基于2个核糖体DNA序列的国产白酒草属、小舌菊属战歧伞菊属(菊科紫菀族)体系学钻研.pdf

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  动物科学学报 2014ꎬ 32(3):216 ~227 Plant ScienceJournal DOI:10�3724/ SP�J�1142�2014�30216 基于2个核糖体 DNA序列的国产白酒草属、 小舌菊属和 歧伞菊属(菊科紫菀族)系统学研究 1 1∗ 1 2 1 1 钟 ꎬ 黎维平 ꎬ 杨秀林 ꎬ 唐 明 ꎬ 廖 威 ꎬ 陈三茂 (1� 湖南师范大学生命科学学院ꎬ 长沙 410081ꎻ 2� 中国科学院华南动物园ꎬ 中国科学院动物资本取可持续操纵沉点尝试室ꎬ 广州 510650) 摘 要:国产白酒草亚族(菊科紫菀族)由白酒草属(Conyza)、 小舌菊属(Microglossa)和歧伞菊属(Thespis) 3个小属构成ꎬ 且国产白酒草亚族各属间及其取非洲白酒草属动物之间的系统发育关系尚无报道ꎬ 故本研究 操纵核糖体 DNA ITS和ETS序列并采用最大简约法和贝叶斯阐发法ꎬ 沉建了国产白酒草亚族的系统发育 树ꎮ 成果表白ꎬ 国产4种白酒草属动物、 歧伞菊和非洲白酒草属动物构成一支ꎬ 而劲曲白酒草的两变种和小舌 菊嵌入田基黄亚族分支ꎻ 小舌菊取Psiadiapascalii近缘ꎮ 基于这些成果ꎬ 我们认为: (1)劲曲白酒草和 Conyza incisa应处置为田基黄亚族的一个的属ꎻ (2)国产4种白酒草属动物和歧伞菊以及大大都非洲白酒草属动物 属于Eschenbachia属ꎬ 并且 Eschenbachia属代表一个新的亚族ꎬ 歧伞菊可处置为Eschenbachia属的一个组ꎮ Eschenbachia属可能从非洲经数次长距离达到我国南部ꎻ (3)Welwitschiella 和小舌菊属应连结属的地位ꎬ Psiadiapascalii、 Conyza scabrida和C� pyrrhopappa可并入小舌菊属ꎮ 环节词:国产白酒草族ꎻ 白酒草属ꎻ Eschenbachiaꎻ 小舌菊属ꎻ 系统发育ꎻ 歧伞菊属 中图分类号:Q949�783�5 文献标识码:A 文章编号:2095 ̄0837(2014)03 ̄0216 ̄12 Molecular Phylogeny of Chinese Conyzaꎬ Microglossa and Thespis (Asteraceae: Astereae) Based on Two Nuclear Ribosomal DNA Regions 1 1∗ 1 2 1 1 ZHONG Cai ̄Xia ꎬ LIWei ̄Ping ꎬ YANGXiu ̄Lin ꎬ TANG Ming ꎬ LIAOWei ꎬ CHENSan ̄Mao (1� College of Life Sciencesꎬ Hunan Normal Universityꎬ Changsha410081ꎬ Chinaꎻ 2� Key Laboratory of Plant Resources Conservation and Sustainable Utilizationꎬ South China Botanical Gardenꎬ ChineseAcademy ofSciencesꎬ Guangzhou510650ꎬ China) Abstract: Chinese subtribe Conyzinae (Asteraceae: Astereae) consists of three small generaꎬ Conyzaꎬ Microglossa and Thespis� To dateꎬ howeverꎬ the molecular phylogenetic relationships among African Conyzaꎬ and Chinese Conyzaꎬ Thespis and Microglossa have not beeninvestigated� The internalandexternaltranscribedspacersof nuclearribosomalDNA were used to reconstruct the phylogeny of Chinese Conyzinae through maximum parsimony and Bayesian analyses� Results showed that four Chinese Conyza species and Thespis divaricata belonged to a clade with most sampled African Conyzaꎬ whereas two Conyza stricta varieties and Microglossa pyrifolia were embedded deeply within the subtribe Grangeinae clade� Microglossa pyrifolia was closely related to Psiadia pascalii� Based on phylogenyꎬ we considered that (1) C� stricta and C� incisa should be treated as a separate genus that belongs to subtribe Grangeianeꎻ (2) four Chinese Conyzaꎬ Thespis and most 收稿日期:2014 ̄01 ̄21ꎬ 修回日期:2014 ̄02 ̄20ꎮ 基金项目:国度天然科学基金赞帮项目(31370265)ꎻ 湖南省生态学沉点学科扶植项目赞帮ꎻ 湖南省教育厅科学研究基金赞帮 (08A046)ꎮ - 钟(1988 )ꎬ 女ꎬ 硕士研究生ꎬ 次要研究标的目的为种子动物分类学(E ̄mail: 860905430@qq�com)ꎮ ∗通信做者(Authorforcorrespondence. E ̄mail: lwprenyi@aliyun�com)ꎮ 钟等: 基于2个核糖体 DNA序列的国产白酒草属、 小舌菊属和 第3期 歧伞菊属(菊科紫菀族)系统学研究(英文) 217 sampledAfrican Conyzabelongtothegenus Eschenbachia� Eschenbachiashould belongtoa new separate subtribeꎬ while Thespis should be treated as a section of Eschenbachia� Eschenbachia may have arrived at southern China by several long ̄distance dispersals from Africaꎻ (3) it is reasonable to maintain the generic status of Welwitschiella and Microglossa and