(5.9)--植物检疫2018-EVA-Qin&Metal.pdf

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1、Evolutionary Applications.2018;114.| Received:16April2018|Revised:8July2018|Accepted:7August2018DOI:10.1111/eva.12701O R I G I N A L A R TI C LEPopulation structure of a global agricultural invasive pest,Bactrocera dorsalis(Diptera:Tephritidae)Yu-jia Qin1*|Matthew N.Krosch2*|Mark K.Schutze2|Yue Zhan

2、g1|Xiao-xue Wang1|Chandra S.Prabhakar2,3|Agus Susanto4|Alvin K.W.Hee5|Sunday Ekesi6|Kemo Badji7|Mahfuza Khan8|Jia-jiao Wu9|Qiao-ling Wang1|Ge Yan1|Li-huan Zhu1|Zi-hua Zhao1|Li-jun Liu1|Anthony R.Clarke2|Zhi-hong Li11DepartmentofEntomology,CollegeofPlantProtection,ChinaAgriculturalUniversity,Beijing,

3、China2SchoolofEarth,EnvironmentalandBiologicalSciences,QueenslandUniversityofTechnology(QUT),Brisbane,Queensland,Australia3DepartmentofEntomology,BiharAgriculturalUniversity,BhagalpurBihar,India4FacultyofAgriculture,PadjadjaranUniversity,Jatinangor,Indonesia5DepartmentofBiology,FacultyofScience,Univ

4、ersitiPutraMalaysia,Selangor,Malaysia6InternationalCentreofInsectPhysiologyandEcology,Nairobi,Kenya7FruitFlyControlProject-ECOWASResponsableComposanteSurveillance.ProjetLuttecontrelesMouchesdesFruits-CEDEAOCRSA,Bamako,Mali8InsectBiotechnologyDivision,InstituteofFoodandRadiationBiology,AtomicEnergyRe

5、searchEstablishment,Savar,Dhaka,Bangladesh9GuangdongInspectionandQuarantineTechnologyCenter,Guangzhou,ChinaThisisanopenaccessarticleunderthetermsoftheCreativeCommonsAttributionLicense,whichpermitsuse,distributionandreproductioninanymedium,providedtheoriginalworkisproperlycited.2018TheAuthors.Evoluti

6、onary ApplicationspublishedbyJohnWiley&SonsLtd*Theseauthorscontributedequallytothiswork.CorrespondenceZhi-hongLi,DepartmentofEntomology,CollegeofPlantProtection,ChinaAgriculturalUniversity,Beijing,China.Email:andAnthonyR.Clarke,SchoolofEarth,EnvironmentalandBiologicalSciences,QueenslandUniversityofT

7、echnology(QUT),Brisbane,QLD,Australia.Email:a.clarkequt.edu.auFunding informationNationalKeyPlanforScientificResearchandDevelopmentofChina,Grant/AwardNumber:2016YFC1200600AbstractBactrocera dorsalis,theOrientalfruitfly,isoneoftheworldsmostdestructiveagri-culturalinsectpestsandamajorimpedimenttointer

8、nationalfreshcommoditytrade.Thegeneticstructuringofthespeciesacrossitsentiregeographicrangehasneverbeenundertaken,becauseunderaformertaxonomyB.dorsaliswasdividedintofourdistincttaxonomicentities,eachwiththeirown,largelynon-overlapping,distribu-tions.Basedontheextensivesamplingofsixapriorigroupsfrom6

9、3locations,ge-netic and geometric morphometric datasets were generated to detectmacrogeographicpopulationstructure,andtodeterminepriorandcurrentinvasionpathwaysofthisspecies.WeakpopulationstructureandhighgeneticdiversityweredetectedamongAsianpopulations.InvasivepopulationsinAfricaandHawaiiarein-ferr

10、edtobetheresultofseparate,singleinvasionsfromSouthAsia,whileSouthAsiaisalsothelikelysourceofotherAsianpopulations.ThecurrentnorthwardinvasionofB.dorsalisintoCentralChinaistheresultofmultiple,repeateddispersalevents,mostlikelyrelatedtofruittrade.Resultsarediscussedinthecontextofglobalquar-antine,trad

