N.J.M.Gremmen 99-161 (F, PRC);Waterfall Camp,matof mixed hepaticson very wetpeaty soil in seepage area, 750m,16Sep 2000 coll. N.J.M.Gremmen 2000-873 (F); Waterfall Camp,densemats on very wetpeaty depositon verticalrockfaceinseepage area, DNA-based identification of Herbertus specieson Gough Island 71 Fig. 1. Phylogramrecovered in maximumlikelihood analysisof the nrITS dataset.Bootstrap percentage
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