Analysing effects of spatiotemporally distributed species interactions in Maculinea systems [Elektronische Ressource] / vorgelegt von Alexander Singer

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tal2006ersitäteectsingofLeipzigspatiotempausorallyEcologicaldistributedtrespPeciesofin5teractionstheimintMaculineadsystemsDissertationEnzurhErlangung15desacultDoktorgradesysicsderRenNaturwissenscMarburghaftenHeilbronn-Son(Dr.Marburg/Lahn,rer.Departmennat.)ofdemMoFellacUFZhCenbforereicvironmenhResearcPhLeipzig-Halleysikermoserstr.derD-04318Philipps-UnivFersitätyMarburgPhvPhilipps-UnivorgelegtMarburgvthofonD-35032AlexanderAnalysingSingereitgutacVTomProf.FProf.acündlicher:bWisselereicter:h.PhderysikPrüfung:dertPhiliHerrpC.ps-ZwUnivhersitätHerralsPDissertationThomasangenommenagamm22.09.hen200625.10.2006Erstgutach...ten.ts.I.Inmotro.duction27.In.tro.duction.9.I.I.Ecology.and.Mo35del.11.1.Mo.delbdev.elopmen.t2.4.213.1.1.Inthetro.duction....I.tro...............36...2.3.2.............2.4.1...the.Conclusion.......1.3.5.......1.4.........temp.dynamics..13.1.2.Ecology.of.Maculinea.systems........36...........p.................the.................amics14.1.2.1.MaculineaMacu.............3.......tation.del.............remarks.................I.dynamics.opul.2.1........14.1.2.

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ResearcAnalysingDepartmeneectsMarburgofUFZspatiotempForallyHeilbronn-SondistributedMospforeciesermoserstr.inPhteractionsD-35032inMarburg/Lahn,MaculineaofsystemsellDissertationCenzurvironmenErlangungLeipzig-HalledesD-04318DoktorgradesyderPhilipps-UnivNaturwissenscthofhaftenaus(Dr.theimrer.2006nat.)tdemEcologicalFdacinghbtreereicEnhtalPhhysikPder15Philipps-UnivLeipzigersit?tacultMarburgofvysicsorgelegtersit?tvRenon5AlexanderMarburgSingerhVtomagFC.acPhhenbHerrereicZwhHerrPhThomasysikmder25.10.2006Philier:pProf.ps-WisselUniveitgutacersit?tter:alsProf.Dissertation.angenommenTamder22.09.?ndlic2006Pr?fung:Erstgutach.Con.ten.ts.I.In.trodelduction.7PIn.tro.duction.9.I.I.Ecology.and.Mo.del2.211.1.Mo.del.devehaelopmen.t.13Comparison1.1.In.tromoduction......I.duction...............2.3.1...Extinction.............Characterisation...HCET.........Implemen.......Summarising.........oral.3513.1.2.Ecology.of.Maculinea.systems..........2.3...........opulation...............41.system.....44..........14of1.2.1.Maculinea..mo...................25.of.............28.................30.Observing.33.ation.In......14.1.2.2.Host.plan.ts......ds...........................................ypical.............36....16.1.2.3.Myrmica.an.ts......Initial.of.............Discussion...................46.dyn.Macu.......46.the.with.del.....2.4.317.1.2.4.Lo.cal.in.teraction.an.d.spatial51d.i.stribu.ti.on.of.sp1.3.5eciestation.the.del....................17.1.2.5.The1.4HCETremarksmo.del..............................I.I.temp.dynamics.2.opul.dynamics.2.1.tro..............18.1.3.Mo.del.dev.elopmen.t............35.Metho.........................................36.Results..........18.1.3.1.Motiv.ation.of.mo.del.d.esi.gn............36.T.p.dynamics.........................2.3.2......18.1.3.2.Mo.del.description......................2.3.3.b.viour.the.........................2.4......19.1.3.3.Sto.c.ha.s.ticit.y.of.the.mo.del............2.4.1.of.amics.the.mo...............2.4.2.of.Macu.del.the.mo......25.1.3.4.P48arameterisationConclusion...................................3..4aCONTENTS.3.PVorkA.for.in.teracting.sp.ecies.53.3.1.Inttro.duction..........y.........85...h.analyse.......parasitic...A.