11、e,andmanagementofthispest.Therecentexpansionoftheflyinto2|QIN et al.1|INTRODUCTIONThehorticulturalsectorisoneofthelargestwithintheglobalagri-culturaleconomy.Globalfruitandvegetableproductionwasesti-matedat676.9and879.2milliontonnes,respectively,in2013,withglobalfruitexportsinthesameyearvaluedatUSD97

12、.02billion(GyanResearchandAnalytics,2014).WhiledevelopednationssuchastheUnitedStateshavelong-establishedhorticulturalsectors,sig-nificantgrowthincommercialhorticultureisalsooccurringinthedevelopingworld.Anexpandedhorticulturesectorisseennotonlyasamechanismtoincreasehumanhealthindevelopingnations,but

13、alsoasawayofincreasinggenerallivingstandardsthroughcashgenerationfromthesaleoffreshcommoditiesforexport(Virchow&Jaenicke,2016;Weinberger&Lumpkin,2007).Oneofthemajorbiologicalimpedimentstohorticulturalproduc-tionandexportintropicalandsubtropicalregionsoftheworldisthefrugivoroustephritidfruitflies(Dip

14、tera:Tephritidae)(Hendrichs,Vera,DeMeyer,&Clarke,2015).Thetephritidor“true”fruitflies(nottobeconfusedwithdrosophilidfruitflies)laytheireggsintosound,near-ripe fruit on plant,where the resultant larvae feed.Dependingoncommodity,andintheabsenceofcontrols,fruitflydamagecaneasilyleadto80%to100%croploss(

15、White&Elson-Harris,1992).Theglobalfruitflyproblemisexacerbatedbyasmallgroup of highly polyphagous,highly invasive pest species whichcompetitivelydominatelocalfaunaiftheyenterandestablishinaregion(Duyck,David,&Quilici,2004;Duycketal.,2006),andwhichcansubsequentlystopfreshcommoditytradebecauseofthequa

16、r-antinerisktheypose(Dohinoetal.,2016).ThetwobestknownoftheseinvasivetephritidsaretheMediterraneanfruitfly,Ceratitis capitata(Wiedemann),andthefocusofthispaper,theOrientalfruitfly,Bactrocera dorsalis(Hendel).Orientalfruitflyisoneoftheworldsmostinvasiveandpo-lyphagouspestsofagriculture,witharecordedh

17、ostrangeofover250fruitsandvegetables(Clarkeetal.,2005).EndemictotheIndo-Asianregion,theflyfirstestablishedoutsidethisnativerangeinHawaiiin1945,whereitremainsamajorpest(Vargas,Piero,&Leblanc,2015).TheflyhassubsequentlyinvadedthecontinentalUnitedStatesonnumerousoccasionsand,whiletheformalreg-ulatorypo

18、sitionisthatitiscurrentlyabsentfromthecontinentalUnitedStates,debateexistsinthescientificliteratureastowhetheritispermanentlyestablishedinCalifornia(Papadopoulos,Plant,&Carey,2013),orisarepeatinvader(Barretal.,2014).Regardlessof the position in the United States,B.dorsalis is invasive andpermanently

19、establishedinseveralSouthPacificcountries(Vargasetal.,2015),hasinvadedandbeeneradicatedtwiceinAustralia(Cantrell,Chadwick,&Cahill,2002),iscurrentlyactivelyinvadingCentralChina(Chen,Zhang,Ji,Yang,&Zheng,2014),andisan“A1”quarantinepestfortheEuropeanUnion(EPPO,2015).However,itisitsinvasion,spread,andes

20、tablishmentinsub-SaharanAfricathathasreceivedmostattentioninrecenttime.TheflywasfirstdetectedinKenyain2003(Lux,Copeland,White,Manrakhan,&Billah,2003),andwithinaspanof14yearshasspreadacrossallofsub-SaharanAfricaandonlysmallpartsofSouthAfricaremainfreeofthepest(Manrakhan,Venter,&Hattingh,2015).Thecost

