中国天牛亚科昆虫系统发育与生物地理学分析(鞘翅目,天牛科)
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摘要
天牛亚科Cerambycinae是天牛科Cerambycidae中较大的类群,包含了很多农林业害虫、检疫性昆虫和访花类昆虫,具有重要的经济意义。此外,该亚科包含种类数量大,生活习性复杂,众多学者对亚科内族属划分意见不一,极大地影响了该亚科分类研究的发展。本文通过检视大量中国天牛亚科昆虫标本、查阅并收集相关文献,筛选了98个特征对中国天牛亚科进行系统发育研究,并对其中10个族进行了族内属间的系统发育探讨。检视标本期间发现天牛亚科3新种,1中国新纪录属,18中国新纪录种,订正2亚种名。对中国虎天牛族Clytini进行了DNA条形码的初步研究,并结合NCBi数据库中数据建树分析并探讨了该族的系统发育关系。整理了中国天牛亚科(仅本研究所涉及种类)昆虫的区系分布,分析了中国天牛亚科昆虫的分布概况,初步分析了中国天牛亚科昆虫的起源和进化。获得的主要结果如下:1.中国天牛亚科系统发育研究1.1检视标本中发现的新单元:
     淡纹露胸天牛Jrtimpaza brevilineata Tian et Chen,2012
     长距刺虎天牛Demonaxproculscuti Li, Tian et Chen,2013
     蒋氏绿虎天牛Chlorophoms jiangi Li, Tian et Chen,2013
     横纹绿虎天牛Chlorophorus copiosus Holzschuh,1991(中国新纪录)
     回纹艳虎天牛Rhaphuma lanzhui Holzschuh,1991(中国新纪录)
     箭纹艳虎天牛Rhaphuma illicata Holzschuh,1991(中国新纪录)
     断纹刺虎天牛Demonax traudae Holzsclmh,1983(中国新纪录)
     蔷藻刺虎天牛Demonax rosae Holzschuh,1983(中国新纪录)
     格氏刺虎天牛Demonax gertrudae Holzschuh,1983(中国新纪录)
     )〈点刺虎天牛Demonax contrarius Holzschuh,1991(中国新纪录)
     独艳虎天牛Rhaphuma unigena Holzschuh,1993(中国新纪录)
     宽带绿虎天牛Chlorophorusproannulatus Gressitt&Rondon,1970(中国新纪录)
     四纹痩棍腿天牛Stenodryas nigromaculatus (Gardner,1942)(中国新纪录)
     白腹痩棍腿天牛Stenodryas ventralis (Gahan,1906)(中国新纪录)
     灰色跑■虎天牛Perissm cinericius Holzschuh,2009(中国新纪录)
     异拟眉天牛Epicyftus insolitus Holzschuh,1991(中国新纪录)
     长纹绿虎天牛chlorophorusruifmembrisgressitt&rondon,1970(中国新纪录)老挺溢銷天牛kuraruaplautagressitt&rondon,1970(中国新纪录)锤腿天牛属nyphasiapascoe,1867(中国新纪录属)帕氏锤腿天牛况pascoeilacordaire(中国新纪录)六纹艳虎天牛rhaphumanishidaihayashi&makihara,1981(中国新纪录)工斑绿虎天牛chlorophorusconiperdaholzschuh,1992(中国新纪录)取消亚种单元:红胸附虎天牛perissusmutabilisvitabilispic,1923黑胸跑■虎天牛perissusmutabilisobscuricolorpic,19371.2系统发育研究结果检视了天牛亚科19族87属301种昆虫,筛选了98个外部形态特征、解剖特征、生物学习性性状并编码,分别利用paup(nj法)",mrbayes(贝叶斯法)建立系统发育树,所得结果如下:①天牛亚科族间的系统发育关系:天牛亚科为单系群;折天牛族pyrestini、扁胸天牛族callidiini、绿天牛族callichromatini、侧沟天牛族obirini、紫天牛族trachydeirni及虎天牛族clytini均为单系群;天牛族cerambycini昆虫属较原始类群,位于分支图基部;绿天牛族callichromatini与紫天牛族trachydeirni亲缘关系较近;折天牛族pyrestini与天牛族cerambycini亲缘关系较近,沟角天牛族hesperophanini与丽天牛族rosaliini亲缘关系较近,虎天牛族clytini和眉天牛族tillomorphini及锥背天牛族thraniini亲缘关系较近,纤天牛族cleomenini和狭鞘天牛族stenopterini、侧沟天牛族obirini及短鞘天牛族molorcliini亲缘关系较近。②各族内属间系统发育关系:(1)折天牛族pyrestini红天牛属erythms为单系群,珠角天牛属pachylocerus与长红天牛属erythresthes亲缘关系较近,突肩折天牛p.pascoei是折天牛属pyrestes最原始的种类。研究结果不支持折天牛属独立成一族。(2)绿天牛族callichromatini是单系群,多带天牛属polyzonus与灿天牛属anubis亲缘关系较近,黑线天牛属embrikstrandia、拟柄天牛属cataphrodisium与厚天牛属亲缘关系较近。柄天牛属却如为并系群,紫绿柄天牛a.cantori与其他柄天牛属种类亲缘关系较远。(3)紫天牛族trachydeirni是单系群,珊糊天牛属dicelosternus与阔嘴天牛属亲缘关系最近,肖亚天牛属力与其他属亲缘关系较远。阔嘴天牛属、肖亚天牛属形成单系群。ii
