[PDF] Genetic and morphological variation of butterflies in relict habitats

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Genetic and morphological variation of butterflies in relict habitatsR R dThesisRformatwRIntegratedRArticleeR R R R byR R R R

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Abstract

Habitat fragmentation and loss are leading threatsRtoRglobalRbiodiversityRandRcanRalterRpatternsR ofRdispersalhRpopulationRdynamicshRandRgeneticsRwithRimplicationsRforRlongitermRspeciesR persistencekRMostRhabitatRfragmentationRresearchRhasRfocusedRonRrecentlyRfragmentedRspeciesR thatRhistoricallyRoccupiedRinterconnectedRhabitatRpatcheskRWeRknowRcomparativelyRlittleRaboutR howRnaturallyRfragmentedRspeciesRmayRrespondRtoRhabitatRlosskRForRtheseRspecieshRlocalRhabitatR patchRqualityRmayRinfluenceRtheRdynamicsRandRgeneticsRofRpopulationsRmoreRthanRtheRstructureR ofRtheRsurroundingRlandscapeRdekgkhRdegreeRofRisolationRofRsuitableRhabitatekRIRexaminedRaspectsR ofRtheRecologyRandRevolutionRofRpopulationsRinhabitingRfragmentedRlandscapeshRusingRtwoR butterflyRspeciesRrepresentingRrelictRpopulationsRthatRareRneRrecentlyRfragmentedRbyR anthropogenicRactivitiesRdMormonRmetalmarkhRApodemiaRmormo), and 2) naturally fragmentedRdbogRcopperhRLycaena epixanthe). I assessed patterns of genetic (amplified fragmentRlengthRpolymorphismhRAFLPeRandRflightirelatedRmorphologicalRvariationhRandRtheirR relationshipRtoRmeasuresRofRsurroundingRlandscapeRstructureRandRlocalRhabitatRqualitykRRR RPopulationRgeneticRanalysisRofRtheRanthropogenicallyRfragmentedRMormonRmetalmarkR revealedRaRhighRdegreeRofRspatialRgeneticRstructurehRindicatingRlimitedRgeneRflowhRdespiteRaR smallRgeographicRscaleRdzomRkmekRManagementRofRthisRendangeredRpopulationRshouldRfocusR onRincreasingRconnectivityRamongRtheRmostRisolatedRsubipopulationsRandRthroughRurbanRareaskR ForRtheRnaturallyRfragmentedRbogRcopperhRgeneticRdiversityRwasRexplainedRbyRvariablesRrelatedR toRpatchRqualityRratherRthanRlandscapeRstructurekRMovementRabilityRinRtheRbogRcopperRdinferredR byRflightRmorphologyeRappearedRtoRdependRonRbothRlocalRhabitatRconditionsRandRtheR surroundingRlandscapekRAlsohRusingRanRAFLPibasedRgenomeRscanRapproachhRIRidentifiedR signaturesRofRselectionRinRtheRbogRcopperRassociatedRwithRfineiscaleRlandscapeRheterogeneitykR iiiR My work on the bog copperRhighlightsRtheRimportanceRofRconsideringRtheReffectsRofRlocalR habitatRconditionshRinRadditionRtoRhabitatRisolationhRforRconservationRofRfragmentedR populationskRR RFinallyhRIRalsoRreviewedRtheRcurrentRliteratureRdqtmRarticleseRtoRevaluateRtheRqualityRofR AFLPRdataRusedRinRecologicalRandRevolutionaryRresearchkRIRdiscoveredRaRpervasiveRlackRofR consistencyRandRtransparencyRinRbothRtheRmethodsRusedRtoRassessRdataRreproducibilityhRandRinR theRdetailsRofRmethodologyRpresentedkRThisRworkRhasRidentifiedRanRimportantRpublishingRgapR inRmolecularRecologyRresearchkRR R

Keywords

AmplifiedRfragmentRlengthRpolymorphismhRAdaptiveRgeneticRvariationhRApodemia mormo, ConservationRgeneticshRFlightRmorphologyhRFragmentedRhabitatshRGeneticRdiversityhR GenotypingRerrorRratehRHabitatRfragmentationhRHabitatRqualityhRHostRplanthRIsolationhR LandscapeRheterogeneityhRLepidopterahRLycaenaRepixanthe, MobilityhROutlierRlocihRPopulationR geneticshRRelicthRSpecialistkR ivR