21、oflostexportmarketstoAfricaduetotheinvasionhasbeenes-timatedat$2billion(Ekesi,DeMeyer,Mohamed,Massimiliano,&Borgemeister,2016).All facets of research and management of this pest havebeenconfoundedbyitsconfusedtaxonomichistory,withtheflyinrecentdecadesbeingknownunderthenameofnotonlyB.dorsalis,butalso

22、B.invadensDrew,Tsuruta&White,B.papa-yaeDrew&HancockandB.philippinensisDrew&Hancock.Asaresultofamajorinternationalcollaborativeeffort(DeMeyeretal.,2015),theselatterthreespeciesarenowrecognizedasjuniorsynonymsofB.dorsalis(Drew&Romig,2013;Schutze,Mahmood etal.,2015;Schutze,Aketarawong etal.,2015).While

23、thesynonymizationclarifiestaxonomicidentityandhelpssomeaspectsofpre-andpost-harvestcontrolandmarketaccess(Dohino etal.,2016;Hendrichs etal.,2015),they also createnewchallenges.Anorganism,whosegeographicrangeextendsfromAfrica,acrossAsiatothePacific,mightbepredictedtoexhibitmacrogeographicpopulationst

24、ructuring(Ascunceetal.,2011;Gloria-Soria etal.,2016;Virgilio,Delatte,Backeljau,&DeMeyer,2010;Zhang,Edwards,Kang,&Fuller,2014).AstheInternational Plant Protection Convention(FAO,2011)recog-nizes“Pest”as“anyspecies,strainorbiotypeofplant,animalorpathogenicagentinjurioustoplantsorplantproducts,”syn-ony

25、mization of taxa does not negate the issue that differentgeographicpopulationsmaystillshowhighlevelsofpopulationstructuringandsobeofpotentialquarantineandtradeconcernatthe“strain”level.To address this issue,this paper presents the most compre-hensiveglobalassessmentofB.dorsalispopulationstructuringy

26、etundertaken.WesampledfromacrosstheentirerangeofB.dorsa-lisoccurrence,includinginvasivelocations,andusedmorphologicaldata(geometricmorphometricanalysisofwingshape)andmoleculartemperateChina,withveryfewassociatedgeneticchanges,clearlydemonstratesthethreatposedbythispesttoecologicallysimilarareasinEur

27、opeandNorthAmerica.K E Y W O R D SBactrocera dorsalis,geometricmorphometrics,microsatellites,mitochondrialgenes,populationstructure|3QIN et al.markers(cox1 and nad6genesandmicrosatelliteloci)todetermineglobal population structuring in this species.Three independentmarkers were used in an integrative

28、 framework(Schlick-Steineretal.,2010),andinalignmentwithpreviousstudiesofB.dorsalis thathaveshownthesemarkerstobeinformativeatdifferenttem-poralandspatialscalesofpopulationstructuringandinvasionbiol-ogy(Boontop,Schutze,Anthony,Cameron,&Krosch,2017;Schutze,Krosch etal.,2012;Shi,Kerdelhue,&Ye,2012).Us

29、ing DIYABCanalysis,ourglobalpopulationdataalsoallowustomakeinformedcommentonthelikelyoriginofB.dorsaliswithintheIndo/Asianre-gion(anissueunderdebate(Choudhary,Naaz,Prabhakar,&Lemtur,2016)andglobalinvasionpathways.Asecondmajorcomponentofourstudyistodocumentmor-phologicalandgeneticchangesassociatedwit

30、hthecurrent,ongo-ingnorthwardinvasionofB.dorsalisintoCentralChina.Althoughwell-documentedintropicalandsub-tropicalChina(Wan,Nardi,Zhang,&Liu,2011),B.dorsaliswashistoricallyabsentfromCentralChina because of climatic unsuitability(specifically cold stress(Stephens,Kriticos,&Leriche,2007;De Villiers et

