Abstract
1) Alloxystinae are major secondary parasitoids of aphids, important in both their ecology and pest management. 2) Two radically differing views of alloxystine taxonomy exist in the literature, in one of which the group is very diverse, in the other it consists of a few variable species. 3) We sequenced a variable nuclear gene region (ITS2) for 28 specimens of a morphologically clearly defined group which, in one view belong to a single species and in the other to four species. We find that the four putative species each carry a different unique allele with no intraspecific variation. We show that the probability of the observed distribution of alleles under the assumption of a single interbreeding population is very small and we reject the view that all specimens belong to a single biological species. 4) We discuss the implications of our results for aphid - parasitoid community ecology and the biological control of aphids with parasitoids.
Highlights
IntroductionTnttnmrthtwrdcllicWoceessf(ee1hohihooneeeMpeaoolcrfoooenv9eesfeeoiecggrtfAtTFluovanmmsdw9deheteeoblctiüliittirloeecal8alrreccpcentmilesnoipliyyaelrdexvrlcr)ptcaalolchtroggaoespeys.enruotslliuauttiir,phontmynsprhhchdmdnmipilivsssSseraerngahetbeieepoprupaewasitdaeorde.rmtosonuoameesphseeemlst-cacrphorbttatTrnecpcrdsrapiaoloieheaipeaptfmsalaifi.efitvahrchtme,neemicesety(hranlrrsiensereec.oiHmesisdcycse2rsieda,peptmedaahenalhtao0ssibordesotsatarl(tssnggr.scimou0ihu1iyhcolGintsgitytdnuihitcafi1b9ruentoikietuqd,sTioafseeacokannth)9lcdiiisinuplpnssecl.tbhnddency1epa9ltuiydeaabmesheaitaduf9le)oteoesshra.bracsrehs.seto7laocstaanassnfairticieapr7fossiocfsiiye(ptWTudhsaentaseco)icihoPmml(,atrsta.tshpstaoudaAgphowoeofupheeirpmenhi1ngsonWaruesuaionihacetveaio9didheilihclgtsfvgauidbaincrta9usenusedutdti,shntheeirnbseaeoest4pieomcssegearitilecnract)ea,obelataeiefvaasa.ua,pyosnothdnedrpv.eaptanpploo.enpsS1lfiaaoaaea1rnpdirreehrsbIfrvc9aitdonrnrm9foaeatrncsovhtOatlkpe9ieaycmtt8sorcceliyineberhuieu4nhycmren6taeftdtrseltiyelantb;uescicsbh;acseehmtyehdoilpidetrlsestemheampolrcessAea,hudKothyepicsnqttrifrpnneeotieussuenidaictoectdlunecpnsrapttoruytufdserrhesraiornihdniusepetmrscmpsw2ymnoirntnewapreaanumpehia8ieibfatnrhbnpestasnetDbcoaectmilnciync.otasttetcorertdgotnstjdhfuaoeoyNoescirnielrasergmnsteeyvmnrevdmeqvcertjaiAEeasgooocte,veiatiuapaohstsnsrmtnopndcrofte-hataoyttptpnleoidb2donetuaagealnnentudaert0wfnaairndderledbsdctsnrcsiroadavsx0icaeacieaseeaoif/iiitnosdeiaeeo0oroovologtotniiactcrfi.dhdydnyneeass-;rr,ttf