     (4)蜡天牛族callidiopini为并系群,蜡天牛属ceresiwm、拟蜡天牛属stenygirnum和瘦棍腿天牛属stenodryas形成单系群;异柱天牛属parasalpinia具有多个特征与另外三属存在较大差异,建议将其移出该族。(5)虎天牛族clytini中脊虎天牛属xyknrechus与丽虎天牛属plagionotus亲缘关系较近,球虎天牛属calloides与跗虎天牛属perissus亲缘关系较近。(6)纤天牛族cleomenini中红胸天牛属det-e、露胸天牛属artimpaza及纤天牛属cleomenes均为单系群,原纤天牛属procleomenes与露胸天牛属artimpaza亲缘关系较近,尼天牛属nida与溢鞘天牛属kurarua亲缘关系较近。(7)扁胸天牛族callidiini除扁胸天牛属callidium以外构成一支,族内小扁天牛属callidiellum与扁鞘天牛属ropalopus亲缘关系较近。杉天牛属semanotus及沟跗天牛属turanium均形成单系群。(8)沟角天牛族hesperophanini中各属均为单系群。锐天牛属zoodes为该族最原始类群,其次为莺天牛属trichoferus’凿点天牛属stromatium和额天牛属gnatholea。(9)天牛族cerambycini为单系群。棱天牛属xoanodem、缘天牛属margites、脊胸天牛属rhytidodera及皱胸天牛属neoplocaederus均为单系群;闪光天牛属aeolesthes较为混乱,其中红绒闪光天牛a.ningshanensis与肿角天牛属neocerambyx樱红肿角天牛n.oenochrous亲缘关系最近;褐天牛属nadezhdiella洁褐天牛n.cantori与肿角天牛属neocerambyx亲缘关系较近;刺角天牛属trirachys与闪光天牛属aeolesthes中除红域闪光天牛aningshanensis外的其他种类亲缘关系较近。(10)双条天牛族xystroceirni中3属均为单系群,茶色天牛属oplatocera较为原始,肋翅天牛属hyphm与双条天牛属xystrocem亲缘关系较近;茶色天牛属oplatocera具有多个特征与另外两属存在较大差异,建议将其移出该族。2dna条形码应用于虎天牛族系统发育研究dna条形码研究涉及天牛亚科cerambycinae虎天牛族clytini9属25种60余条序列,种内遗传距离差异达0-1.9%,使用barcode进行虎天牛族种类鉴定时,遗传距离差异阈值选择低于2%为同种较为合适。barcode在种间的遗传距离差异阈值为2.9%-16.5%,大于种内遗传距离差异的最大值,证明barcode可应用于该族昆虫的种类鉴定。根据本研究得到的dna条形码序列结合ncbi已有序列联合建树分析了虎天牛族亲缘关系,结果表明:矮虎天牛属amamiclytus与剌虎天牛属demonax亲缘关系较近,归入刺虎天牛属较合适;绿虎天牛属chbrophorus是该族中较原始类群,艳虎天牛属亲缘关系较为杂乱,附虎天牛属perisms■为单系群。通过与外部形态特征建树结果对比得到:绿虎天牛属chlorophorus是虎天牛族中较原始的类群,刺虎天牛属demonax与艳虎天牛属rhaphuma亲缘关系较近,矮虎天牛属amamiclytus与刺虎天牛属demonax亲缘关系较近。3天牛亚科生物地理学研究本研究针对中国天牛亚科19族87属301种的分布区系进行了研究,就中国天牛亚科属级阶元分布看,只分布在古北界的属有3属,只分布在东洋界的属有25属,古北界和东洋界都有分布的59属。因此,古北界总计分布62属,东洋界分布84属,以东洋成分占优势。其中华南区最为丰富,分布82属,其次是华中区分布75属,西南区分布74属,华北区分布57属,东北区分布36属,蒙新区分布29属,青藏区分布33属。而且多数属为跨区分布:7区的18属、6区的9属、5区的15属、4区的14属、3区的13属、2区的10属、1区7属就中国天牛亚科种级阶元分布看,只分布在古北界的有17种,只分布在东洋界的有176种,古北界和东洋界都有分布的108种,因此,古北界共计分布125种,占总种数的41.5%,东洋界分布284种,占我国总种数的94.4%,我国天牛亚科种类分布以东洋界最为丰富。各动物地理区中分布的种数分别为:蒙新区35种;青藏区28种;东北区55种,华北区101种,华中区165种,华南区228种,西南区183种。在隶属于东洋界的三个地理区中,华南区天牛亚科种类最为丰富,占总数的75.7%,其次是西南区和华中区,分别占我国已知种类的60.8%和54.8%。中国天牛亚科跨区分布和广布的种类较少,分布在1区的90种,2区的84种,3区的39种,4区的45种,5区的27种,6区的11种,7区的5种。分布3区以上的仅127种,占我国己知种类的42.2%;单区和2区分布的种较多,共计174种,占我国已知种类的57.8%,其中,只分布在华南区的就有43种,华南区-西南区分布41种。利用特有简约性分析(pae法)对中国天牛亚科87属301种昆虫进行生物地理学研究发现:天牛亚科昆虫的演化顺序是:首先在蒙新区和青藏区形成,依次是东北区、华北区、华中区、华南区、西南区。因青藏区存在地理位置特殊,地质历史复杂,缺乏生物多样性调查等因素,造成了该区分布物种少的现状,也造成了青藏区在演化顺序中位置靠前。除此区外,蒙新区和东北区比其他生物地理分布区形成要早,分析其原因与第四纪末的冰期活动有关。总体来讲,中国天牛亚科昆虫生物多样性南部地区较北部地区丰富,主要原因可能是:间冰期南部地区森林避难所的形成促进了新物种的形成;北部地区的生态环境较为恶劣造成了生物多样性降低。iv
Cerambycinae is a big group in Cerambycidae,many species are seirous pests for forestand farming. In addition, Cerambycinae conclude many species with complex behavior.Many different classificaitons were hold by several taxonomists, which hindered theresearch progress of Cerambycinae. Based on98characters and more than300speciesof Cerambycinae,the phylogenetic study on Cerambycinae and10selected tribes ismade.3new species,18new record species were found in specimen checking,2subspecies were abolished. The preliminary research of barcode on Clytini, whichcombined data ofNCBI is made. This part of research aimed to analysis the relationshipof several genus of Clyitni. The zoogeography of Cerambycinae is analyzed. The resultsof the dissertation are listed as follow:
     1Systematic of Cerambycinae from China '
     1.1TaxonomyThe new taxa which founded in this research:Artimpaza brevilineata Tian et Chen,2012Demonax proculscuti Li,Tian et Chen,2013Chlorophorus jiangi Li,Tiaci et Chen,2013Chlorophorus copiosus Holzschuh,1991(new record)Rhaphuma lanzhui Holzschuh,1991(new record)Rhaphuma illicata Holzschuh,1991(new record)Demonax traudae Holzschuh,1983(new record)Demonax rosae Holzschuh,1983(new record)Demonax gertrudae Holzschuh,1983(new record)Demonax contrarius Holzschxih,1991(new record)Rhaphuma unigena Holzschuh,1993(new record)Chlorophorus proannulatus Gressitt&Rondon,1970(new record)Stenodryas nigromaculatus (Gardner,1942)(new record)Stenodryas ventralis (Gahan,1906)(new record)Perissus cinericius Holzschuh,2009(new record)Epiclytus insolitus Holzschuh,1991(new record)Chlorophorus ruifmembris Gressitt&Rondon,1970(new record)Kuraruaplauta Gressitt&Rondon,1970(new record)Nyphasia Pascoe,1867(new record genus)N. pascoei Lacordaire (new record) rhaphumanishidaihayashi&makihara,1981(newrecord)chlorophorusconiperdaholzschuh,1992(newrecord)subspeciesabolished:perissusmutabilisvitabilispic,1923perissusmutabilisobscuricolorpic,19371.2phylogeneticanalysesofcerambycinaebasedon98morphologicaldata,whichfrom301species,87genus,19tribes,andthephyligeneticrelationshipsofcerambycinaeisreconstructed.theresultsareasfollows:thechinesecerambycinaeisamonophyleticgroup.(1)obriiniisamonophyleticgroup,stenopteirnicombinedwithkuraruaofcleomeniniassistergroupofobriini.(2)pyrestiniisamonophyleticgroup,erythrusisamonophyleticgroup,pachylocerusisclosedtoerythresthes,pyrestesisamonophyleticgroupwithoutp,pascoei.bayestreedidnotsupportpyrestesasanewtirbepyrestini.njtreeshowedpyrestespartwithotherthreegenuseswithstrongsupport.(3)callichromatiniisamonophyleticgroup;polyzonuswasshowedclosedtoanubis,embrikstrandiaclosedtopachyteria.cataphrodisiumclosedtopachyteneaphrodisiumisaparaphyleticgroup.bayestreehadsomedifferentwithnjtree,therelationshipofmanyothergenushavenodeifniteconclusion.(4)trachyderiniisaparaphyleticgroup.dicelosternusisrelatedtoeuryphagus.thistribeisrelatedtocallichromatini.trachyderiniispartto3branchs,oneispurpuricenus,oneisdicelosternusandeuryphagus,andoneisamarysius,anoplistesandfalsanoplistes.(5)callidiopiniisaparaphyleticgroup.ceresium,stenygrinumandstenodryasareformtoonebranchwithoutparasalpinia.(6)xylotrechusisrelatedtoplagionotus,calloidesisrelatedtoperissus.thephylogenticrelationshipofclytinicannotdiscusswhenusemorphologiccharactersonly.(7)procleomenesofcleomeniniisthesistergrouptomolorchini.dere,artimpaza,cleomenesaremonophyleticgroup;procleomenesrelatedtoartimpaza^nidaisrelatedtokurarua.