31、al.,2016).Nevertheless,B.dorsalis is now able to successfully overwinterincentralChineseprovinces,suchasHubeiProvince(Hanetal.,2011).ThisposesagreatconcernnotonlyforChina,butmustalso to temperate Europe and North America.Climate modelspredicttheseregionstobe“unsuitable”forB.dorsalis(DeVilliersetal.,

32、2016)but,giventheChinesesituation,mustnowbecon-sideredatthreat.Understandinghowthisinvasionisprogressing,whetherthereisongoinggeneflowwiththesourcepopulation/s,andwhethertherearemorphologicalormolecularcharacteristicsassociatedwiththeinvasionfront,canhelpinformmanagementandpreventionofnovelinvasions

33、intoEuropeandNorthAmerica.WithintensivesamplinginChina,weassesswhethertherearemorphologicaland/orgeneticsignaturesassociatedwiththeinva-sionfrontinChinawhichmighthelpinformtheriskposedbythisinvasivepopulation.Takentogether,weaddressthefollowingspecificresearchques-tionswiththesecomprehensivedata:(a)

34、whethermacrogeographicpopulationstructureisdetectableinB.dorsalispopulationsacrossitsentirerange;(b)whatisthelikelyregionoforiginforthespeciesanditsassociatedglobalinvasivepathways;and(c)whatvariationoccursininvasiveCentralChinapopulations.Theresultsaredis-cussedwithrespecttothemanagementofthispest.

35、2|MATERIALS AND METHODS2.1|Sample collectionTwo-thousandeight-hundredandsixty-sevenB.dorsalisadultswerecollectedfrom63locationsandassignedintosixapriorigroups(CentralChina(CC),SouthernChinaandfarnorthernSouth-EastAsia(SCNA),SouthernSouth-EastAsia(SSA),SouthAsia(SA),Africa(AF)andHawaii(HI)basedonbiog

36、eographicalfactors.Thirty-fiveloca-tionsweresampledwithinChina,belongingto16provinces.Detailsof the sampling sites and sampling sizes are given in SupportingInformationTableS1,andthelocalitiesareshowninFigure1.Allsampleswereidentifiedusingavailabletaxonomickeyspriortocon-ductingmolecularanalyses(Lia

37、ng,Yang,Liang,Situ,&Liang,1996;White&Elson-Harris,1992).Thesamplinginvolvedtheuseofmale-onlylures,sofemaleswerenotcollected.Thelegsoraportionofthebodywereremovedforgeneticanalysisandonewing(usuallytheright)forgeometricmorphometricshapeanalysis.TherestofthebodyandtheDNAwerestoredat20Cwithvoucherrefer

38、encesformorphologicalandmolecularverificationatthePlantQuarantineandInvasionBiologyLabinChinaAgriculturalUniversity.2.2|Geometric morphometric analysisUsually,therightwingwasdissectedfromeachflyforslidemount-ing,imagecaptureandanalysis,theleftwasusedinsteadiftherightwingwasdamaged.Wingswereslidemoun

39、tedusingDPXmountingagentandair-driedpriortoimagecaptureusinganAnMoDino-Eyemicroscopeeyepiececamera(model#AM423B)mountedonaLeicaMZ6stereomicroscope.Fifteenwinglandmarkswereselectedfol-lowingSchutze,Jessup,&Clarke,2012anddigitizationusingtps-DIG2v2.16(Rohlf,2010).Rawlandmarkcoordinatedatawereimportedi

40、ntothecom-puterprogramMORPHOJv1.04a(Klingenberg,2011)forshapeanalysis.DatawerefirstsubjectedtoProcrustessuperimpositiontoremoveallbutshapevariation(Rohlf,1999).Multivariatere-gressionofthedependentwingshapevariableagainstcentroidsize(independentvariable)wasconductedtoassesstheeffectofwingsizeonwings