pwrtwrwntmtAttcoFciccacgaacCfsshhohhsaaeepooiolinpaaanreyelhielennieeeeraHrlerrm-imdpllthrsersslnvrdniggarowphygcmeyecsoeenIoiaimmseesxhiiuGaaTcphiaorfmwnmtiwtee..mnysimersaotitouurScmceiovbes)oscToekligionnwaoowrOw2eotdeliodhtpnndebmhinediilurnnwnueettioesurecveepceiirryyndosaaavnfornersarrethhsiu,,f(iuie,nsenetosacdrrdngvopcr1oos(ridr,tbroop1asitiaineE9airleceoiCreoes8mutmmrorpafnacbs8coeuwivrpierwfonmtnhgotinttel6cpmebdameliee,oehagaiswaGsao,tc)crteionsoccdioebtrepeeetuanolitasihlhaheiirfrsrsstoonermbepnp,gvlwlelreueiiesittucntolbminciecenainseihhvheyaodtefuontgcshafgoltryirwnoneeeibaayesloitgafelgvbtnaserhtdneaeedth&ecuewsohernebrshs)iycdttteDnis.tracoiihowuvirec.atnbwtavsyrodvaueiehnBfNaotpweevcydsmeeFuTneccoerecrvhpcdhaiaoaAoihsistdrhrvobaoaionauryhrMeu.AesenudiesibefnrgibrtarpafunpraveioplHioredreoltyonlülvqdewagabgrehliwselenlitffuioeelorbenuoyigolniicisltemptewrtnssxwmeldseeigegeiysfoaltseewaape,plysrlhspyarpoledonrl-wstroetseewkFeeoatadovteueor1tetchaotcfehnesesecifeeodrfiwv9atiadriFniitttdoiridmrfex8etfidtlwhhaeftcsg,aieeaerssithsoe7sonesehsbhweelu.ior.omittip,o.toenegnhofyassahaeBeiTadas(sdWnmsasusteltyrt(HclprntstthEohrgoajms1sthwpuahioosnoieodhelsneoryev9tadepedalaciaroseoaiefsmy9nelgrsietitiarb(busrss,nldmcpeenwh9erec1reoleeseee.poiadodstoaaha)ife9qq(cxfnloeei.teslfcoewruc1oufdsho8uluyarospBsaceiriesyhp9Iwnniee6nctneenspeesesrnsr8aghthnn)oienctcghpihhsist6testdaaniccnmoocoonpeiitptaooie)rnvvhceeeicseeuuonooiotr.eaesssfihoddhdddeeeeessrr,,fftttDeTrbayblsheir1e., HUoKs;t Hdeeta=ilsHaenndlelyo,caOlixtfieosrdoshfitrhee, UAKH;oxBysata= sBpeacyirmeuetnhs, tGheartmwaenrye. aTnhaelysloecda. tiAosn =ofAosucrotl,onBge-rtkesrhmirec,omUKm;unHitoy =stuHdoypeis, Ascot.
MethodsTaxonomic background AAbAttaafsshhaeulbeHlltemecbloTllhooooofuoiAhxnaxxlnAtueiymgyyldegsll0ssCasoltiht.ttoa,lrxo5iihyy.xntny-aahyea2tsApreAhsestilapiptaeCmnnrpiirhnaaddaaghcmisanerardaehieea,detriaowsiiifwlnpiinircsctsdlaiahiiionssetsdcaiomeat)c(tosomi1tehioaptaf9dhwolnnra71Heirdbtshf82ptoaoepoi0a)hacmfmrcdothiktycttmionlawhlaoapaycpeatlgtomtkaetasreiaAlynercreotcagaaghpsgkpsa.tleehiuhlarptnyiIeefre,ortaeoalsiuiarrhsidrlnaasnilioslkitts,iodhsdPifeoottiaoeialPosdvfyferlishamiwdwawdeaatpsooehanoheghfdoirdnrirliccblaodfeodithpetnuspighcth.tpslhoedssiyyIedpyepoAnltbsvenlhwfalirceolEdip(iaooislnwnseuhcoxy,ssoriphsyedmoaauniscnnpcoabtistiaddhdesss.f tdshtaialrWltyweapleawivrraoeesr,fiketpeorrdiitdoowsraittthsohatthameagutrtvmaoiccumktpriitfoxhicfegamrmtoioouurnmpp.hm(Toinylho)e.cgyoicncatalrnlaysbtveteodryeofstihinmeerdilasbreyctaothxnea