(8)callidiiniisaparaphyleticgroup.callidiellumisrelatedtoropalopus.semanotus,turaniumaremonophyleticgroup.(9)hesperophaniniisamonophyleticgroup.zoodesisrelatedtostromatium.gnatholeaisrelatedtotrichoferus.everygenusisamonophyleticgroup.(10)phoracanthiniisamonophyleticgroup.(11)cerambyciniisamonophyleticgroup.xoanodera,margites,rhytidodera,neoplocaederusaremonophyleticgroup.aeolesthesisaparaphyleticgroup,a.ningshanensisrelatedtoneocerambyx^andclosedton,oenochrous.nadezhdiellaisaparaphyleticgroup.n.cantonbranchedwithneocerambyx.trirachysrelatedwithii aeolestheswithouta.ningshanensis.(12)xystroceirniisaparaphyleticgroup.3genusesaremonophyleticgroup.hyphusrelatedtoxystrocera.combinedwithmetendosternitetoanalysishyphusandxystrocerashouldbeintwotribes.2dnabarcoderesearchofclytiniinthisstudy,morethan60genesequenceswereobtainedfrom25speciesof9generainchinesecerambycinae.theresultsareasfollows:thedivergenceamongspeciesingenusvariedfrom2.9-16.5%.2%asthedivergencethresholdinclytiniisproper.thedivergenceshowedbarcodecannotuseidentiifcationindifferentgenus.withthedataofncbi,wegotphylogentictreetoshowtherelationshipofclytini:(1)amamiclvtusrelatedtodemonaxspp.,butnotrhaphuma,inaddition,amamiclytuswasconcludedinthebranchofdemonax.(2)therelationofothergenusisconfused,whichneedmoreevidencetoclear.3zoogeographyofcerambycinaeaccordingtothegeneraldataofgeographicaldistirbution,thefaunaofcerambycinaeinchinaisanalyzed.theresultshowsthattheorientalrealmhas84genuses,accountingfor96.6%ofthetotal,andhas26endemicgenuses;thepalaearcticrealmhas62genuses.andonlyhas3genuses.82genusaredistirbutedmsouthchinasub-region,and75genusaredistributedincentralchinasub-region.74genusaredistributedinsouthwestchinasub-region,57genusaredistirbutedinnorthchinasub-region.36genusdistirbutedinnortheastchinasub-region,29genusdistirbutedinmengxinchinasub-region,33genusdistributedinqiangzangchinasub-region.inspecies,theorientalrealmhas284species,accountingfor94.4%.andhas171endemicspecies;thepalaearcticrealmhas125species,accountingfor41.5%.andhas19endemicspecies.228speciesaredistributedinsouthchinasub-region,and165speciesaredistributedincentralchinasub-region.183speciesaredistirbutedinsouthwestchinasub-region,101speciesaredistributedinnorthchinasub-region,55speciesdistirbutedinnortheastchinasub-region.35speciesdistirbutedinmengxinchinasub-region,28speciesdistributedinqiangzangchinasub-region.weusepaetoanalysisthedataandgettheresult:theformingorderofcerambycinaeisbeginininnermongoliaxinjiangregionandqinghai-tibetarea.northeastarea.northchina,centralchina.southchina.southwestarea.qinghai-tibetareaisaspecialregionwithcomplexhistory.asaconclusion,thespeciesinthisareaisthefewest.
引文
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