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Molecular Phylogeny of Myricaria (Tamaricaceae): Implications for Taxonomy and Conservation in China

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Molecular Phylogeny of Myricaria (Tamaricaceae): Implications for Taxonomy and Conservation in China Botanical Studies (2009) 50: 343-352. CONSERVATION Molecular phylogeny of Myricaria (Tamaricaceae): implications for taxonomy and conservation in China YongWANG1,YifeiLIU2,SongbaiLIU1,andHongwenHUANG2,* 1Wuhan Botanical Garden, The Chinese Academy of Sciences, Wuhan, Hubei, 430074, P.R. China 2South China Botanical Garden, The Chinese Academy of Sciences, Guangzhou, Guangdong 510650, P.R. China (ReceivedSeptember23,2008;AcceptedMarch4,2009) ABSTRACT. ThegenusMyricariabelongstothefamilyTamaricaceae,whichconsistsofthirteenspecies, tenofwhicharedistributedinChina. Theyareriparianorlake-sideshrubsandnaturallyoccurineastern Asia,extendingtocentralAsiaandEurope,withasuggestedcenteroforiginanddiversityintheHimalayan region. Mostofthespeciesarethreatenedbyincreasinghabitatfragmentationandanthropogenicdisturbances likedamandhighwayconstructionandover-grazing. Informationonmolecularphylogeneticrelationshipsis criticalforunderstandingthetaxonomyanddevelopingconservationstrategiesforMyricariaspeciesinChina. Inthepresentstudy,DNAsequencedatafromthenuclearribosomalinternaltranscribedspacerregionandthe plastidpsbA-trnHintergenicspacerwereusedtoinferthephylogenyofthegenus. Thirteenmorphological traitswerealsousedinconjunctionwiththemolecularphylogeneticrelationships. Thephylogeneticanalysis revealedabasalcladeofM. eleganstootherMyricariaspecies. Molecularevidenceresolvedonesuspicious specimenMyricaria sp.thatwascloselyrelatedtoM. wardii. Furthermore,theresultsrevealedthatthree widespreadspecies—M. paniculata,M. bracteataandM. squamosa—withlittlemorphologicaldifference havedistinctDNAsequences. Ontheotherhand,M. pulcherrimaandM. platyphyllawerefoundtobe groupedwiththeabovethreewidespreadspeciesdespitetheirmorphologicalcharactersbeingsimilarto thatofM. elegans. Also,M. laxiflorawasfoundinamorebasalphylogeneticpositionthanM. paniculata althoughthetwospeciesaremorphologicallysimilar. Ourphylogeneticanalysesprovidedmolecularevidence insupportingthehypothesisthatthecenteroforiginandevolutionforMyricariaspeciesistheHimalayan region. Thepresentstudyprovidesusefulbaselinedataforformulatingconservationprioritiesandfurther taxonomicdelineation. Keywords:Conservation;Morphology;Myricaria;Phylogeny;Tamariacaceae;Taxonomy. INTRODUCTION Myricariaspecies. Theyre-treatedaspecimenunder M. germanica(L.)Desv.collectedinChinaasanew ThegenusMyricariaDesv.belongstothefamily speciesM. paniculataP.Y.ZhangetY.J.Zhangand Tamaricaceae,whichcomprisesthreegenera,including upliftedM. germanicavar.laxifloraFranch.asaspecies TamarixL.,ReaumuriaL.,andMyricariaDesv.,andabout M. laxifloraP.Y.Zhanget.Y.J.Zhang. Inthisextensive 110specieswidelydistributedinEurope,Africa,andAsia revision,ZhangandZhang(1984)alsocombinedM. (ZhangandZhang,1984;Zhang,2005). Althoughthe alopecuroidesSchrankintoM. bracteataRoylebased equivocaltaxonomy,whichincludesfromthreetofive onthemorphologicalsimilarity. Otherwise,twospecies generawithinTamaricaceae,hasremainedcontroversial, M. hediniiO.PaulsenandM. laxaW.W.Sm.previously