treat Psiadia pascaliiꎬ C� scabrida and C� pyrrhopappa as members of Microglossa. Keywords:ChineseConyzinaeꎻ Conyzaꎻ Eschenbachiaꎻ Microglossaꎻ Molecularphylogenyꎻ Thespis [8] AlthoughtribeAstereae isthesecond largest World Conyza . inAsteraceae (Compositae)ꎬ phylogenetic rela ̄ Howeverꎬ Old World Conyza is not mono ̄ tionships within Astereae are still far from re ̄ phyletic� As early as 1990ꎬ based on certain [1 ̄5] [8] solved � ChineseAstereae subtribe Conyzinae morphological featuresꎬ Nesom concluded that consists of three generaꎬ Conyza Less�ꎬ Micro ̄ Old World Conyza was strongly heterogeneous [6] glossa DC� and Thespis DC� ꎬ which have not and its taxonomy was farfrom resolved� In addi ̄ yet been phylogenetically evaluated based on tionꎬ some molecular phylogenetic analyses [8] molecular data. have supported Nesom ꎬ showing that Old Conyzawas established in 1832 by Lessing [3] World Conyza was polyphyletic or at least and typified by C� chilensis Sprengelꎬ a New biphyletic[1ꎬ2ꎬ11ꎬ13]. For exampleꎬ 14 sampled = World species ( Conyza primulifolia (Lam�) Conyza species of Madagascar and Indian [6ꎬ7] Cuatrec� and Lourteig) � Morphological studi ̄ Ocean islands formed several smallerꎬ mostly es[7 ̄10] and molecular data[1ꎬ2ꎬ4ꎬ11 ̄13] revealed that weakly supported clades with two to five species New World Conyzawas not closely relatedto Old in phylogenetic treesꎬ indicating that African World Conyza� Furthermoreꎬ phylogeneticanaly ̄ [3] [3] Conyza was polyphyletic � Strijk transferred ses[1ꎬ2ꎬ12] suggested that New World Conyzawas threeAfrican Conyza species (C� scabrida DC�ꎬ at least biphyletic and northern and southern C� pyrrhopappa Sch� Bip� and C� ageratoides American Conyza were all derived from Erigeron DC� ) to a newly circumscribed Psiadia s�s�ꎬ [14] L� Accordinglyꎬ Chen et al� merged a few whereas other African Conyza species were New World Conyza species introduced to China paraphyleticwithrespect to FrappieriaCordemoyꎬ [7] [3] into Erigeron� Converselyꎬ Nesom suggested which was resurrected by Strijk to accommo ̄ that OldWorld species identified as Conyzawere date the smallerof the two clades of Psiadia. related to genera outside Conyzinaeꎬ and Ne ̄ Up to nowꎬ howeverꎬ molecular phyloge ̄ [10] som and Robinson stated that “species of netic analysesꎬ including Old World Conyzaꎬ Conyza nativetoAfrica apparently are more simi ̄ have mainly sampled African species[1 ̄5ꎬ11ꎬ13]. lar and perhaps moreclosely relatedto generaof Three Conyza speciesꎬ C� blinii H� Lév� (Es ̄ subtribeGrangeinae”� Similarlyꎬ Brouillet etal�[1] chenbachia blinii (H� Lév� ) Brouillet )ꎬ C� suggested that the subtribe Grangeinae should perennis Hand� ̄Mazz� (Eschenbachia perennis be expanded to include African Conyza� Re ̄ (Hand� ̄Mazz� ) Brouillet) and C� muliensis Y� centlyꎬ as a small part of Old World Conyzaꎬ L� Chen (Eschenbachia muliensis (Y� L� Chen) Conyza species (excluding C� stricta Willd� ) Brouillet)ꎬ are endemic to Chinaꎬ though their native to China were assigned to Eschenbachia molecular data