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A version of Chapter oRwasRpublishedRinRMolecular Ecology withRDariaRKoscinskiRandRNushaR KeyghobadiRasRcoiauthorskRDrkRKoscinskiRandRDrkRKeyghobadiRcontributedRtoRstudyRdesignhR helpedRwithRdataRinterpretationRandRprovidedReditorialRcommentsRonRtheRmanuscriptkR RARversionRofRChapterRpRwasRpublishedRinRConservation Genetics withRSylvieR DesjardinsRandRNushaRKeyghobadiRasRcoiauthorskRDrkRDesjardinsRprovidedRlogisticalRsupportR inRtheRfieldhRandRbothRcoiauthorsRcontributedRtoRstudyRdesignhRprovidedRaccessRtoRequipmenthR helpedRwithRdataRinterpretationRandRprovidedReditorialRcommentsRonRtheRmanuscriptkR RChaptersRqRiRsRareRcoiauthoredRwithRNushaRKeyghobadiRwhoRcontributedRtoRstudyR designhRprovidedRequipmenthRhelpedRinterpretRdataRandRprovidedReditorialRcommentsRonRtheR manuscriptskR R vR

AcknowledgmentsR

First and foremost I would like toRthankRmyRsupervisorRDrkRNushaRKeyghobadiRforRherR invaluableRsupportRandRguidanceRthroughoutRmyRdegreekRYouRhaveRbeenRanRinfluentialRmentorR toRmeRbothRacademicallyRandRpersonallyhRandRIRconsiderRyourRleadershipRsomethingRtoRaspireR tokRToRallRtheRmembersRofRmyRlabhRpastRandRpresentwRLiamRFrappehRHeidiRKellerhRChungiYungR dChristinaeRKimhRDrkR Gordana Rasic, Noah Shapiro, Jenna Siu, Eliot Winkler and KristinaRZilichRthankRyouRforR yourRfriendshipRandRsupportkRYouRhaveRmadeRmyRgraduateRexperienceRtrulyRspecialRandR somethingRthatRIRwillRalwaysRlookRbackRonRwithRfondRmemorieskRInRparticularRIRwouldRlikeRtoR thankRSheriRforRteachingRmeRtheRinsRandRoutsRofRworkingRwithRAFLPsRandRKatieRforRhelpingR meRtoRlearnRhowRtoRnavigateRtheRdauntingRworldRofRRRandRmodelRselectionRanalyseskRAsRwellhRIR amRdeeplyRgratefulRtoRDariaRwhoseRcollaborationhRmentorshipRandRfriendshipRhasRbeenRcentralR toRallRmyRsuccessesRthroughoutRmyRdegreekR RIRwouldRalsoRlikeRtoRthankRtheRmembersRofRmyRadvisoryRcommitteeRbothRpastRandR presentwRDrkRBethRMacDougalliShackletonhRDrkRBrockRFentonhRDrkRCherylRPearceRandRDrkR AdamRYateskRYourRvaluableRadviceRandRexpertiseRhasRgreatlyRcontributedRtoRtheRdevelopmentR ofRmyRprojectkR RIRhaveRbeenRveryRluckyRtoRhaveRhadRtheRchanceRtoRworkRwithRmanyRwonderfulRpeopleR throughoutRmyRdegreeRandRwouldRlikeRtoRthankRallRofRtheRvolunteersRandRresearchRassistantsR whoRhaveRhelpedRwithRmyRlabRandRfieldRworkwRWesleyRChickhRAmyRDanghRJennaRDonaldhR OrvilleRDyerhRJaredRHobbshRCurtisRIrvinehRHeidiRKellerhRChungiYungRdChristinaeRKimhRKatieR MillettehRGeorgeRPanghRKevinRSchreiberhRDennisRStiJohnhRLucyRReisshRSueRSeddonhRKatelynR WeaverRandRKatieRWhitekRAsRwellRIRamRalsoRextremelyRappreciativeRtoRtheRfollowingRpeopleR viR who offeredRtechnicalRandRlogisticalRsupporthRserviceRandRaccessRtoRequipmentRasRwellRasR providedRvaluableRadviceRtowardsRmyRprojectwRDrkRBrianRBranfireunhRJasonRDromboskiehRArenR FischerhRJenniferRHillyerhRAngelaRMarinashRDrkRJeremyRMcNeilhRJustinRPetershRAlanRNoonhRDrkR BrentRSinclairhRRickRStronksRandRtheRstaffRatRHarknessRLaboratoryRforRFisheriesRResearchkR RFinancialRsupportRforRmyselfRandRthisRresearchRprojectRwasRgenerouslyRprovidedRbyR TheRCentreRforREnvironmentRandRSustainabilityhRCanadaRResearchRChairsRProgramhR GovernmentRofROntariohRNationalRSciencesRandREngineeringRResearchRCouncilRofRCanadahR TheRXercesRSocietyRforRInvertebrateRConservationhRTheRUniversityRofRWesternRUniversityR DepartmentRofRBiologyhRMalcolmRFergusonRAwardRinRLifeRSciencehRandRaRpartnershipRofRtheR