btWavaaiggfca4EFinorpnarieuunevceaedlrneptrstgeltineaugsoselrnnutnydtcsoiutwhbahwixinpiensseceen5uieadhsogtngiaclowcecososrglpinhtdo,udsneserthoausersh,ueb1ever(gppecn9ya1sntemr,dtwy9c9oodutociish1tne8htparnerfuyoeeno6giiodrbctlprmftns)ndieehstooivyelorecatretfiihmmhausrrdawpaetanficmauaiituiaclhdctsseorraa;trte,algslaeftsepbdrshir(trsacnoroiieEsetAsricocuha:rdrie;vit.ni)rteyeta(neerwd(yoicrrecdni)etsgeiv(cnahuhecrtsdcr)otracespmuaeolliliumutyoenefnioulhs,rcsmnsaupaoeEhoedearlsrrmaodvn.eeecaanfcpelncsrstWryadhndrhiatoantepahiedahlgbhoftsctrluyiefeneaoeeaofaeisnintxc-slrtrirsHhmnw,htntayeeeceeataegnsse1oeoriretrolotl9rrnietwlfntelknwhi7sohvogloe;e4apouyeseifrtsrtswp(srdsamhtoptiuee.hrisnin’)bcesotsessdWohdceifyearpellatihdpgMbAeeeusfnioshmfsc,nctm.btehseoeitugbeetvvrynnil,.dteennsoiihhkcccnywtFgFeodsesttcityrreeisbniheoot&ii3s)trryghexxs-.ff, rgeulsastoedn,(Msheonwksei&t toEvbeenhouuitss,id1e9t9h1e).victrix group and not closely Specimens analysed bo(MefItünshpleleeocArui.eerstv.laiocSl.nt,prgie1x-ct9eig9mrr9moe;unupssntpotuhufdabyetlaiwcsohhefeahdaranedvaieantapcp)hlruwoiddve-eipsdhaiaoirvnanesatihlrtloeiysaidrcseotducnodstyhmidrmaeeseruewtndaiexttlyoal aa1sphiasidfasouafrnutdhllhslpoisestctipeolsfanftothsue.ndspneecaimrbeynsbustenqouteninceodu,r tshtueidryhsoitset.s,Tahbolset (i)thhnaoHshtpvioouioinihlsssrcwlgttitsdsstretshtspihvuaixeeziendwscerdymW.i,vheaoWfossteorhisssitiuteeeeatswntpiipwddcooonaounoeclrilye.ldtaouphsuttdhaisoorteevaodngEebvidsoeavru1suree1iqsnaasiuhrtttsmiieruptoiadeeiecntsmctttvaoih(smcab.1iekres9eeiTsranm7shbos2paesl,ferefec,prtltioihoeemeaemrsslsbilp.gtefeetMrehcdconcorimaueaomccelsrolr,mryo1nhet0sc.loh)iaienspotdpbtehusitcudfigiofnotsheliwnpo.ritteauihIotnnirhnes.tf (((iiiivif)ivmmtysdacncffoocarrlct(((u))hosrr11oiayaionto2suinufslcoocctooecf9n9trputeurdmmarcafhiommsiarr8de7auRsnnecgiacppeeslar,pcG6cu2nerkteinihitnkhhprt,aonabi)tu.b)iAGisidieiieihco.nohsnrraeredeskvaIreiwoisaeliliaignlssssnyihosiodnsshilertr.soisaya(pmgigdHrmnstGssahEtotnnsrhyeiiiUhhiigrdaaatncccietvscewdeteweehrevadrnraaaahiexaeatHeHoaiilCeliysdeiniehwusocntrllfntlgla)hyyacehuifyhedlllbUmbh.rryuedrwrduublp.estutiteiiravdeKEschiienbeihcdaeeAtetsgiuirieoerreheverceenm,,nn.ihmdopnxelcrdrtnlettlatroiiuOadiocon.tybwtn1fhnc(ilcsriwgleuraxphxpna9coaouaodtWofarhM.rl.esuyale8ehstrlsasdsmlrhoooiil.s2itWtiWeefssütotv,utcunamnso)odWht.inlt.efhtoryrenaroeelih(eha,elcmedtln1veetytemihtesoFthrssos9nhsaphfoisdemocieeeBetre8nreisrenfrihGsoqrifqmpsit2rsyfemswregmeimutomurtrit)eeamadthhwuiktoouhaem.yelrqnriae)oies.rosmnmcsesiene.urgenssrshtescd(rycUiaoeoaeutkattct1neWfeeihrnrdfnnafiarnKUf9fdwedcaepsecepirsybGoe9woheudro,aqevyhh.bsw(9odsmvenrdimuanap1iiimaocpaa)lirtdacnnpeo9eegcmhmtnsbeaalhiorndhcunt8ieECcsstriatloraiitaicy6aixsoiimWlnlwprsdcmnaniosxl)atymp)yinfhlhatpyndeeuteopCeoealoioiCsbh,snbcsslnsxstdllthlbeeDiyyofcisfsylaeuhosydmoaodngipanmdspdagsiusFfurxasetpoafvAieti(prEiyamiuwnentcriyeecBonepneoenzolvrigddcsarlmysKiucregpemeotasitelrremissieahVhulnteeitmbxbtnvyfreoslibo^saahyirisrilöeitet.fseescuorrroocwphuasfllbobeeenuAtottklstmxpeiasiioohayadddnnesss---.r.lt.