have not yet been reported. [14] Moench ꎬ the oldest name representing Old Thespis DC� isasmallgenuswiththreespe ̄ 218 植 物 科 学 学 报 第32卷 cies and distributed in south ̄easternAsia� Thes ̄ Microglossa pyrifoliaand Thespis divaricata (Ap ̄ pis divaricata DC� is the only species of the ge ̄ pendix) were sampled in China for sequence nus from China� The systematic position of this variations in nrDNA ITS and ETS� The ITS and genus has beenproblematicfora longtime� Ling ETS sequencesof C� sumatrensis (Retz�)Walk ̄ [6] er and C� japonica (Thunb� ) Less� were pro ̄ et al� placed the genus within subtribe Conyzi ̄ [15] [4] naeꎬ whereas Zhang and Bremer treated it as duced from previous studies ꎬ andthoseof the an isolated genus of Astereae and Bremer[16] other seven species were newly generated (Ap ̄ thought it to be a member of subtribe Asterinae� pendix). [9] [2] Nesom included it tentatively in subtribe Lag ̄ Brouillet et al� divided Astereae into six enophorinae Nesomꎬ whichwas followed by Ne ̄ phylogenetic lineagesꎬ that isꎬ the basal linea ̄ [10] gesꎬ palaeo ̄South American cladeꎬ New Zea ̄ som and Robinson � Although its genetic sta ̄ [14] land cladeꎬ Australasian lineagesꎬ South Ame ̄ tuswas supported by Chen etal� ꎬ its system ̄ atic position remains uncertain and no molecular rican lineages and Northern American lineage� [4] phylogenetic analyses have involved Thespis so Accordingto Lietal� ꎬ there isaseventhcladeꎬ far. Bellidinaeꎬ and an eighth cladeꎬ Grangeinae� With approximately 10 speciesꎬ Microglossa To determine systematic positions of the three [14] Chinese generaꎬ all eight phylogenetic lineages DC� is distributed in Africa and tropical Asia� Microglossa pyrifolia (Lam� ) O� Kuntze occurs were represented by more or less species in our widely from Africa to tropical Asiaꎬ and in China ITS matrix andsevenphylogenetic lineages inthe it is a traditional medicine and the only represen ̄ combined matrix� Therewere moreaccessions in [6ꎬ14] [6] the ITS matrix than in the combined matrix be ̄ tative of the genus � Ling et al� assigned the genus within Conyzinaeꎬ while Zhang and causetherewas more ITS data available in Gen ̄ [15] [16] [1ꎬ4ꎬ5ꎬ11ꎬ12ꎬ17 ̄26] [3] [5] Bremer and Bremer placed it in the Eriger ̄ Bank � Strijk andStrijk etal� sug ̄ [9] gested that African Conyza species were nested on ̄Conyza group� Howeverꎬ Nesom placed it in the Psiadia group of subtribe Baccharidinaeꎬ withtwo large cladesꎬ so in our ITS matrix some [10] African Conyza and their relatives represented and Nesom and Robinson treated it as an un ̄ placed genera in tribe Astereae� Thus farꎬ how ̄ twoclades (one clade: C� ageratoidesꎬ C� scabri ̄ everꎬ its phylogeny has not been evaluated by daꎬ C� pyrrhopappaꎬ C� strictaꎬ Psiadia pas ̄ molecular data. calii Labat and Beentjeꎬ P� punctulata (DC� ) Based on nuclear ribosomal DNA (nrDNA) Vatkeꎻ the other clade: eight Conyza species ITS and ETS (internal and external transcribed and PsiadiaargenteaCordem�) inorderto reval ̄ spacers of 35S ribosomal DNA)ꎬ the present uate