RESEARCHARTICLE Predicting the current potential and future world wide distribution of the onion maggot, Delia antiqua using maximum entropy ecological niche modeling ShuoyingNing1,3☯,JiufengWei2☯,JinianFeng1,3* 1 KeyLaboratoryofPlantProtectionResourcesandPestManagement,MinistryofEducation,Entomological a1111111111 Museum,CollegeofPlantProtection,NorthwestA&FUniversity,Yangling,Shaanxi,P.R.China,2 Collegeof a1111111111 Agriculture,ShanxiAgriculturalUniversity,Taigu,Shanxi,P.R.China,3 StateKeyLaboratoryofCropStress a1111111111 BiologyforAridAreas,NorthwestA&FUniversity,Yangling,Shaanxi,P.R.China a1111111111 ☯Theseauthorscontributedequallytothiswork. a1111111111 *[email protected] Abstract OPENACCESS Climatechangewillmarkedlyimpactbiology,populationecology,andspatialdistribution Citation:NingS,WeiJ,FengJ(2017)Predicting patternsofinsectpestsbecauseoftheinfluenceoffuturegreenhouseeffectsoninsect thecurrentpotentialandfutureworldwide distributionoftheonionmaggot,Deliaantiqua developmentandpopulationdynamics.Onionmaggot,Deliaantiqua,larvaearesubterra- usingmaximumentropyecologicalniche neanpestswithlimitedmobility,thatdirectlyfeedonbulbsofAlliumsp.andrenderthem modeling.PLoSONE12(2):e0171190. completelyunmarketable.Modelingthespatialdistributionofsuchawidespreadanddam- doi:10.1371/journal.pone.0171190 agingpestiscrucialnotonlytoidentifycurrentpotentiallysuitableclimacticareasbutalsoto Editor:HugoRebelo,UniversityofPorto, predictwherethepestislikelytospreadinthefuturesothatappropriatemonitoringand PORTUGAL managementprogramscanbedeveloped.Inthisstudy,MaximumEntropyNicheModeling Received:February11,2016 wasusedtoestimatethecurrentpotentialdistributionofD.antiquaandtopredictthefuture Accepted:January18,2017 distributionofthisspeciesin2030,2050,2070and2080byusingemissionscenario(A2) Published:February3,2017 with7climatevariables.Theresultsofthisstudyshowthatcurrentlyhighlysuitablehabitats forD.antiquaoccurthroughoutmostofEastAsia,someregionsofNorthAmerica,Western Copyright:©2017Ningetal.Thisisanopen accessarticledistributedunderthetermsofthe Europe,andWesternAsiancountriesneartheCaspianseaandBlackSea.Inthefuture,we CreativeCommonsAttributionLicense,which predictanevenbroaderdistributionofthispestspreadmoreextensivelythroughoutAsia, permitsunrestricteduse,distribution,and NorthAmericaandEurope,particularlyinmostofEuropeancountries,Centralregionsof reproductioninanymedium,providedtheoriginal UnitedStatesandmuchofEastAsia.Ourpresentdayandfuturepredictionscanenhance authorandsourcearecredited. strategicplanningofagriculturalorganizationsbyidentifyingregionsthatwillneedto DataAvailabilityStatement:Allrelevantdataare developIntegratedPestManagementprogramstomanagetheonionmaggot.Thedistribu- withinthepaperanditsSupportingInformation files. tionforecastswillalsohelpgovernmentstooptimizeeconomicinvestmentsinmanagement programsforthispestbyidentifyingregionsthatareorwillbecomelesssuitableforcurrent Funding:Thisresearchwasfinanciallysupported bytheSpecialFundforAgro-scientificResearchin andfutureinfestations. PublicInterest(GrantNo.201303027).Thefunder hadnoroleinstudydesign,datacollectionand analysis,decisiontopublish,orpreparationofthe manuscript. CompetingInterests:Theauthorshavedeclared thatnocompetinginterestsexist. PLOSONE|DOI:10.1371/journal.pone.0171190 