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Table of Contents

AbstractRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRii

CoiAuthorshipRStatementkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRiv

AcknowledgmentskkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRv

List of TablesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRxii

List of FiguresRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRxvi

List of AppendicesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRxviii

List of Symbols and AbbreviationskkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRxix

List of Software PackagesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRxxi

Chapter 1RkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRn

1 Introduction and literature reviewRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRn

1.1 Habitat fragmentationRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRn

1.2 Relict species and populationsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRo

1.3 Habitat fragmentation versus fragmented habitatsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRp

1.4 Role of habitat quality in fragmented landscapesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRr

1.5 Evaluating the evolutionary potential of fragmented populationsRkkkkkkkkkkkkkkkkkkkkkkkkkkRs

1.6 The role of movement in fragmented landscapesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRt

1.7 Butterflies: important flagship species and a model system for the study of habitat

fragmentationRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRv

1.8 Population genetic studies of butterflies using AFLPs and nonilethalRtissueR

samplingRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnm

1.9 The quality of AFLP data in ecological and evolutionary researchRkkkkkkkkkkkkkkkkkkkkkkRnp

1.10 Dissertation structureRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnq

1.11 Literature CitedRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnu

Chapter 2RkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRpr

viiiR

2 A call for more transparent reporting of error rate: The quality of AFLP data in

ecologicalRandRevolutionaryRresearchRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRpr

2.1 IntroductionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRpr

2.2 MethodskkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRpu

2.2.1 Literature searchkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRpu

2.2.2 Data collectionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRqm

2.3 ResultsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRqo

2.3.1 Overall trendsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRqo

2.3.2 ................................kkkkkkkkkkkkkkRqu

2.4 DiscussionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRqv

2.4.1 Genotyping error reporting in AFLPRstudiesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRqv

2.4.2 Conclusions and recommendationskkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRrr

2.5 Literature CitedRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRrv

3 FineiscaleRgeneticRstructureRofRanRendangeredRpopulationRofRtheRMormonRmetalmarkR

butterflyRdApodemia mormo) revealed using AFLPsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRsp

3.1 IntroductionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRsp

3.2 MethodskkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRst

3.2.1 Study speciesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRst

3.2.2 Sample collectionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRsv

3.2.3 DNA extraction and AFLP analysisRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRsv

3.2.4 Data analysisRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRtp

3.3 ResultsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRts

3.3.1 AFLP analysis and phenotype scoringRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRts

3.3.2 Genetic diversityRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRtt

3.3.3 Population genetic structureRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRtt

3.4 DiscussionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRuo

3.4.1 AFLP analysis and phenotype scoringRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRuo

ixR

3.4.2 Population genetic structure of the Mormon metalmarkRkkkkkkkkkkkkkkkkkkkkkkkkkkRup

3.4.3 Genetic variationRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRut

3.4.4 Future management and recommendationsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRut

3.5 Literature citedRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRuv

Chapter 4RkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRvu

4 Local patch characteristics determine patterns of genetic diversity in a glacial relict,

peatlandRspecialistRbutterflykkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRvu

4.1 IntroductionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRvu

4.2 MethodskkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnmn

4.2.1 Study speciesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnmn

4.2.2 Study areakkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnmo

4.2.3 Sample collectionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnmq

4.2.4 DNA extraction and AFLP marker selectionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnmt

4.2.5 AFLP AnalysisRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnmu

4.2.6 Landscape evaluationRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnnm

4.2.7 Data analysisRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnnp

4.3 ResultsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnns

4.3.1 AFLP analysis and phenotype scoringRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnns

4.3.2 Population structureRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnnv

4.3.3 Genetic diversityRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnom

4.3.4 Effects of landscapeRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnom

4.4 DiscussionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnpn

4.4.1 AFLP analysis and phenotype scoringRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnpn

4.4.2 Population genetic structureRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnpo

4.4.3 Genetic variationRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnpp

4.4.4 Landscape effects on genetic diversityRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnpq

xR

4.4.5 Management and Conservation ImplicationskkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnpv

4.5 Literature CitedRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnqn

Chapter 5RkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnrq

5 Flight morphology corresponds to both broadiRandRfineiscaleRlandscapeRstructureRinRaR

highlyRspecializedRglacialRrelictRbutterflyRdLycaena epixanthe)RkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnrq

5.1 IntroductionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnrq

5.2 MethodskkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnrv

5.2.1 Study speciesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnrv

5.2.2 Study areakkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnsm

5.2.3 Specimen collection and preparationRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnsn

5.2.4 Measurement of morphological charactersRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnso

5.2.5 Landscape evaluationRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnsr

5.2.6 Statistical analysesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnss

5.3 ResultsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnsv

5.4 DiscussionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRntr

5.5 Literature CitedRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnun

Chapter 6RkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnvm

6 Molecular signatures of selection associatedRwithRfineiscaleRlandscapeRheterogeneityRinR

aRrelictRbutterflyhRLycaena epixanthe kkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnvm

6.1 IntroductionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnvm

6.2 MethodskkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnvr

6.2.1 Study sites and data collectionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnvr

6.2.2 DetectingRpotentiallyRadaptiveRlociRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomm

6.2.3 Identifying landscape variables associated with putative outlier lociRkkkkkRomn

6.3 ResultsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomp

6.3.1 AFLP analysis and phenotype scoringRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomp

6.3.2 OutlierRlocusRdetectionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomp

xiR

6.3.3 Associations between landscape variables and putative outlier lociRkkkkkkRomq

6.4 DiscussionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomv

6.4.1 Detecting potentially adaptive loci using an FST-basedRoutlierRapproachRonm

6.4.2 Identifying landscape variables affecting potentiallyRadaptiveRlociRkkkkkkkkRonn

6.4.3 Conclusion and future perspectivesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRonp

6.5 Literature CitedRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRonr

Chapter 7RkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRoor

7 General DiscussionkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRoor

7.1 AFLPswRanRalternativeRmarkerRsystemRforRassessmentsRofRneutralRandRadaptiveR

geneticRvariationRinRbutterfliesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRoos

7.2 Genetics of relictRpopulationsRinRfragmentedRlandscapesRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRoou

7.2.1 Relict populations in anthropogenically fragmented habitatsRkkkkkkkkkkkkkkkkkRoou

7.2.2 Relict populations in naturally fragmented habitats the importance of

localRhabitatRpatchRcharacteristicsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRopn

7.3 Mobility in naturally fragmented relict populationsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRopq

7.4 ConclusionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRopr

7.5 Literature CitedRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRopt

Appendix A: Laboratory protocol used to optimize AFLPs for Lycaena epixanthe kkkkkkRoqp

Appendix B: Landscape data reported in Chapters 3irRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRorm

AppendixRCwRChapterRpRSupplementaryRMaterialkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRorn

Appendix D: Chapter 4 Supplementary MaterialRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRors

Appendix E: Permission to reproduce published materialRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRospR

CurriculumRVitaeRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRosp

xiiR

List of TablesR

Table 2.1 Replicate sample size, as a percentage of the total number of samples in the

datasethRforRstudiesRwhichRassessedRreproducibilityRofRAFLPRdatasetskkRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRrm

Table 2.2 The stage at which replicate samples were generated for studies which assessed

reproducibilitykkRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRrn

Table 3.1RSummaryRofRrepresentativeRstudiesRusingRAFLPsRtoRmeasureRgeneticRdiversityRinR

naturalRpopulationsRofRLepidopterakRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRsu

Table 3.2 Summary of the AFLP phenotype scoring results for all selective primer

combinationskkRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRtv

Table 3.3 Within subipopulationRandRglobalRgeneticRdiversityRforRtheRBritishRColumbiaR

populationRofRApodemia mormo. kkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRtv

Table 4.1 Collection record of the 13 studied populations of Lycaena epixanthe inRAlgonquinR

ProvincialRParkhROntariohRCanadakkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnms

Table 4.2 Summary of the AFLP phenotype scoring results for Lycaena epixanthe for all selectiveRprimerRcombinationsRfollowingRaRzRqaRmismatchRerrorRrateRcriterionkkkkkkkkkkkkkkkkkkkRnnu Table 4.3 Pairwise FST values for the surveyed populations and subipopulationsRdNR©RnreRofR

