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Strong Incongruence Between the ITS Phylogeny and Generic Delimitation in the Nemosenecio-Sinosenecio- Tephroseris Assemblage (Asteraceae: Senecioneae)

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Strong Incongruence Between the ITS Phylogeny and Generic Delimitation in the Nemosenecio-Sinosenecio- Tephroseris Assemblage (Asteraceae: Senecioneae) Botanical Studies (2009) 50: 435-442. PHYLOGENETICS Strong incongruence between the ITS phylogeny and generic delimitation in the Nemosenecio-Sinosenecio- Tephroseris assemblage (Asteraceae: Senecioneae) Liu-YangWANG1,2,5,PieterB.PELSER3,BertilNORDENSTAM4, and Jian-Quan LIU2,* 1Key Laboratory of Adaptation and Evolution of Plateau Biota, Northwest Plateau Institute of Biology, Chinese Academy of Sciences, Xining 810001, P.R. China 2Key Laboratory of Arid and Grassland Ecology, School of Life Science, Lanzhou University, Lanzhou 730000, P.R. China 3Oklahoma State University, Botany Department, 104 Life Sciences East Stillwater, Oklahoma 74078-3013, USA 4Swedish Museum of Natural History, P. O. Box 50007, SE-104 05 Stockholm, Sweden 5Graduate University of Chinese Academy of Sciences, Beijing 100049, P.R. China (Received January 9, 2009; Accepted March 9, 2009) ABSTRACT. ThethreegeneraSinosenecio,NemosenecioandTephroserisformacloselyknitgroupnested inthesubtribeTussilagininaeofthetribeSenecioneae(Asteraceae).Thegenericlimitsinthisassemblage remain unclear and need revision. In this study, we analysed sequences of the internal transcribed spacer (ITS) region of nuclear ribosomal DNA available from GenBank and sequenced 19 accessions of an additional 13 species encompassing all three genera. Phylogenetic analyses based on the ITS variation of 27 species in this assemblage and seven species from related genera of the Tussilagininae suggested that neither Sinosenecio norTephroserisismonophyletic.ThesampledspeciesofSinoseneciowerescatteredindifferentcladesor subcladesofthephylogenetictree.Fourspeciesofthisgenus,includingthegenerictypespecies(S. eriopodus,S. hederifolius,S. homogyniphyllusandS. subcoriaceus) are clustered in a tentative clade with genera such as Ligularia,Cremanthodium,Parasenecio, Farfugium and Tussilago.TheremainingtenSinoseneciospecies compriseahighlysupportedcladetogetherwith13TephroserisspeciesandfourNemoseneciospecies.Within thisclade,10TephroserisspeciestogetherwithtwoSinoseneciospecies(S. newcombeiandS. koreanus) compriseamonophyleticsubcladewhiletheremaining11speciesfromallofthreegeneraareclustered intoanothercladewithmoderatestatisticalsupport.Withinthelattersubclade,T. changii was revealed to becloselyrelatedtofourSinosenecio species,andthreeNemoseneciospeciescomprisingamonophyletic lineage.ThesetwolineagesformapolytomousradiationwiththeothertwoSinoseneciolineages.Thegeneric delimitations of the three genera clearly need some adjustments, which is also supported by previous studies of gross and floral morphology. Two Sinosenecio species(S. newcombeiandS. koreanus)shouldbetransferred to Tephroseris,andthegenusSinosenecio shouldbere-circumscribedtocontainthosespeciesclustered intheLigularia—Tussilagoclade.MostoftheotherdescribedspeciesunderSinosenecioandT. changii shouldeitherbetransferredto an enlarged Nemosenecio concept,oranewgenusneedstobeestablishedto encompass them. However, the morphological distinctions between these genera require further investigation. Keywords: Asteraceae;Internaltranscribedspacer(ITS)region;Nemosenecio;Molecularphylogeny; Sinosenecio;Tephroseris;Tussilagininae. INTRODUCTION etal.,2008).Sinoseneciocontainsabout38speciesof which37occurinChina,Korea,andIndo-Chinawith Nemosenecio (Kitam.) B. Nord., Sinosenecio B. a distinct center of diversity in the Sichuan province of Nord.andTephroseris(Reichenb.)Reichenb.areclosely China.Sinosenecionewcombei (Greene) J. P. Janovec relatedgeneraoftribeSenecioneaeoftheAsteraceae & T. M. Barkley, however, is endemic to the Queen (Liuetal.,2006;Pelseretal.,2007).Nemosenecio has CharlotteIslandsofCanada.Tephroseris containsaround six species, five of which are found in China and one in 50speciesmainlyfoundintemperateandarcticEurasia. Japan(JeffreyandChen,1984;Nordenstam,2007;Zhang SixspeciesarerecognizedinNWNorthAmerica,oneof whichisendemic(BarkleyandMurray,2006).Although *Correspondingauthors:E-mail:[email protected]; these three genera have traditionally been regarded [email protected];Fax:+86-931-8914288;Tel: asmembersofthesubtribeTussilagininaeDum.,the +86-931-8914305. Nemosenecio-Sinosenecio-Tephroserisassemblagehas 436 Botanical Studies, Vol. 50, 2009 also been recognized at the subtribal level (Jeffrey and to be revised to resolve monophyletic genera. For example, Chen,1984).TephroseridinaeC.JeffreyetY.L.Chenwas Goldenetal.