February3,2017 1/15 Predictionofcurrentpotentialandfutureworldwidedistributionofonionmaggot Introduction Climateisoneoftheprincipalfactorsdefiningthepotentialrangeofinsectsandclimate changedirectlyaffectsthedistributionofspecies[1–2].Considerableevidencesuggeststhat theaverageglobaltemperatureswillincreaseby2–4˚Cbetweenthepresentand2100under severaldifferentgreenhousegasemissionscenarios[3–4].Globalwarmingwilllikelyaffect almostallaspectsofinsectlifehistoryandpopulationdynamicssuchasdevelopmentrate,vol- tinism,anddistributionrange[5–8].Risingtemperaturescansignificantlyinfluencethekey physiologicalcharacteristicsthataffectthedistributionandseasonalactivityofinsectpests[9– 10]andchangethelikelihoodofseverepestdamagebyturningclimaticallyunsuitablehabitats intosuitableonesorviceversa[11–12].Becauseofthesensitivityofinsectstoweathercondi- tions,globalclimatechangewillpotentiallydrasticallyalterpestoutbreaks[13–15]. Theonionmaggot,Deliaantiqua,isaworldwideandseriouschronicpestwithanarrow hostrangewithinthegenusAlliumsuchasOnion(AlliumcepaL.),Scallion(Alliumfistulosum L.)andGarlic(AlliumsativemL.)thatsignificantlydamagesplantsinsubtropicalregions throughouttheworld[16–17].Theinitialoccurrenceofonionmaggotwasreportedinthe USAinsomeregionsofWisconsinintheearlytwentiethcentury[18].Thispestcontinuesto causeseriousdamageinAsia,particularlyinChinaandJapan,Europe,andNorthAmerica [19–21].Theonionmaggothasthreeorfourgenerationsannually,andthefirstgeneration causesthegreatesteconomiclossbecausethelarvaekillyoungplants[22].Second-andthird- generationlarvaecauselittledamagetothecroprelativetothefirstgeneration,butfeeding injurycandistortbulbsandallowentryofpathogens,bothofwhichrendercropsunmarket- able[23–24]. Manystudieshaveinvestigatedtheecologyandphysiologyoftheonionmaggot,andthe currentworldwidedistributionofthepestisfairlywellknown.However,itislikelythatthe futuredistributionofthispestwillbeexpandedbecauseofthecontinuedeffectsofglobal warming.Currently,noonehasuseddistributionrecordsandcurrentandpredictedclimactic datatopredictfuturechangesinthedistributionofthispest.MostofAlliumplantshavevery higheconomicvalueandtheyalsorankhighpositionamongvegetablesproductionthrough- outtheworld[25].Thistypeofinformationwillbeabsolutelynecessaryforscientistsand farmersinvolvedinproductionofAlliumcropsthroughouttheworldtodevelopfuturemoni- toringandmanagementstrategiesthancanminimizelosses,particularlyinareaswherethe pesthasnotcausedseriousdamagepriortoglobalwarming. Severaldistributionmodelshavebeendevelopedtoprovideinformationaboutthefuture potentialeffectsofclimatechangeonthefuturedistributionofinsectspecies.Themostwell knownmodelstopredictthedistributionofinsectsbasedonclimacticinputsinclude:GLM, GAM,BRT,BIOCLIM,CLIMEX,GARPandMaxEnt,etc[15,26].Inrecentcomparisons usingseveraltypesofalgorithmstopredictspeciesdistribution,MaxEntwasregardedasthe best-performingmodelusingpresence-onlydataandshowedabetterperformancecompara- bleto16otheralgorithms,suchasgenerallinearmodels(GLM)[27–28].Inseveralprevious studies,MaxEntwasappliedtopredictthepotentialdistributionofspecies,suchas:Stinkbug (Halyomorphahalys)[29],Winterannualgrass(BromustectorumL.)