MyricariaismostcloselyrelatedtoTamarixsinceitwas describedbasedonspecimenscollectedfromTibetwere erectedasagenusfromTamarix and the first species of discardedbecauseofconfoundingsynonymywithM. M. squamosaDesv.wasdescribedin1825byDesvaux prostrataHook.f.etThomsexBenth.etHook.f.and basedonthedistinctmorphologicaldifferenceinstamens, M. squamosa. However,differencesinmorphological stigma, and leaflet (Zhang and Zhang, 1984). The most characteristicsbetweenspeciesorinfraspecifictaxaare recentrevisionbyZhangandZhang(1984)principally notalwaysclearcutandconsistent,andtherehasbeen consideredthetaxanativetoChina,butitincluded13 debateastowhatdegreeofmorphologicaldifferences observedjustifiesthedelineationoftaxa.Forexample, thetaxonomicstatusofthespeciesM. elegansRoylehas *Correspondingauthor:E-mail:[email protected]; remaineddebatableandcontroversial(Zhangetal.,2000; Tel:+86-20-37252778;Fax:+86-20-37252831. Huaetal.,2004;Zhangetal.,2005). 344 Botanical Studies, Vol. 50, 2009 Myricariahasawidegeographicdistributionextending GorgesRegion.ConstructionoftheThreeGorgesDam fromeasternAsiatoEurope,itsoriginalnativecenter raisedwaterlevelsandsubmergeditsmostdistribution anddiversitybeingtheQinhai-TibetPlateau(QTP), range(Wangetal.,2003;Liuetal.,2006).Several Chinaandadjacentregions(ZhangandZhang,1984). populationsofM. paniculataarealsoseverelydisturbed Itwasgenerallyacceptedthatthecommonancestorof bydamandhighwayconstruction(Wangetal.,2006). In MyricariaoriginatedinthePaleo-Mediterraneansea addition,desertificationandsoilerosionintherangeof region;likeothergenerainTamariacaceae,theHimalayas Myricariahasseverelydamagednaturalhabitatsofmany wassuggestedasthecenteroforigin.Thephylogenetic speciesandimpactedsurvivaloflocalpopulations.The divergenceandspeciationofMyricariaoccurredwiththe populationsizesofM. pulcherrimahavebeensharply riseoftheQTPduringtheTertiary(ZhangandZhang, decliningduetothedisappearanceoflakes/wetlandsinthe 1984;Zhangetal.,2001). Ofthe13speciesincluded TenggerDesert.Myricaria platyphyllaisalsoendangered inthegenusMyricaria,tenspeciesandonevarietyare becauseofovergrazingandhabitatdegradation(Wanget naturallydistributedinChinaandonlythreespecies—M. al.,2006). Awell-resolvedphylogenyisneededtodevelop germanica(Linn.)Desv.,M. dahurica(Willd.)Ehrenb. andprioritizetheconservationandmanagementstrategies andM. longifolia(Willd.)Ehrenb.—aredistributedin in consensus with defining evolutionary significant species EurasiaextendingfromcentralAsiatowesternEurope and/ortaxa(RusselloandAmato,2004;Dávalosetal., (ZhangandZhang,1984;Wangetal.,2006). Myricaria 2005;Ewenetal.,2006). speciesarealldeciduousshrubsofupstanding,recumbent The primary purposes of this study are : 1) to orcreepingforms,dependingonthehabitat,andare reconstructamolecularphylogenyofMyricariaincluding distributedinover14provinceswithinandaroundthe alltenspeciescollectedfrom17differentgeographic QTPandadjacentregionsinChina(ZhangandZhang, regionsinChinaandtwospeciesobtainedbyothersources 1984;Wangetal.,2006). Exceptforthethreewidespread usingsequencesofinternaltranscribedspacers(ITS)of speciesM. bracteata,M. paniculataandM. squamosa, nuclearribosomalDNAandthechloroplastpsbA-trnH mostMyricariaspeciesinChinahavenarrowdistributions intergenicspacer;2)toassessthetaxonomicdistinctness inmontaneareasclosetoriversorlakes. Theirvertical andrankofthespeciesincomparisontothereported distributionsinaltitudesarevariable,rangingfrom70to phylogeneticrelationshipspreviouslyestablishedbased 5200metersabovesealevel(a.s.l.)