the relationships among theseAfrican plants study aimed to reconstruct the phylogeny of Chi ̄ and the Chinese taxa� Inthe ITS matrixꎬ two se ̄ nese Conyzaꎬ Microglossa and Thespis and to quences (HE978363 and GU045830) published [5] discuss their classification. by Strijk et al� were recorded in GenBank and were obtained from the same speciesꎬ Conyza 1 Materials and methods ulmifolia� The two are quite differentꎬ however� 1�1 Taxonsampling and source of sequences Comparisonof our ITStreewiththat of Strijketal�[3] Four Conyza species native to Chinaꎬ three showed that HE978363 was the ITS sequence of [5] American Conyza species introduced to Chinaꎬ C� incisa instead of C� ulmifolia� As a resultꎬ 钟等: 基于2个核糖体 DNA序列的国产白酒草属、 小舌菊属和 第3期 歧伞菊属(菊科紫菀族)系统学研究(英文) 219 we treated HE978363 as the ITS of C� incisa nealing (56℃ꎬ 40 s) and extension (72℃ꎬ [5] 1min)ꎬ and a finalextensionof 10 min at 72℃. accordingto Figures2and3 ofStrijk etal� The ITS sequence GU045827 published by Strijk The PCR fragments were purified with a [5] UNIQ ̄10 Spin Column PCR Product Purification et al� was obtained from C� stricta sampled from easternAfrica� We regarded it as C� stricta Kit (Sangon Biotech Co�ꎬ Ltdꎬ Shanghaiꎬ Chi ̄ var� stricta distributed in both Asia and Africaꎬ na) following the manufacturer�s instructions� Di ̄ and was thus distinguished from the other sam ̄ rect sequencingwas performed inbothdirections pled varietyꎬ C� stricta var� pinnatifida Kitam� by Sangon Biotech Co�ꎬ Ltd. Conyza stricta var� pinnatifida is restricted to 1�3 Sequence alignment and phylogenetic ana ̄ southern Asia and was sampled from Yunnanꎬ lyses [4] China (Appendix). Because some taxa of the Accordingtoourpreviousstudy ꎬ bounda ̄ ITS matrix were unavailable for ETSꎬ the com ̄ ries ofthe ITSand ETS regionsweredetermined� bined ITS/ ETS matrix included only 34 ingroup All DNA sequences were aligned initially using [32] taxa (Appendix). In the two matricesꎬ Chrysan ̄ Clustal X 1�83 and then adjusted manually in [33] themum coronarium L� and Dendranthema indi ̄ BioEdit � The ITS region was analysed sepa ̄ cum (L� ) Des Moul� of tribe Anthemideae and rately and in a combined data set with the ETS Calendula officinalis L� of tribe Calenduleae (Ap ̄ region� The incongruence length difference test[34] pendix) were selected as outgroups for the roo ̄ was carried out to test the homogeneity between ting of the phylogenetictrees because inmolecu ̄ ∗ data sets using PAUP version4�0b10with 1000 lar phylogenetic analyses[27ꎬ28] Anthemideae and replicates� Maximum parsimony (MP) and Baye ̄ Astereae are sistersꎬ and Calenduleae is a sister sian inference (BI) were performed for the data to tribes Gnaphalieaeꎬ Anthemideae and Astere ̄ ∗ [35] sets using PAUP version 4�0b10 and Mr ̄ [36] ae. Bayes version3�1�2 ꎬ respectively� In the MP 1�2 DNA extractionꎬ polymerase chain reaction analysisꎬ characters were unorderedandequally (PCR) and sequencing weightedꎬ gaps were scored as missing dataꎬ Total genomic DNA was isolated from fresh and a heuristic search