[30],andFruitfly(Bactro- ceradorsalis)[31],andSpinypocketmice(Rodentia:Heteromyidae)[32],etc. Inthisstudy,wecollectedtheoccurrencedataforonionmaggotbasedonextensiverefer- encesandweusedMaxEnttoestimatethecurrentpotentialdistributionofD.antiquaandto predictthefuturedistributionofthisspeciesin2030,2050,2070and2080.Basedontheresults ofthesepredictions,weprovideatheoreticalreferenceframeworkforthemanagementand preventionofonionmaggot. PLOSONE|DOI:10.1371/journal.pone.0171190 February3,2017 2/15 Predictionofcurrentpotentialandfutureworldwidedistributionofonionmaggot Methods Occurrencedataandstudyarea Occurrencedatawerecollatedfromsurveyspublishedintheresearchliterature.Atotalof 128occurrencerecordsforDeliaantiquawereusedinouranalysis.Geo-coordinatesforeach chosendatapointwereeitherreferencedfrominformationpresentedintheliteratureorby usingGoogleEarthcoordinates(S1Table). Occurrencedataareoftenbiasedduetodifferencesinsamplingintensity.Inorderto reducetheimpactforclusteredoccurrencedataonMaxEntmodels,weusedaraster(gridsize approximately10km2)andrandomlyselectedonerecordpercellinthisstudy[33–34].This methodreducedthenumberofoccurrencesto99pointsusedforfurtherstudy. Somestudiesusingenvironmentalvariablesoverlargeareas(eg.allovertheworld)have ledtoconservativepredictionsofcurrentspeciesdistributionandunder-estimatedtheextent ofclimatechange[33–34].Moreover,somestudiessuggestedthattheonionmaggotisa chronicpestofcultivatedAlliumsp.throughouttheHolarcticregion[16,35].Therefore,we selectedNorthAmerica,EuropeandWestAsia,East-AsiaandJapanrespectivelyasthetarget areasinourmodeling. Environmentallayers Currentenvironmentalparameters. Nineteenenvironmentalvariableswhichhad30’spa- tialresolution(~1km)wereusedinourpresentstudy(Bio1-19)(http://www.worldclim.org/) [26,36].Correlationanalyseswereusedtoselectbioclimaticlayers.Weselectedlayersthatwere nothighlycorrelated(r<0.9)becausemanystudieshaveshownthathighlycorrelatedvariables affecttheresultsforspeciesdistributionmodeling(S2Table)[37–40].Furthermore,manypre- viousstudiesofthebiologyandpersistenceofonionmaggotinfestationsshowthattemperature isabasicimpactfactorforsubterraneanpests,suchasDeliaantiqua,becauseitdeterminesthe extentofspreadandinducesspecificphysiologicalbehaviors[18–20,22].Temperaturenotonly determinesthesuitabilityofhabitatsforthispest,butalsoinduceskeyphysiologicalprocesses suchasdiapause[41–42].Thesevenbioclimaticlayersincludedinourmodelingstudieswere obtainedfromWorldclim(Table1)[36]. Futureenvironmentalparameters. Forfutureclimaticpredictions,weuseddownscaled predictionsavailableintheWorldclimdatabasefromover24differentGCMsusedinthe IPCCFourthAssessmentReport[4].TheIPCCSRES(IntergovernmentalPanelonClimate ChangeSpecialreportofEmissionscenarios)includedtwoemissionsscenarios,theA2andB2 scenarios.Toillustratetheworst-caseclimatechangeprediction,wechoseA2asthefuturecli- matescenario.TheA2emissionsscenarioassumesrelativelyrapidpopulationgrowthandrel- ativelyhighglobalCO emissionsincreasingtoalmost5timesthe1990valuesby2100.TheB2 2 Table1. FactorsincludedaspredictorvariablestomodelthepotentialglobaldistributionsofDelia antiqua. EnvironmentalVariables Bio01 Annualmeantemperature(˚C) Bio02 Meandiurnaltemperaturerange(mean(periodmax-min))(˚C) Bio04 Temperatureseasonality(CofV) Bio05 Maxtemperatureofwarmestmonth(˚C) Bio06 Mintemperatureofcoldestmonth(˚C) Bio13 