ResultsAbbUvBtztotigii“ssshhsdnnpheiytyenreccJKeesedtIaFseolgle3cthscsonsymecrowotcra0eeehltnihiabiuht)uixskdm9tneegnrctrra,sdih9epebkaedweneiaaAee6eolfniudn(dcto”ed2G4.stassuceimFuas1i-tfetanlnenoi5lfhli1eqofsblardieigog)eeuequemffllr.a.nsmlenesAeuebeet1eastiTpnnehe.sshqawtsdn.cohtciehwlieuwiynTaeiheavopafe,e,grsesfaelihtfmounnel,ecA.rlrlrateeutleulecerteheh.alemlrele1qlieestedillwgnfitduiuxepasnsooncmwwnseieeechthuneneenrch(yaianasrdels1rwitce(pyeslvdebhta0Eeytkpusoees.dhsr)iuibsMdertenaneIpohfeleaeiqbobnfcdteferefBtndsueroolmwedstdeoiieeiLthllvcindonhslemannsseAoiegaeuetlepscyr(ceufn.rineeiaFittstoplfs“tdaehsceswdu,iinovcrebgfpiedstmiieapochtn.1eehectchrlteuhatfcsadtealiew1eisrvtlycieftntmt)afsiieohopixsiteacsdceStaeiora”“dbihmloicfslnsitnrvapereetelvnoetiaisoiow)le(sca?muslonomtonlasrtionhieegrafifapfnldCoetaillioantbreueeaextdtecoibdhhcoitmdl”ephohneteiseitshamPshensbisaft5pnieefrais.ehanedio0ogtrarrrnr2ahAenAoIokreml.sssd8nd7deees-.:r..t theTnaubllleh2y.poPtahireswisisoefpinrotebrabbrieleitdieinsgopfotphuelaotbiosnesrv. eSdeedatetaxtufnodrear full explanation.victrix tscheki fuscicornis tscheki5 x 10-8 fuscicornis3 x 10-4 0.003 leunisii2 x 1Q-5 5 x 10-4tbApbtuphol(fsohotepraeoosbhpagmeeerlpmsnipotieidcseuscaodlwoiroalylaetviazeteenlinsotetysnsfsihgsdategogpieoatimonmrnerdctteus0ephcafhcaa.v.seiloiuitt0eesedrseapr5saicup“pibsltcssassltoplaaiasoylogekisccthnmnitecineuhdhooctidsipleeamenfatkaasithpotciinotitan”nraotdzsiitthansnctyottnaecamhgh0ttdeleuoo.ila(a0psys0ulicl9aerts.es.caho.sv4sfonpesboe9T“uauoalhr)vTohlnsmb1ayi(ts0ihtcgiTphs(spilet0ebileiarttsoto.ihlrbiay0oeixgalpipd9f”nsertibrp))aavo.b4e2rolipeibowsca)nThf.aaasp=aithaebcuIcahyHtehdisaiuslsreiylisis5opltnmwcyytfnrihaga0moienellsxt.ibnel4hssopensaoeo9eoofibcplnrt1waifbottatlo0tetaieimenhebuotrAnrlyddeearlt.