was implemented with leaf materials or silica gel ̄dried leaves� Amplifi ̄ 1000 random additional sequence replicates and cation and sequencing were performed using sub ̄tree pruning ̄regrafting branch swapping� [29] primers ITS1and ITS4 forthe ITS regionꎬ and Support for clades was estimated by bootstrap [30] [31] using primers Ast ̄8 and 18S ̄ETS for the with 1000 replications� The MaxTrees setting in ∗ ETS region. PAUP was set to 5000 for the searches and - The PCRmixturecontained 1μL (50 100ng) bootstraptests� For BI analysis of the ITS region × of sample DNAꎬ 2 2 μL of primer (10 pmol)ꎬ and combined data setꎬ the best ̄fitting modelof 2+ × 5 μLof 10 PCRbufferꎬ3μLof Mg (25mol/ L)ꎬ each sequence partition (ITS1ꎬ ITS2ꎬ 5�8Sꎬ [37] 0�8μLof deoxyribonucleotide triphosphate (each ETS)was determined using MrModeltest2�2 � 25 mol/ L)ꎬ 0�5 μL of Taq DNA polymerase + The SYM G modelwas chosenforthe5�8S re ̄ + + (5 U/ mL) and sterile water for a final volume of gionꎬ and the GTR I G model for the ITS1ꎬ 50 μL� The PCRprocedureswereasfollows: ini ̄ ITS2 and ETS regions� The Markov chain Monte tial denaturation for 4 min at 95℃ followed by Carlo algorithmwas runfor 1000000 generationsꎬ 30 cycles of denaturation (94℃ꎬ 1 min)ꎬ an ̄ resulting in an overall sampling of 10000 trees� 220 植 物 科 学 学 报 第32卷 The first 3000 trees were discarded as a conser ̄ subclade but mixedwithothergenera (Fig�1and vation burn ̄inꎬ and the remaining trees were Fig�2). Microglossa pyrifolia was nested within used to construct the 50% majority rule consen ̄ clade2 and was not closely related to most Chi ̄ sus tree. nese Conyza (Fig�1and Fig�2)ꎬ whereas Thes ̄ pis divaricata belonged to clade 7with most Afri ̄ 2 Results can and Chinese Conyza sampled. 2�1 Characterizationof nucleotide data 3 Discussion The ITSdataset consistedof680alignednu ̄ cleotidesꎬ of which383 (56�32%) were variable 3�1 Phylogenetics of Chinese Conyza and the and 278 (40�88%) were parsimony ̄informative� taxonomic implications = The ILD test (p 0�123) failed to reject the Three American Conyza speciesꎬ C� bona ̄ combination of the ITS and ETS markers� There ̄ riensisꎬ C� canadensisand C� sumatrensisꎬ are foreꎬ acombinedanalysisofthetwo regionswas widely distributed throughout China� They were ∗ performed using PAUP and MrBayes� Thecom ̄ found to be nested in the North American clade = bined ITS and ETS data set consisted of 1335 with highsupportedvalues (clade 1: PP 1�00 = = = alignednucleotidesꎬ ofwhich715 (53�56%)were and BS 99 inFig�1ꎻ PP 1�00and BS 100 variable and 495 (37�08 %) were parsimony ̄ in Fig�2) of Astereae and closely related to Eri ̄ informative� Near the end of the ETS region Eri ̄ geron� Inparticularꎬ Erigeronbreviscapusshared geron breviscapus (Vant�) Hand� ̄Mazz�ꎬ Cony ̄ an 84 bp segment of ETSꎬ a synapomorphic za canadensis (L� ) Cronq�ꎬ C� bonariensis characterꎬ with C� canadensisꎬ C� bonariensis (L� ) Cronq� and C� sumatrensis had an insert and C� sumatrensis� These resultssupport previ ̄ as long as 84 bpthat was their synapomorphy. ous