Lycaena epixanthe inRAlgonquinRProvincialRParkhROntariohRCanadakRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnoo

Table 4.4 Mean population genetic diversity for Lycaena epixanthe inRAlgonquinRProvincialR

ParkhRCanadakkRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnos

Table 4.5 Effect of landscape and patch predictors on estimates of genetic diversity in Lycaena epixanthe for a) all 13 surveyed peatland sites, and b) for all sites except Mizzy

LakeRdMLekRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnot

Table 5.1 Collection records for adult male and female butterflies of Lycaena epixanthe inR AlgonquinRProvincialRParkhROntariohRCanadakRUTMRcoordinatesRdntNeRrepresentRtheRcentroidR

ofRbutterflyRcapturekRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnsp

xiiiR Table 5.2 Absolute mean morphology measurements (± SE) of adult male and female Lycaena expixanthe collectedRfromReightRpeatlandsRinRAlgonquinRProvincialRParkhROntariohR

CanadakkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRntn

Table 5.3 Summary of model selection results for morphological traits in male and female

Lycaena epixanthe.RkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnto

Table 6.1 Collection record of the 15 studied populations of Lycaena epixanthe inRAlgonquinR

ProvincialRParkhROntariohRCanadakkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnvu

Table 6.2 Details of the five selective primer combinations used and their contribution to the

finalRAFLPRdatasetkkRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomr

Table 6.3 AFLP loci identified as putatively under divergent selection for Lycaena

expixanthe kkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRoms

Table 6.4 Summary of model selection results for the five candidate outlier loci identified to

beRunderRdivergentRselectionRbyRDFDISTRandRBayeScankRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomt

Table 6.5 The relative importance of landscape variables acting as potential drivers of genetic

variationRinRtheRfiveRcandidateRoutlierRlociRforRLycaena epixanthe.RkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table A.1 OligonucleotidesRusedRforRtheRAFLPRanalysisRofRLycaena epixanthe kkkkkkkkkkkkkkkkkkRoqr

Table A.2 ProtocolRforRannelaingRaeREcoRI and b) MseI adaptorskRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table A.3 RestrictioniligationRdRiLeRprotocolRoutliningRforRaeREnzymeRMasterRMixhRbeRRiLR

MasterRMixhRandRceRRiLRReactionRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table A.4 PCRRprotocolRforRPreiSelectiveRAFLPRamplificationkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table A.5 PCRRprotocolRforRSelectiveRAFLPRamplificationRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table BknRSummaryRofRtheRlandscapeRvariablesRcollectedRforRtheRnrRAlgonquinRProvincialR

ParkRpopulationsRofRLycaena epixanthe kkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

xivR Table CknRSummaryRofRtheRAFLPRphenotypeRscoringRresultsRforRallRselectiveRprimerR

combinationsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

TableRCkoRWithinRsubipopulationRandRglobalRgeneticRdiversityRforRtheRBritishRColumbiaR

populationRofRApodemia mormo................................kkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table C.3 GeneticRstructureRsummaryRforRtheRBritishRColumbianRpopulationRofRApodemia

mormo kkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

TableRCkqRRobustnessRofRtheRgeneticRbarriersRidentifiedRbyRBARRIERRbasedRonRnmmRpermutedR

geneticRdistanceRmatricesRdFST)kRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table C.5 SummaryRofRtheRprincipalRcoordianteRanalysisRdPCoAeRofRallRstudiedRsubi

populationsRbasedRonRaRgeneticRdistanceRmatrixRdFST)RkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table C.6 SummaryRofRtheRmantelRtestsRresultsRexaminingRtheRrelationshipRbetweenRgeneticR differentiationRdFST/(1iFST) and geographical distance (ln transformed) among all studied subi

populationsRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table D.1 CaptureRrecordRforRindividualsRofRLycaena epixanthe surveyedRinRnpRpeatlandsRinR

AlgonquinRProvincialRParkRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table D.2 PairwiseRFSTRvaluesRforRnpRsurveyedRpopulationsRofRLycaena epixanthein