Precipitationofwettestmonth(mm) Bio14 Precipitationofdriestmonth(mm) doi:10.1371/journal.pone.0171190.t001 PLOSONE|DOI:10.1371/journal.pone.0171190 February3,2017 3/15 Predictionofcurrentpotentialandfutureworldwidedistributionofonionmaggot scenariopredictsslowercontinuingpopulationgrowthanddiversetechnologicalchanges resultinginglobalCO emissionsjustdoubletheir1990valuesby2100.Thesescenarioscov- 2 ered7different30yearrunningmeanperiods:2010–2039(2020s),2020–2049(2030s),2030– 2059(2040s),2040–2069(2050s),2050–2079(2060s),2060–2089(2070s)and2070–2099 (2080s).Inthisstudy,4meanperiodswereusedforfutureprediction:2030,2050,2070and 2080.Alldimensionsweresetat30’spatialresolutionforfurtheranalysis. Ecologicalnichemodeling Ecologicalnichesandassociatedpotentialgeographicrangescanbeapproximatedusing correlativealgorithmsthatrelateknownpoint-occurrencedatatodigitalGISdatalayers, andsummarizespatialvariationsintheselayersinmultidimensionalenvironmentalspace [43]. ThemaximumentropymethodasimplementedinMaxEntversion3.3.3softwareisusedto createmodelsofthesuitablecurrentandfuturenichedistributionsofspecies[44–47]andhas beenwidelycontributedtoestimatespeciesdistributions[48–49].Thissoftwarewasusedin ourstudybecauseofthreereasons:1)ithasbeenbetterthanotheralgorithmsinpredicting species’distributionsthatreflectthetruephysiologicalorclimaticconditions[50];2)itper- formsbetterthanothermodelssuchasBIOCLIMandGARPinpredictingtheeffectsofcli- matechangeondistribution[27,51];3)itfindstheprobabilitydistributionofmaximum entropy(thatwhichisclosesttouniform)subjecttoconstraintsimposedbytheobservedspa- tialdistributionsofthespeciesandenvironmentalconditions[45]. Inthisstudy,thealgorithmwasrunforthevariablesatcurrentclimaticconditionswith5 replicates,500iterationsand10000randombackgroundpoints.Then,weperformedmodel- ingprojectionsforfutureclimatescenarios(2030,2050,2070,and2080).WeletMaxEntselect bothsuitableregularizationvaluesandfunctionsofclimatevariablesautomatically,whichit achievesbasedonconsiderationsofsamplesize.MaxEntoutputsacontinuousindex,ranging from0to1,thatisanindicatorofrelativesuitabilityforthespecies,basedontheprincipleof maximumentropy,asconstrainedbytheinputoccurrencedata.Weranthemodelsusingthe defaultregularizationvaluesthathavebeentunedtoperformwellacrossavarietyoforganisms andregions[52]. Modelevaluation Toassesstheperformanceofthemodels,theareasunderthecurve(AUC)metricsofthe receiveroperatorcharacteristics(ROC)curveswereusedinthecurrentstudy[53].TheAUC wasobtainedbythethresholdindependentreceiveroperatingcharacteristic(ROC)analysis [49].AROCcurveshowstheperformanceofamodelwhoseoutputdependsonathreshold parameter,ittestswhetheramodelclassifiesspeciespresencemoreaccuratelythanrandom predictions.AperfectmodelhasanAUCvalueof1,butperformanceisexcellentwhenthe AUCvalueis>0.9[54].Intheprocessofmodeling,80%ofoccurrencelocalitieswereran- domlyselectedastrainingdata,whiletheremaining20%servedfortestingtheresultingmod- els[55]. Toimprovedisplaysofpredictionsinthisstudy,thelogisticoutputofMaxEnt,which rangesfrom0(unsuitability)to1(highsuitability),wasused[52].Conversionofthecontinu- oussuitabilityindexmapstobinaryhabitatandnon-habitatchartsrequiredaprobability thresholdtodeterminethepotentialchangeincurrentandfuturehabitatsofspecies.Todefine