Summary
TnttnmrthtwrdcllicWoceessf(ee1hohihooneeeMpeaoolcrfoooenv9eesfeeoiecggrtfAtTFluovanmmsdw9deheteeoblctiüliittirloeecal8alrreccpcentmilesnoipliyyaelrdexvrlcr)ptcaalolchtroggaoespeys.enruotslliuauttiir,phontmynsprhhchdmdnmipilivsssSseraerngahetbeieepoprupaewasitdaeorde.rmtosonuoameesphseeemlst-cacrphorbttatTrnecpcrdsrapiaoloieheaipeaptfmsalaifi.efitvahrchtme,neemicesety(hranlrrsiensereec.oiHmesisdcycse2rsieda,peptmedaahenalhtao0ssibordesotsatarl(tssnggr.scimou0ihu1iyhcolGintsgitytdnuihitcafi1b9ruentoikietuqd,sTioafseeacokannth)9lcdiiisinuplpnssecl.tbhnddency1epa9ltuiydeaabmesheaitaduf9le)oteoesshra.bracsrehs.seto7laocstaanassnfairticieapr7fossiocfsiiye(ptWTudhsaentaseco)icihoPmml(,atrsta.tshpstaoudaAgphowoeofupheeirpmenhi1ngsonWaruesuaionihacetveaio9didheilihclgtsfvgauidbaincrta9usenusedutdti,shntheeirnbseaeoest4pieomcssegearitilecnract)ea,obelataeiefvaasa.ua,pyosnothdnedrpv.eaptanpploo.enpsS1lfiaaoaaea1rnpdirreehrsbIfrvc9aitdonrnrm9foaeatrncsovhtOatlkpe9ieaycmtt8sorcceliyineberhuieu4nhycmren6taeftdtrseltiyelantb;uescicsbh;acseehmtyehdoilpidetrlsestemheampolrcessAea,hudKothyepicsnqttrifrpnneeotieussuenidaictoectdlunecpnsrapttoruytufdserrhesraiornihdniusepetmrscmpsw2ymnoirntnewapreaanumpehia8ieibfatnrhbnpestasnetDbcoaectmilnciync.otasttetcorertdgotnstjdhfuaoeoyNoescirnielrasergmnsteeyvmnrevdmeqvcertjaiAEeasgooocte,veiatiuapaohstsnsrmtnopndcrofte-hataoyttptpnleoidb2donetuaagealnnentudaert0wfnaairndderledbsdctsnrcsiroadavsx0icaeacieaseeaoif/iiitnosdeiaeeo0oroovologtotniiactcrfi.dhdydnyneeass-;rr,ttf pwrtwrwntmtAttcoFciccacgaacCfsshhohhsaaeepooiolinpaaanreyelhielennieeeeraHrlerrm-imdpllthrsersslnvrdniggarowphygcmeyecsoeenIoiaimmseesxhiiuGaaTcphiaorfmwnmtiwtee..mnysimersaotitouurScmceiovbes)oscToekligionnwaoowrOw2eotdeliodhtpnndebmhinediilurnnwnueettioesurecveepceiirryyndosaaavnfornersarrethhsiu,,f(iuie,nsenetosacdrrdngvopcr1oos(ridr,tbroop1asitiaineE9airleceoiCreoes8mutmmrorpafnacbs8coeuwivrpierwfonmtnhgotinttel6cpmebdameliee,oehagaiswaGsao,tc)crteionsoccdioebtrepeeetuanolitasihlhaheiirfrsrsstoonermbepnp,gvlwlelreueiiesittucntolbminciecenainseihhvheyaodtefuontgcshafgoltryirwnoneeeibaayesloitgafelgvbtnaserhtdneaeedth&ecuewsohernebrshs)iycdttteDnis.tracoiihowuvirec.atnbwtavsyrodvaueiehnBfNaotpweevcydsmeeFuTneccoerecrvhpcdhaiaoaAoihsistdrhrvobaoaionauryhrMeu.AesenudiesibefnrgibrtarpafunpraveioplHioredreoltyonlülvqdewagabgrehliwselenlitffuioeelorbenuoyigolniicisltemptewrtnssxwmeldseeigegeiysfoaltseewaape,plysrlhspyarpoledonrl-wstroetseewkFeeoatadovteueor1tetchaotcfehnesesecifeeodrfiwv9atiadriFniitttdoiridmrfex8etfidtlwhhaeftcsg,aieeaerssithsoe7sonesehsbhweelu.ior.omittip,o.toenegnhofyassahaeBeiTadas(sdWnmsasusteltyrt(HclprntstthEohrgoajms1sthwpuahioosnoieodhelsneoryev9tadepedalaciaroseoaiefsmy9nelgrsietitiarb(busrss,nldmcpeenwh9erec1reoleeseee.poiadodstoaaha)ife9qq(cxfnloeei.teslfcoewruc1oufdsho8uluyarospBsaceiriesyhp9Iwnniee6nctneenspeesesrnsr8aghthnn)oienctcghpihhsist6testdaaniccnmoocoonpeiitptaooie)rnvvhceeeicseeuuonooiotr.eaesssfihoddhdddeeeeessrr,,fftttDeTrbayblsheir1e., HUoKs;t Hdeeta=ilsHaenndlelyo,caOlixtfieosrdoshfitrhee, UAKH;oxBysata= sBpeacyirmeuetnhs, tGheartmwaenrye. aTnhaelysloecda. tiAosn =ofAosucrotl,onBge-rtkesrhmirec,omUKm;unHitoy =stuHdoypeis, Ascot.
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