(2001),promptedbytheextraordinary consideredasdistinctfromtheTussilagininaebyowning biogeographicalconsequenceofthetransferofSenecio narrow,cylindricalanther-collars,polarized,scattered,or newcombei Greene from the Queen Charlotte Islands radialendothecialcellwallthickenings,andconfluent, totheotherwiselargelyChineseSinosenecio; Janovec contiguousorseparatestigmaticareas(JeffreyandChen, andBarkley(1996)performedaphylogeneticstudyof 1984). These character states are, however, certainly not ITSsequencestoexaminethetruerelationshipsofS. uniquetoNemosenecio,Sinosenecio,andTephroseris newcombei.Theseauthorsfoundthatthetwospeciesof and have also been observed in the Tussilagininae sensu Sinosenecioincludedintheirstudies(S. koreanus(Kom.) JeffreyandChen(Liu,1999).Inaddition,Jeffreyand B.Nord.fromKoreaand S. newcombei)didnotforma Chen(1984)suggestedthegameticchromosomenumber monophyleticgroupandarenestedwithinTephroseris. of24asdiagnosticforTephroseridinae,althoughawide Thenon-monophylyofSinoseneciowasalsoconcluded range of other chromosome numbers have also been inanotherstudyusingITSsequencedata(Liuetal., observed (Jeffrey and Chen, 1984; Nordenstam, 2007; 2006).Inthatstudy,thesinglespeciesofTephroseris Pelseretal.,2007)andthemoretypicalTussilaginoid and Nemosenecio and two of the three Sinosenecio chromosomenumberofx=30hasbeenrecordedfor speciesincludedformedawellsupportedcladeinwhich twoSinosenecio species(Liu,2004).Ofthethreegenera, S. bodinieri(Van.)B.Nord.andS. globigerus(Chang) Sinosenecioespeciallyshowscharacterstatesofboththe B. Nord. comprised a strongly supported subclade. TephroseridinaeandTussilagininaesensuJeffreyand SinoseneciosubcoriaceusC.Jeffrey&Y.L.Chen,the Chen(1984)(Liu,2000).Becauseofthelackofdiagnostic third species included, however, appeared to be more charactersforTephroseridinaeanditsphylogenetic distantlyrelatedandwasplacedinalargepolytomywith positiondeeplynestedwithinTussilagininaesensuJeffrey several other Tussilagininae clades. The results of these andChen(1984),itiscurrentlynotrecognizedasa two studies were confirmed by Pelser et al. (2007), who subtribeand Nemosenecio,Sinosenecio,andTephroseris includedtheNemosenecio,Sinosenecio,andTephroseris areplacedinTussilagininaebymostauthors(e.g.,Bremer, ITSsequencesgeneratedbyGoldenetal.(2001)andLiu 1994;Liuetal.,2006;Pelseretal.,2007). etal.(2006)andafewotheraccessionsofSinosenecio Inadditiontodifficultiesconcerningthesubtribal andTephroserisintheirSenecioneaeITSphylogeny. delimitation, also the generic delimitation of the Thisstudyindicatedwithstrongbootstrapsupportand Nemosenecio-Sinosenecio-Tephroserisassemblagehas posteriorprobabilities thatSinosenecio newcombeiand been somewhat problematic. The three genera have S. koreanusaremorecloselyrelatedtoTephroseristhan mainlybeendefinedonthebasisofleafcharacters totheotherfourSinoseneciospeciesthatwereincluded. (JeffreyandChen,1984),althoughNordenstam(1978) Furthermore,Tephroseris changii proved to be distantly also indicated differences in habit and floral morphology. relatedtotheotherTephroserisspeciesandinsteadformed BothNemosenecio andTephroseris have pinnately-veined acladewithSinoseneciobodinieri,S. globigerus,and leaves, but those of Nemosenecioarepinnatisect,whereas S.septilobus(Chang)B.Nord.InthesamewayasLiu the leaves of Tephroserisaresubentireoronlyshallowly etal.(2006),Pelseretal.(2007)foundS. subcoriaceus lobed.Incontrasttothesetwogenera,mostspeciesof tobeonlydistantlyrelatedtotheothermembersofthe Sinosenecio have palmately-veined leaves with sinuate- Nemosenecio-Sinosenecio-Tephroserisassemblageand dentate to sinuate-denticulate margins, although the leaves nested within a large, but poorly resolved clade composed ofSinosenecio hainanensis(Chang&Tseng)C.Jeffrey& ofCremanthodium,Ligularia,Parasenecio,andother Y. L. Chen are pinnately-veined. Although Nemosenecio mostlyAsianTussilagininaegenera. is easily distinguished by its deeply incised leaves, the As a first step towards arriving at a monophyletic differencesbetweenSinosenecioandTephroserisarenot genericdelimitationoftheNemosenecio-Sinosenecio- alwaysclear.Tephroseris changiiB.Nord,forexample, Tephroserisassemblage,anITSphylogenyofthisgroup was regarded as a member ofTephroseris based on ispresentedthatincludesamuchlargersamplingofits its pinnately veined leaves, even though this species species(27)togetherwithaselectionofotherSenecioneae resemblessomeoftheSinoseneciospeciesinhabit,anther genera.Onthebasisofthisphylogeny,wediscussits shapeandphyllarynumber(JeffreyandChen,1984). incongruencewiththecurrentmorphology-basedgeneric Inrecentyears,phylogeneticstudiesofDNAsequences delimitation and explore alternative classifications to have been successfully applied to resolve the systematic obtainstrictlymonophyleticgenera. positions and generic delimitations of several Senecioneae genera(e.g.,KnoxandPalmer,1995;Swensonand MATERIALS AND METHODS Bremer, 1997, 1999; Panero et al., 1999; Bain and Golden,2000;WagstaffandBreitwieser,2004;Liuet Sequence data and sampled species al.,2006;Pelseretal.,2002,2003,2007;Wagstaffetal., TheITSsequencesincludedinthisstudywereeither 2006).Thesemolecularphylogeneticstudiessuggested newlyobtained(19accessionsrepresenting13species)
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