reports[1 ̄4ꎬ7ꎬ11 ̄13] and the taxonomic treatment [14] 2�2 Phylogenetic analyses of Chen et al� that these three Conyza were Phylogenetic analyses using ITS and the placed inthe genus Erigeron� Although previous combined data sets yielded generally consistent studiesalsoshowedthat NewWorld Conyzawere phylogenetic trees (Bayesian treesꎻ see Fig�1ꎬ not closely related to Old World Conyzaꎬ only Fig�2)ꎬ though combined analyses resulted in one Asian speciesꎬ C� japonica (Eschenbachia [4] more supported clades and more resolved rela ̄ japanica)ꎬ was included � Our study showed tionshipsthanthe independent analysis basedon that inthe trees (Fig�1 and Fig�2) the three in ̄ ITS data� In boththe ITStreeandcombinedtreeꎬ vasive alien Conyza species were far from the the genus Conyzawas not monophyletic and oc ̄ Conyza native to China. curred within three big clades with higher Baye ̄ Our phylogenetic trees (Fig�1 and Fig�2) sian posterior probability (PP) and bootstrap demonstrated that Chinese Conyzawere biphyle ̄ = = support (BS) (clade 1: PP 1�00 and BS tic because Conyza stricta var� stricta and var� = = 99 in Fig�1ꎬ and PP 1�00 and BS 100 in pinnatifida belonged to clade 2 and were not Fig�2ꎻ clade 2: PP = 1�00 and BS = 68 in close to clade 7 where the other three Chinese = = Figs�1and 2ꎻ clade7: PP 0�99and BS 90 Conyza occurred (Fig�1and Fig�2). Conyza in ̄ in Fig�1ꎬ and PP = 1�00 and BS = 100 in cisa andthetwo C� strictavarietiesoccurred ina = = Fig�2). Furthermoreꎬ ineachof the three clades well ̄supported sub ̄clade (PP 1�00ꎬ BS 95ꎻ the Conyza were not grouped in an independent Fig� 1). Unexpectedlyꎬ var� pinnatifida initially 钟等: 基于2个核糖体 DNA序列的国产白酒草属、 小舌菊属和 第3期 歧伞菊属(菊科紫菀族)系统学研究(英文) 221 1.00/96 Aster verticillatus AIS = Astereae incertae sedis 1.00/100 Aster tataricus AL = Australasian lineages AL Aster ageratoides var. lasiocladus BE = Bellidinae Sheareria nana BL = early-branching lineages 0.99/- Baccharis neglecta SA BS = Eschenbachia Aster lavanduliifolius GR = Grangeinae 1.00/- Calotis hispidula NA = North American lineage NZ = New Zealand clade Lagenophora pumila AL OG = outgroup 1.00/100 Myriactis nepalensis PSA = palaeo South American clade Myriactis wightii SA = South American lineages Conyza chilensis 1.00/78 Conyza sumatrensis 1.00/100 Conyza bonariensis 1.00/54 1.00/99 0.96/- Conyza canadensis NA 1 Erigeron breviscapus 1.00/87 Symphyotrichum subulatum Eurybia sibirica 0.98/64 Bellis perennis BE 1.00/88 Galatella dahurica Tripolium vulgare Dichrocephala auriculata 1.00/- Nidorella polycephala 3 Grangea maderaspatana 1.00/95 Conyza stricta var. stricta 1.00/68 Conyza stricta var. pinnatifida 2 Conyza incisa GR 0.99/53 Welwitschiella nereifolia 1.00/100 Conyza ageratoides 0.94/- 5 Psiadia punctulata 4 1.00/100 Microglossa pyrifolia 1.00/- Psiadia pascalii 6 Conyza scabrida Conyza pyrrhopappa 0.97/77 Psiadia argentea 0.99/96 Conyza attenuata 1.00/90 Conyza pinnata 0.98/51 Conyza lim

  基于2个核糖体DNA序列的国产白酒草属、小舌菊属和歧伞菊属(菊科紫菀族)系统学研究.pdf

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