habitatsandnon-habitatsforDeliaantiqua,the“maximumtrainingsensitivityplusspecificity” thresholdwasemployedinourstudy[56]. PLOSONE|DOI:10.1371/journal.pone.0171190 February3,2017 4/15 Predictionofcurrentpotentialandfutureworldwidedistributionofonionmaggot Thepredictedresultswereclassifiedintofourlevelsofsuitability:unsuitable(0-threshold), marginallysuitable(threshold-0.4),moderatelysuitable(0.4–0.6)andhighlysuitableareas (0.6–1). WeusedArcview3.3(ESRI)tooutputourmapinourstudies. Results Statisticalmodelevaluation Basedonknownoccurrencesofonionmaggotandcurrentclimatedata,wegeneratedgeo- graphicdistributionmapspredictingareaswhereonionmaggotmightoccur.Themodelper- formanceforthispestwasbetterthanrandom(AUC=0.94fortrainingdataandAUC= 0.923fortestdata);thus,themodelproducedhighlyeffectiveinpredictingthesuitablehabitat areaforthisspecies.Our“maximumtrainingsensitivityplusspecificity”thresholdvalueof 0.29wasobtainedfromthe5timereplication. Currentandpotentialdistribution ThepresentdistributionofDeliaantiquaintheworldaccordingtotheoccurrencerecordswe obtainedfromtheliteratureisillustratedinFig1.ThecurrentclimatepredictionsfromMax- Entshowedthefollowingpotentialdistributionpatternsforthevariouspartsoftheworld:(1) wideareasofNorthAmerica,particularlytheCentralandEasternpartsoftheUnitedStates (especiallytheAtlanticoceancoastalareas,southerncoastalregionsofAlaska,andsome Northernandcentralstates)andtheEasternSouthernareasofCanadaborderingtheUS(2) EuropeandwesternAsia,particularlytheEasternEuropeanandWesternAsiancountries neartheCaspianseaandBlackSea(especiallyEngland,Ireland,Denmark,Ukraine,Turkey andKazakhstan)(3)EasternAsia-MostofChina,Japan,NorthKoreaandSouthKorea(Figs 2–5). Themodelpredictionsforcurrentdistributionandtheactualrecordeddistributiondata shownintheoccurrencerecordsfittogetherverywell.Seventy-ninepercentoftheoccurrence recordsfromaroundtheworldoccurredintheareasthatthemodelpredictedwouldbehighly Fig1.Thecurrentglobaldistributionofonionmaggot,Deliaantiqua.ReprintedfromESRIdataunderaCCBYlicense,with permissionfromEnvironmentalSystemsResearchInstitute,Inc.,originalcopyright2016. doi:10.1371/journal.pone.0171190.g001 PLOSONE|DOI:10.1371/journal.pone.0171190 February3,2017 5/15 Predictionofcurrentpotentialandfutureworldwidedistributionofonionmaggot Fig2.FuturepotentialsuitablehabitatsforDeliaantiquaontheNorthAmericacontinentpredictedby MaxEnt.Thelettersshowpredictionsforthecurrent,2030,2050,2070and2080timeframes(A:Current;B: 2030;C:2050;D:2070;E:2080).Red=highlysuitableareas;Blue=Moderatelysuitableareas;Yellow= Marginallysuitableareas. doi:10.1371/journal.pone.0171190.g002 suitableareasforonionmaggotdevelopment.Themodelpredictionsforareasaroundthe worldthatwouldbemoderatelyormarginallysuitableforonionmaggotdevelopmentare extensive,mainlyincluding:NorthAmerica,CentralandWesternEurope,EasternAsia,and theCaspianandBlackSearegions.IntheUSAandEuropethesepredictedregionsthatareless optimalfordevelopmentofthispestareactuallymuchlargerthanthepredictedhighlysuitable areas.InChina,JapanandNorthKorea,thereareverylargeregionspredictedtobehighly suitable,whichareslightlylessthanthesumofmoderatelyandmarginallysuitableregions. ButinSouthKorea,almostallregionsarepredictedtobemoderatelysuitable. Futureclimatepredictions