AlgonquinRProvincialRParkhROntariohRCanadaRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table D.3 SummaryRofRtheRoptimalRAFLPRphenotypeRscoringRparametersRandRassociatedR mismatchRerrorRratesRandRnumberRofRretainedRlociRforRallRselectiveRprimerRcombinationsRkkkkkRomu Table D.4 SummaryRofRtheRoverallRgeneticRstructureRandRintrapopulationRgeneticRdiversityR

resultsRforRLycaena epixanthe inRAlgonquinRProvincialRParkRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table D.5 SummaryRofRanalysisRofRmolecularRvarianceRdAMOVAeRresultsRforRtheRAlgonquinR

ProvincialRParkRpopulationsRofRLycaena epixanthe kkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

Table D.6 SummaryRofRtheRprincipalRcoordinateRanalysisRdPCoAeRforRallRstudiedRpopulationsR

ofRLycaena epixanthe kkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

xvR Table D.7 SummaryRofRtheRmantelRtestRresultsRexaminingRtheRrelationshipRbetweenRgeneticR differentiationRdFST/(1iFST) and geographical distance (log transformed) among all studied

populationsRofRLycaena epixanthe kkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRomu

xviR

List of FiguresR

Figure 2.1 AFLP studies surveyed, categorized by reporting of dataset reproducibility.RkkkkkkRqq Figure 2.2 Within each taxonomic group, the proportion of surveyed AFLP studies where reproducibilityRwasRassessedRandRgenotypingRerrorRrateRwasRreportedRdGE), reproducibility wasRassessedRbutRgenotypingRerrorRrateRwasRnotRreportedRdNGE) and where reproducibility

wasRnotRreportedRatRallRdNR)..RkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRqr

Figure 2.3 Average number of polymorphic AFLP loci retained for final analysis in studies,

groupedRbyRreportingRofRdatasetRreproducibilitykkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRqs

Figure 3.1 Locations of the 14 studied subipopulationsRofRApodemia mormo inRtheR

SimilkameenRRiverRValleyhRBritishRColumbiahRCanadakRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRtm

Figure 3.2 Principal coordinate analysis (PCoA) of all 14 subipopulationsRofRApodemia

mormok kkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRum

FigureRpkpRRelationshipRbetweenRgeneticRdifferentiationRdFST/ (1iFST)) and geographical distanceRdlnRtransformedeRamongRallRstudiedRsubipopulationsRofRApodemia mormok kkkkkkkkkkkkkRun Figure 4.1 Map of the study system and sampling locations for Lycaena epixanthe outliningR theRlocationRofRAlgonquinRProvincialRParkRinROntariohRCanadaRdaehRtheRstudyRregionRwithinR AlgonquinRProvincialRParkRdbeRandRtheRlocationsRofRtheRnpRstudiedRpopulationsRdbhRcekRkkkkkkkkRnmr Figure 4.2 Principal coordinate analysis of all surveyed Lycaena epixanthe populations in AlgonquinRProvincialRParkRbasedRonRaRcovarianceRmatrixRofRpairwiseRFST values.RkkkkkkkkkkkkkkkRnop Figure 4.3 Relationship between genetic differentiation (FST/(1iFST)) and geographical distanceRdlog10 transformedeRamongRaeRallRnrRstudiedRpopulationsRandRsubipopulationshRbeR westernRpopulationsRdBUGhRDLhRMLahRMLbhRWHRandRWReRandRceReasternRpopulationsRdBABhR

EOShRMINahRMINbhROPLhRSBRandRZENekRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnor

xviiR Figure 4.4 Relationships between peatland size (ha) and a) mean cranberry density, b) cranberryRabundancehRandRceRtheRcoefficientRofRvariationRinRcranberryRdensityhRforRpeatlandsRinR

AlgonquinRProvincialRParkhRCanadakRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnov

Figure 4.5 Relationship between catch per unit effort (number of individuals / hour) of

Lycaena epixanthe and a) mean cranberry density andRbeRpeatlandRsizekRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnpm

Figure 5.1 Map of the study system and sampling locations of Lycaena epixanthe: (a) locationRofRAlgonquinRProvincialRParkRinROntariohRCanadayRdbhRceRlocationRofRtheReightRstudiedR

populationskRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnsq

Figure 6.1 Sampling locations of Lycaena epixanthe inRAlgonquinRProvincialRParkhROntariohR

CanadakkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRnvv

xviiiR

List of Appendices

Appendix A: LaboratoryRprotocolRusedRtoRoptimizeRAFLPsRforRLycaena epixanthe kkkkkkkkkkkkRoqpR

AppendixRBwRLandscapeRdataRusedRinRChaptersRqisRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRorm

Appendix CwRSupplementaryRmaterialRforRChapterRpRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRorn

Appendix EwRSupplementaryRmaterialRforRChapterRqkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRors

AppendixRFwRPermissionRtoRreproduceRpublishedRmaterialRkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkkRosp

xixR

List of Symbols and AbbreviationsR

ǻc Difference of Akaike information criterion value between model x and the model with the lowest AIC value ߚ ߚ wi Akaike model weight w+(i) relative variable importance

AIC Akaike information criterion

AICc Second order Akaike information criterion

AMOVA Analysis of molecular variance

ANOVA Analysis of variance

AFLP Amplified fragment length polymorphism

Area Total bog area (ha)

ATP Adenosine triphosphate

BSA Bovine serum albumin

bp Base pair

DNA Deoxyribonucleic acid

dNTP Deoxyribonucleotide triphosphate

EDTA Ethylenediaminetetraacetic acid

FDR False discovery rate

For Proportion of forest habitat

FST Fixation index among sites (the degree of between- relative to within-site genetic variation)

GE Genotyping error rate reported

GIS Geographical information system

GPS Geographical positioning system

He Expected heterozygosity

IBD Isolation-by-distance

ISI Institute for Scientific Information

ISSR Inter simple sequence repeat

logLik Log likelihood

MgCl2 Magnesium chloride

NA Information not available

NaCl Sodium chloride

NGE Reproducibility assessed but genotyping error rate not reported

NGS Next-generation sequencing

nf Number of fixed loci np Number of private loci

NR No assessment of reproducibility reported

PCoA Principal coordinates analysis

PCR Polymerase chain reaction

pH Power of hydrogen

PPL Proportion of polymorphic loci

Prox Mean proximity of potentially suitable wetland habitat

RAPD Random amplified polymorphic DNA

xxR rfu Relative florescence units

RJ-MCMC Reversible jump Markov Chain Monte Carlo

R-L Restriction-ligation

SNP Single nucleotide polymorphism

SSR Simple sequence repeat (microsatellite)

TE Tris-EDTA buffer

Tris Tris(hydroxymethyl)aminomethane

Wat Proportion of open water habitat

BAB Bab Lake

BUG Buggy

COS Costello Creek

DL Dizzy Lake

DT D. Thompson Lake

EOS Eos Lake

KB Kearney

MIN Minor Lake

ML Mizzy Lake

OPL Opeongo Lake

SB Spruce Bog

SUN Sunday Creek

WH Wolf Howl Pond

WR West Rose Lake

ZEN Zenobia Lake

RR xxiR

List ofRSoftwareRPackagesR

AFLPDATRhttpwllwwwknhmkuiolnolenglishlresearchlncblaflpdatlR AFLPSCORERvknkpbRhttpwllwwwkshefkackuklmolecollsoftwarelaflpscore AFLPiSURVRvknkmRhttpwllwwwkulbkackbelsciencesllagevlaflpisurvkhtmlR AICcmodavgRvknkpmRhttpwllCRANkRiprojectkorglpackage©AICcmodavg

ArcGISRvknmkmRESRIRdRedlandshRCAe

ArlequinRvkpknRhttpwllcmpgkunibekchlsoftwarelarlequinplR BARRIERRvokoRhttpwllwwwkmnhnkfrlmnhnlecoanthropologielsoftwarelbarrierkhtmlR BayeScanRvkoknRhttpwllcmpgkunibekchlsoftwarelbayescanlR

EndNoteRxqRThompsonRReutersRdNewRYorkeR

FAFLPcalcRhttpwllwwwkyorkkackuklresldasmahapatralpublicationskhtmlR

FAMDRvknkorRhttpwllwwwkfamdkmekuklfamdkhtmlR

FRAGSTATSRvkqkmRhttpwllwwwkumasskedullandecolresearchlfragstatslfragstatskhtml GenAlExRvkskmRhttpwllwwwkanukedukaulBoZolGenAlExlR GENEMAPPERRvkqkmRAppliedRBiosystemsRdForestRCityhRCAeR

IBMRSPSSRvkomRIBMRCorpRdArmonkhRNewRYorkeR

ImageJRvknkrmRRhttpwllimageknihkgovlijl

JMPRvkuRSASRInstituteRInckRdCaryhRNCeR

lmeqRvkmkvvvvvvioRhttpwlllmeqkriforgekriprojectkorgl

MchezaRhttpwllpopgenlnetlsoftlmchezalR

MuMInRvknkvkrRhttpwllriforgekriprojectkorglprojectslmuminl RRvkpkmkmRRRDevelopmentRCoreRTeamhRhttpwllwwwkriprojectkorglR