41、hape(i.e.,allometry)(Drake&Klingenberg,2008;Schutze,Jessupetal.,2012).Thestatisticalsignificanceofthisregressionwastestedbypermutationtests(10,000rep-licates)againstthenullhypothesisofindependence.Subsequentanalysesusedtheresidualcomponentsasdeterminedfromtheregression of shape on centroid size to c

42、orrect for allometriceffect.Thesizeofeachwing(centroidsize)wascalculatedinMORPHOJv1.04a.Centroidsizeisanisometricestimatorofsizecalculatedasthesquarerootofthesummeddistancesofeachlandmarkfromthecenterofthelandmarkconfiguration.ANOVAwithposthocTukeystestwasusedtoassessforsignificantdifferencesamongsa

43、mplesites.Sampleswereaprioriassignedtotheabovesixgroups(asforcentroidsizeanalysis)and16provinceswithinChina,fromwhichsubsequentcanonicalvariatesanalysis(CVA)wasappliedtodeter-minerelativedifferencesinwingshapeamonggroups(Kroschetal.,2013).Significant differences were determined via permutationtests(

44、1000permutationrounds)forMahalanobisdistancesamonggroups.Weregressedgeographicdistance(km)betweeneachpairofsamplingsitesdeterminedbyGoogleEarth7.1.7.2606againstMahalanobisdistancescalculatedfromCVAtotestfor“isolation-by-distance”(IbD)effects(Wright,1943).Thestrengthoftheassocia-tionwasdeterminedbyli

45、nearregressionanalysisusingtheprogramSPSSv17.0.4|QIN et al.2.3|Genetic analysisTotalgenomicDNAwasextractedfromeachindividualflyfollow-ingthemanufacturersprotocolfromtheTIANampGenomicDNAkit(DP304,TIANGEN,China)foranimaltissue,andslightmodifica-tionsweremadetoincreaseDNAconcentration(Jiangetal.,2013).

46、Elevenmicrosatellitelociwereusedinthisstudy:MS3,MS3B,MS4,MS5,MS6,Bd9,Bd19,Bd42,Bp198,Bp200,Bi5;technicaldetailsaregiveninDai,Lin,andChang(2004),Aketarawong,Bonizzoni,Malacrida,Gasperi,andThanaphum(2006),Shearmanetal.(2006)and Khamis etal.(2008).Fluorescently labeled fragments werevisualizedonABIPRIS

47、M377GeneticAnalyzerwithROX-500sizestandard.AllelesizewasanalyzedbysoftwareGeneScanversion3.7(AppliedBiosystems,Beijing,China).Gene amplification and sequencing methods were reportedpreviously(Jiangetal.,2013).TheprimersdesignedforthisstudyareshowninSupportingInformationTableS2.Bothdirectionsofthecox

48、1(dividedintotwofragmentsatfirst)andnad6sequencesfromeachindividualwerereviewedusingChromas(version2.33)andassembledusingDNAMAN5.2(LynnonCorporation,Quebec,Canada).Todeletelow-qualitysections,allsequenceswerealignedwiththestandardsequencesofB.dorsalisfromNCBIusingMEGA7.0(Kumar,Stecher,&Tamura,2016)t

49、o generate 1,488-bp cox1 sequencesand525-bpnad6sequences,4,868sequenceswerede-positedinGenBankwithaccessionnumbersMG687532-MG689973forcox1andMG689974-MG692399fornad6.2.3.1|Marker characteristics and intra-population genetic diversityFormicrosatellitedata,thenumberofalleles(NA),numberofef-fectivealle

50、les(NE),observedheterozygosity(HO),andexpectedheterozygosity(HE)werecalculatedusingPOPGENE1.32(Yeh,Yang,&Boyle,1999).Allelicrichness(AR)andgenediversity(HS)werecalculatedusingFSTAT2.9.3.2(Goudet,2002).Frequencyofnullallele(AN)wasestimatedusingGENEPOP4.1(Raymond&Rousset,1995).Forthesequences,thenucle