TheMaxEntmodelswithA2emissionscenariosforpotentialdistributionofDeliaantiquafor 2030,2050,2070,and2080areillustratedin(Figs2–5).Tosimplifythefigures,theglobaldis- tributionissubdividedintothreeregions:NorthAmerica,EuropeandWestAsia,EastAsia. a)NorthAmerica. InNorthAmerica(Fig2),theMaxEntmodelcurrentlypredictsthata fewregionsinNorthernandNorth-easterncoastalstatesofAmerica,suchasNewYork,New Jersey,Ohio,Maryland,Indiana,IllinoisandIowa,havehighlysuitableclimateforDeliaanti- qua.Also,themodelpredictsthatmostoftheMid-northernandNorth-easterncoastalareas ofAmericaaremoderatelyandmarginallysuitableclimaticareasforthisspecies.Inthefuture PLOSONE|DOI:10.1371/journal.pone.0171190 February3,2017 6/15 Predictionofcurrentpotentialandfutureworldwidedistributionofonionmaggot Fig3.FuturepotentialsuitablehabitatsforDeliaantiquaonEuropeandWestAsiacontinent predictedbyMaxEnt.Thelettersshowpredictionsforthecurrent,2030,2050,2070and2080timeframes(A: Current;B:2030;C:2050;D:2070;E:2080).Red=highlysuitableareas;Blue=Moderatelysuitableareas; Yellow=Marginallysuitableareas. doi:10.1371/journal.pone.0171190.g003 predictionsin2030to2080,theselessoptimalsuitableareaswillgraduallyexpandtoinclude moreoftheNorthernandMid-westernstatessuchasKansas,Oklahoma,ColoradoandUtah. Inaddition,themodelpredictsthathighlysuitableareasforonionmaggotinSouthernAlaska willgreatlyincreasebetween2030to2080. InCanada,theSouthernareasofOntarioandQuebecjustnorthoftheUSborderwill remainmostlymoderatelysuitableforthepestinfutureclimacticpredictions.Theunsuitable areasinNewfoundlandandNewBrunswickwillgraduallybecomemarginallysuitablefor onionmaggotby2030andthisareawillcontinuouslyincreaseorbecomeevenmoresuitable regions.Inaddition,thepredictionsshowedthatSouthernNovaScotiaandSouthernNew Brunswickwillbehighlysuitableforthepestby2030,andthensteadilyincrease(Fig2). Ingeneral,MaxEntpredictsthathighlysuitableareasforonionmaggotinNorthAmerica willgraduallyincreaseuntil2080.Also,themoderatelyandmarginallysuitableareaswillgrad- uallyincreasebymorethan39%and32%inthefuture,respectively.Thehighlysuitableareas willalsoincreaseandthenwillstabilizeto10.24sqkmby2080(Fig6A). b)EuropeandWestAsia. TheMaxEntmodelpredictedthatafewareasinNorth-West- ernEuropeandtheWesternAsianregionsneartheCaspianseaandblackSeawillbehighly suitableforOnionmaggotbetween2030to2080,particularlyinSouthernEngland,Southern PLOSONE|DOI:10.1371/journal.pone.0171190 February3,2017 7/15 Predictionofcurrentpotentialandfutureworldwidedistributionofonionmaggot Fig4.FuturepotentialsuitablehabitatsforDeliaantiquaonChina&theKoreacontinentusing MaxEnt.Thelettersshowpredictionsforthecurrent,2030,2050,2070and2080timeframes(A:Current;B: 2030;C:2050;D:2070;E:2080).Red=highlysuitableareas;Blue=Moderatelysuitableareas;Yellow= Marginallysuitableareas. doi:10.1371/journal.pone.0171190.g004 Norway,SouthernSweden,Netherlands,Belgium,South-westernRussia,SouthernTurkey andSouthernKazakhstan,etc.By2080,therangeofmaximallysuitableareasforthispestwill spreadtomanyregionsinNorthernEuropesuchasIcelandandNorth-westernRussia.Also, themodelpredictsthatmoderatelyandmarginallysuitableareasfortheonionmaggotwill greatlyincreasebetween2030to2080.By2080,themodelpredictsthatmostcountriesof Europewillhavesuitablelivingconditionsforthispest,includinglargepartsofWestern