(Wangetal.,2006). onmorphologicalandmolecularanalysisofanumberof Anattempttodocumentmorphologicalvariationand ChineseendemicMyricariaspeciesavailable;and3)to taxonomicrevisionofMyricariaspecies(Zhangand addressconservationconcernsaboutMyricariaspecies Zhang,1984)basedononlymorphological datafailed endemictoChinainconsensuswith phylogeneticpattern toproducesatisfactoryresults,butcalledinterspecific ofthegenus. relationships among some species into question. Moreover,recentextensivefieldsurveysfoundthat MATERIALS AND METHODS taxonomic status and/or classification of species such asM. paniculata,M. squamosaandM. bracteataappearedtobe Plant material vagueandproblematic(Wangetal.,2006). Ontheother hand,previousstudiesinphylogenetic relationshipsand Several extensive field surveys were conducted during phytochemistrysampledonlyalimitednumbersofspecies thesummersfrom2002to2005inwestern,northwestern, inMyricariaandfocusedonsomespecificquestions southwestern,andcentralChina. Sourcesofplantmaterial (Chengetal.,2000;Jetter,2000;Gaskinetal.,2004;Hua usedinthisstudyarelistedinTable1,including11taxa etal.,2004). ThephylogenyandtaxonomyofMyricaria andonesuspiciousnewtaxonwhichwerecollectedat17 isstilldebatable. Moredetailedinformationisneeded differentgeographicallocationsfromnineprovincesor concerningtheoriginandphylogenyofMyricariaspecies. autonomousregions. Thedataoftwoothernonnativetaxa obtainedfromGenBank(ITSsequencenos:AF484746 Myricariaspeciesarepredominanttaxaintheriparian andAY572413)werealsoincluded. Twospecies,Tamarix plantcommunitiesinwesternChinaandplayimportant androssowiiLitv.andReaumuria songarica(Pall.)Maxim, rolesinecosystemservicestolocalharshenvironments fromrelatedgenerainTamaricaceaewereincludedin andsustainabledevelopmentinthewesternregion(Wang theanalysesasoutgroups. OneunclassifiedMyricaria et al., 2003). However, recent field surveys and studies specimen,tentativelynamedMyricariasp.,wasfound haveindicatedthatthemajorityofMyricariaspecies inthemountainareaofYunnanProvinceinsouthwest intheirnativehabitatsofwesternChinaarethreatened Chinaandwasobviouslymorphologicallydifferentfrom duetoincreasinganthropogenicdisturbanceandhabitat alldocumentedMyricariaspecies.Itwasalsoincludedin fragmentation(Wangetal.,2006). Theriparianhabitats thephylogeneticanalyses. Allvoucherspecimenswere areveryfragileandseverelydegradedbyanthropogenic depositedinWuhanBotanicalGardenHerbarium(HIB). activitiesinwesternChina.Anincreasingnumberofdams Leafmaterialswerecollectedandimmediatelydriedin havebeenconstructedforhydroelectricityandagricultural silicagelforDNAextraction,andsamplingsiteswere irrigationinthepasttwodecades. Awell-knownexample recordedusingaglobalpositioningsystem(GPS,Garmin, isM. laxiflora,aspeciesdistributedexclusivelyonthe eTrex). riverbanksoftheYangtzeRiverValleywithintheThree Wang et al. — Molecular phylogeny of —Molecularphylogeny etal. Wang Table 1. ListofMyricariaspeciesandoutgroupsusedinthisstudyandaccessionnumbersforsequencesdepositedinGenBank. Habitatsof Altitude Longitude GenBankaccessionnumber Taxon Naturaldistribution
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