............................Extinction...approac...e53dynamics3.2.PV88A.in.single.sp.ecies.systems4.5.1...4.5.2.......99...........5.3...................ds............54.3.2.1.Theory....term.......transien.....of.......4.4.1.to.ex.error.......approac.sp.....................the.......p..54953.2.2.Proto.col.for.PVinuenceAeanalysistro.........101...................Metho.........Results.........of..55.3.2.3.Description.of4.2measures.of.p.ersistence..............4.3.....................4.3.1.ersistence55opulation3.2.4.Assessmen.t.of.viabilit81yan.p.........Applicabilit.PV...............t.A-approac.dieren.es.n.4.4.2.the.............Is57appropriate3.2.5inSummasys-ry.of.th.e.ln(1.P.0.(.t.))4.5-metho.d..................dology...............atterns.extinction..58.3.3.DevConclusionelopme.n.t.of.the.m.ulti-sp.ecies.PVIVAspatialmethoAssessingdfrom.5.1.....................F............58.3.3.1.Ov.erview.onexemplaryb.eh.a.viour.of.oscillating.p.opulations104...................104......58.3.3.2.Analytical.dev.elopmen.t.of5.3.3theexamplePV.A.metho.d..........79.Metho....................60.3.3.3.Extrap.olation.for.the.ln(1.P.0.).-PV80A-methoResultsd...............................63.3.4.Application.of.ln(181PLong0p)oftopdieren.t.systems................4.3.2.during.initial.t.hase..............64.3.4.14.4InuyencestheofAsystemhc.haracteristics.on.measures.of.p.ersistence..a.sin.us.mo.de87lUsing.h.PV.h.analyse.t.yp.of.ti.ction.87.Statistical.of.approac.........................4.4.3.the.h.to.complex.teracting.ecies64tems?3.4.2.Sto.c.ha.s.tic.v.a.ri.a.ti.o.ns.inuenc.e.deterministic.p.erio.dicit93yDiscussion.a.predator-.prey.mo.del............................94.Metho.of.analysis.........................94.P.of.opulation.in71systems3.5.Discussion........4.5.3...................................97.The.of.structure.5.landscap.structure.PV.101.In.duction..............74.3.5.1.Extinction.dynamics..................5.2.ramew......................................741023.5.2AnPVapplicationA.for.in.teracting.sp.ecies.systems..................5.3.1.ds....................75.4.Distribution.of.extinction.times.79.4.1.In5.3.2tro.duction..................................105.Conclusion.the.stud.......................105.resultsCONTENTS.5mo5.47.2.4Discussionof...7.2.2.of...8.....of...of...host.....................an.....eness.ess.................174.138....1087.15.4.1.The.meaning.of.p.opulation.p.ersistence.for.theseriesassessmen.tseriesof.landscap.es-108.5.4.2.Suitabilit.y.of.landscap159e.structureoffor.colonisingestingan.d.established.sp.ecies...108del5.4.3.Analysing.spatialastructure.s.and.sp.atial.in.teractions....A...........Analysis......109.5.4.4.Unde.rs.ta.nding.certies.mo.dels.of.habitat.suitabilitby.and.landscapprope.indices.111Maps5.5dynaConclusion..7.2.6.i...Quan.......of.opulation.Revision.......for.s...7.3.1.................Prop.................sp.tro......111.6173Spatialersionhost.distribution.andsystemviabilit.y.1131746.1SummaryIn.tro.duction..............141.index.tro...................143.....................7.2.1................113ulations6.2.Metho.ds............7.2.3.n.of.within.......tication.of.........148.spatial.an.........eectiv.e.con.....151.cell.............7.2.8.eectiv114macroscopic6.2.1.V157ariationanalysisof.host.plan.t.distribution....Spatial.y.t.........elopmen.index.............7.3.2.index......114.6.2.2.Distinguishing.scenario.group165sand.index.........7.4.....................ects:.implicit.8.1........115.6.2.3.V.ariation.of.initial.MyrmicaDescriptionhosteldanthet.distribution......Spatially.ariables.............6.5....115.6.2.4.V.ariation.of.Myrmica.host.an.t.buddin.g.range.............7.spatial.143.In.duction........116.6.3.Results..........................7.2.........................................144.Selection.scenarios..........116.6.3.1.Inu.ence.of.h.ost.plan.t.distribution145.Sim...................................145.Time.of116um6.3.2ersInuMyrmicaencetsofcellsin.i.tial.host.an.t146dQuanisoftribuertiestiontime...................7.2.5.of.distribution.host.t.mics......120.6.4.Discussion..150.'Cell.eness'.Eectiv.dynam.c.tribution.cells.........7.2.7.tifying.eectiv.........................156.Comparison.cell.en.and.p.data.....7.2.9.of.and132.6.4.1.Spatial.analysis.on.the.Maculinea.system..........7.3.index.suitabilit.of.plan.distribution..............1321636.4.2DevSummatrytheof.e.ects......................164.T.the.................................7.3.3134erties6.4.3applicationsHyptheothesis..................168.Discussion.......................................170.Prosp.a.atially.mo1371736.4.4InPductionopulation.dynamical.pro.cesses.pro.v.oking.high.risk.of.extinction.........137.6.4.5.Conserv.ation.asp8.2ectsof.mean.v.of.Macu.del...............8.2.1.implicit.v.........................6195CONTENTS2108.2.2.RulesMeto.transformontheofspatiallyteexplicit.moherdel182inviabilitto.a.spatially.implicitressv)ersionc...t.....85.203.A.1.res.....Ca.line.....meters.T...ersistence.......A.Career.........Synopsis.187.App.P.analysis.solu.for.p....175.8.2.3.Appro205ximationcforethe.exp.ected.p.opulationA.1.2sizesline.inn.horizon.!...209.of.m............176y8.2.4knoMoAdelWissenscequationserdegang...............V.1.Summary.Zusammenfassung.VI.endix.A.opulation.y.205.Analytical.ti.s.measu.of.ersistence................176.8.3A.1.1Comparisonlofulationimplicitthandregressionexplicitparametersmo.del.v.ersions........205.Reg.ion.para.for.i.time.(.H.1......179A.1.38.4asuresDiscussionp.t.and.1.......................Bibliograph.210.c.wledgemen.223.cademic.225.haftlic.W.227.troP7artductionIInAllInobligatorytroIndicatorsductionanEcologicaltheminulti-spseeeciesnarrosystemsHareaimdrivconcept,enbutteriesbetyrestrictedman1998a;yeciesspatiallythedistributedhasproThcesses.MaculineaThereokisofnoopulationsgeneralclosetheoryofonvthetele,inplanterplagyetofspsmallescaleareecologicalliprospcessesconservandjecthoprowparasitismtheyDirectivaectConservdynamicsFattulargerdistributionscalesal(e.g.bitatLevin,Maculinea2000;toWando(Wotton,ynho,2001;MaculineaGreen2004;etofal.,t20ev05).depNonetheless,regionconservVationetofinspwhereeciesvcommpunitieseaninthreatenedorder(IUCN,toBpreservBecauseearebioonsibilitdivexampleersitEUyetrequireshasunderstandingwthehocomplexisystemsHabofeanmanolytintheteractingwingspeeciesthe(Poraloianispetonal.,Maculinea2000;isolatedEbPenmangen&eJonsson,l2005,spandplanrefaneusrences1986;therein).1998a;But,ThereforEuropaeciesparticularetconserv&ationEacaction,iitoneisosucienandtMyrmicatospdevielopthelandscapeteartallymanagemen2005;t2005;measures,2005).whiniaccommhfewarestronglysuitableed,foremthestudycommeunitspyasundervulnerablecon-redsideration.TheyHainvingtheinDirectivmindofaiconcreteEuropcaseincreasedstudyfor,Thisitiisofeasierhto1recogn2002,issefromproIncesseswandrevtheirspatiallyeectsMaculineaonhostspoferformancetofandspListeciesT(Shrader-Fforrecandhettev&eMcCoeects.y,ollo1993;theKrausew&sRuxton,dy2inuence002;spatiotempCrohostwleyandeteciesal.teraction,surviv2005).ofInpthisonthesis,hawsites.earasiticpresenoftusanlivanalysisinframewreorkationfortheircomplexecicsystemshostoftsinhostteractingtsspgenecies.MyrmicaIteidemann,isThomasorienal.,tatedWat2001).aarecenetraleanobspjectiv(buteAlsofal.,natureThomasconservSet-ation:2004).thehpthemersistencesoftosporecieswihostntsashabitat.eralByhostmeanstofecies,aendsimnulationonmogeographicaldel,(Thomasaal.,mTultitude&ofarga,scenariosThomasisal.,testedWitekforal.,theirThesuitabwiteractionlitMyculineatoeciessustainunities,sponlyeciesspcommareunities.inKnoolvwledgemakonthsuitabaierfectlitsystem.yvhelpsEuroptoMaculineadetectecieswhiclistedhnearlyproporertiesinoIUCNflistthe2004).scenariosarenoticeablystedaectAnnexspofeciesEU'ssurvivabitatsal.e.Isomenthethisecconestext,endemic,aemethoandrespforythetheirevation.aluationforofexpresseslandnscapinstallationestheisresearcsuggested,prowhicMacManh(Setteleaccoal.,un2005).tsiforthesissparisenectheiject.esthisbork,ehaeviourtoandeal,ecologicalwprodistributedcesses.ofInonmoretsdetailed1inButteriesvtheestigationsionatfewestronEuropglyRedinuencingasscenarios,andmecohanismssareHabitatanalysedationwhicManagemenh9prothe10terestingINTRorderODUCTIONspaectsepsys-optoulationMaculineadynamicsaoftationthetheoreticalsersistencepofecieselopandvinenconsequencesuitabilitpcersistencesofexptheo.buttery(PVppter,opbulation.teractingFesromhapterourtesndings,methowhostethecancomplexgivelopmenethadvisesapproforclosedtheThismanagemenHotaandsequenceevcaluationhapterofopulationclosedteractinhabitatthesites.isBut,opwithproourextinctiongenericInapprooralactakhtiallywmethoeicdoenotcrestrictelytohosttheoncaseystudycoferoneiMaculineaorkspspatialecies.,Observwationsdonpreliminarytransienthistybtlyehaoviouretitionsorthishost-parasite-mectageohanismstoofafterthehaptersMaculineaestigationsystemtarethirdrelevvidesanfortyinonapbroaderwhileconctextmethoofassessinginMaculineateractingResultsspceciesnewsystems.tAofmaeciesjorthirdpartthesisofectheteractionsthesisiist.dedicatedintoe,theisdevwelopmenevtanofaccordingnewdemands.methosix,ds.isWithtoainuencenalyti-tcaltandersistencenGuidedumericalresults,calculations,enwwedenhtoanceoralasrecenandtlythebofyforWissel.&theSt?hcgivkonersimplication(1991);withStephanEac(1992);iStephanis&tiWisselcan(1994b);indepWisseltheetinal.leads(1994);ortSearliertelterevetofal.adv(1997);asFloranktietoneal.other.(2002);theGrimmws&inWisselin(2004)haptersuggestedwapproacThehcforproptheopulationfoundationviabilitpyviabilitanalysisanalysis(PVA)A)inong-ssingleeciessptems,ecies,inandfourthmakhaetheitdapplicabletestedforpinofteractingpspulations.eciesofsystems.othFhaptersurther,videwineinsighsuggesttoadynamicsmethoindspfsystems.otherpartlandthescapspatiotempespevialuationinwhicarehentaknesaccouninIni-to,accounctvspaeciesdpdeverceptionedofhthehspatialaluastructure.lThesedscapmethostructuresdstoshouldeciesopInenhapternewthispathsdinextensivcommappliedunitinyestigateandoflandscapplaneandecologyan.distributionThepthesisofconsistsbutteries.ofbthreeearlierparts.inThesevrstthparthapter,isededicatedtertoeepainshorttheinspatiotemptrodynamductioncofofthesystemecologicalwsystemonanddevdevtelopmenatindexofhabitataysimFinallyulationinmoeighdel.cInapter,theerstecpreviewhapter,modevelelop-andmenximationtsomeofresults.thehstohaptercnhasticthesisspatiallyaexplicitenrtuandle-basedbsimreadulationendenmoofdelothers.Macustructureisfewpresenccasionsted.toThehsecondreppartofdealsresults.withwnon-spatialer,analysisformofpresentheissystem.anAnus,oitvlerviewwsonenhost-parasite-relydynamicsosectopicalculatedthebTheyofthecMacufollomothedelofisvgivs.en