Europe,WesternRussiaandNorthernKazakhstan. InEuropeandWestAsia,highly,moderatelyandmarginallysuitableareasforonionmag- gotarepredictedtocontinuouslyincreaseuntil2080,andtheareawillstabilizeto146.35sq km,960.2sqkmand550.65sqkm,respectively(Fig6B). c)EastAsia. InChina,almostalloftheNortheastern,NorthcentralandCentralregions andsomeoftheNorthwesternandSoutheasternregionsarecurrentlyhighlysuitableforthe onionmaggot.TheMaxEntmodelshowsthatthetotalareaofmaximumsuitablelandwill slightlyincreaseinthefuturealthoughthemajorpartofthisareawillshiftfromsouthtonorth PLOSONE|DOI:10.1371/journal.pone.0171190 February3,2017 8/15 Predictionofcurrentpotentialandfutureworldwidedistributionofonionmaggot Fig5.FuturepotentialsuitablehabitatforDeliaantiquaontheJapancontinentusingMaxEnt.The lettersshowpredictionsforthecurrent,2030,2050,2070and2080timeframes(A:Current;B:2030;C:2050; D:2070;E:2080).Red=highlysuitableareas;Blue=Moderatelysuitableareas;Yellow=Marginallysuitable areas. doi:10.1371/journal.pone.0171190.g005 andwillincludescattereddistributionsinmanyprovinces.Thesuitableareaswilldecrease graduallyinsouthernpartsfrom2030–2080andNorthernXinjiangwillbecomemoresuitable. Inaddition,mostofWesternandtheNorthernmostChineseregionswillremainunsuitable foronionmaggotoutbreaks(Fig4). PLOSONE|DOI:10.1371/journal.pone.0171190 February3,2017 9/15 Predictionofcurrentpotentialandfutureworldwidedistributionofonionmaggot Fig6.ComparisonoffuturepotentialsuitablehabitatareasforDeliaantiquabyMaxEntfortimeframesondifferentcontinents: Current,2030,2050,2070and2080.Red=highlysuitableareas;Blue=Moderatelysuitableareas;Yellow=Marginallysuitableareas.A: NorthAmerica;B:EuropeandAsia;C:China&Korea;D:Japan. doi:10.1371/journal.pone.0171190.g006 Currently,NorthernJapan,WesternNorthKorea,andafewregionsinNortheastofSouth KoreaarehighlyclimaticallysuitableforDeliaantiqua(Figs4and5).Themodelpredictsthat theproportionanddistributionofhighlysuitableareaswillcontinuouslyincreaseinKorea andNorthernJapanfrom2030–2080.Incontrasttotheriseinsuitabilityinnorthernregions, SouthernJapanwillbecomeprogressivelylesssuitableinthefuture,andthenwillbecome mostlymoderatelyormarginallysuitableby2080. ThepredictionsforhighlyandmarginallysuitableregionsinChinaandKoreawillincrease 21%and5%,respectively.Incontrast,themoderatelysuitableareaswilldecrease22%by2080 (Fig6C).TheoverallpredictionsforhabitatsuitabilityinJapanwillremainfairlyconstant, exceptthatmanyhighlysuitableregionswillbecomeonlymarginallysuitablehabitat(Fig6D). Discussion Inthisstudy,theMaxEntmodel’scurrentpredictionsofthemostsuitablehabitatsforthe developmentoftheonionmaggotgenerallyagreedwithavailablehostoccurrencerecords. AnalysesshowedthatMaxEntproducedhighlyaccuratepredictionsofAUCvaluethatwas greaterthan0.9.Futuremodelpredictionsfrom2030–2080derivedfromaIPCCclimate changescenarioshowedthatclimaticchangeswouldgreatlyaffecttheworld-widedistribution ofthispest,butthespecificeffectswillvaryamongdifferentlocations.Ingeneral,themodel predictedthattherewouldbeanexpansionofhighlyandmoderatelysuitablehabitatsinmost areasinresponsetoglobalwarming,butthesemoresuitablehabitatswillactuallydecreaseina PLOSONE|DOI:10.1371/journal.pone.0171190 February3,2017 10/15
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