SPSSRvkRnskmRSPSSRInckRdChicagoeR

R 1R

Chapter 1R

1RIntroduction andRliteratureRreviewR

nknRHabitatRfragmentationR Habitat fragmentation is the breaking up ofRaRpreviouslyRcontinuousRexpanseRofRhabitatR intoRaRnumberRofRsmallerRsizedRpatcheshRisolatedRfromRoneRanotherRbyRanRunfavourableR landscapeRmatrixRdNossRetRalkRommsyRHabelRandRZachosRomnoekRHabitatRfragmentationRisR oftenRaccompaniedRbyRhabitatRlosshRandRisRconsideredRaRleadingRthreatRtoRbiodiversityR worldwideRdCaughleyRnvvqyRDebinskiRandRHoltRommmyRSalaRetRalkRommmyRFoleyRetRalkRommrehR withRnegativeRimpactsRthatRhaveRbeenRdocumentedRacrossRaRwideRrangeRofRtaxonomicR groupsRincludingRplantsRdHobbsRandRYatesRommpehRinvertebratesRdDidhamRetRalkRnvvsehR amphibiansRdStuartRetRalkRommqehRbirdsRandRmammalsRdAndrénRnvvqekRPredictingRwhetherR populationsRinRhighlyRfragmentedRlandscapesRwillRbeRableRtoRsurviveRandRevolveRinRsmallhR isolatedRhabitatRpatchesRhasRthusRbecomeRaRmajorRresearchRthemeRforRconservationR biologistsRdFazeyRetRalkRommrekRChangesRinRlandscapeRstructureRandRcompositionRassociatedR withRhabitatRfragmentationRresultRinRtheRalterationRofRmanyRbiologicalRprocesseshRincludingR geneRflowRandRgeneticRdrifthRwhichRinfluenceRlevelsRofRgeneticRdiversityRwithinRandRamongR localRpopulationsRdKeyghobadiRommtekRAlterationsRinRpatternsRofRpopulationRgeneticR variationRmayhRinRturnhRaffectRindividualRfitnessRandRevolutionaryRpotentialhRwithR implicationsRforRbothRshortiRandRlongitermRpopulationRviabilityRdFrankhamRetRal. ommoekR ConsequentlyhRconservationRbiologistsRhaveRincreasinglyRbecomeRconcernedRwithRstudyingR theRgeneticRimpactsRofRhabitatRfragmentationRonRpopulationsRdPertoldiRetRalkRommtekRR R 2R RWhileRhabitatsRcanRbeRfragmentedRbyRanthropogenicRactivitiesRsuchRasRagricultureR andRurbanizationhRtheyRcanRalsoRbeRnaturallyRfragmentedRdueRtoRabioticRdekgkhRgeologyhR climateeRandRbioticRdekgkhRpresenceRofRhostshRfoodeRfactorsRdHampeRandRJumpRomnnyRHabelR andRZachosRomnoekRTheRprocessRofRnaturalRhabitatRfragmentationRmayRtakeRplaceRslowlyR overRlongerRtimeRscalesRdekgkhRmillenniayRLomolinoRetRalkRomnmehRinRcontrastRtoR anthropogenicRhabitatRfragmentationRwhichRtypicallyRoccursRwithinRaRrelativelyRshortRtimeR periodRdikekhRcenturieshRdecadesRorRevenRyearsyRLindenmayerRandRFischerRommsekRR R

1.2RRelictRspeciesRandRpopulationsR

Species or populations which occur as small and geographically isolated remnants of a formerlyRmoreRwidespreadRdistri(CasseliLundhagenRomnmyR LomolinoRetRalkRomnmeRandRtheirRformationRcanRbeRattributedRtoReitherRhistoricalRorR contemporaryRhabitatRfragmentationkRForRexamplehRmanyRarcticialpineRspeciesRwhichRareR currentlyRrestrictedRtoRdisjunctRareasRonRmountainRtopsRandlorRmoreRNorthernRlatitudesR dekgkhRmountainRavensRfloweringRplanthRDryas octopetala, SkredeRetRalkRommsyRvioletRcopperR butterflyhRLycaena helle, Habel et al. 2011eRareRrelictsRspeciesRwhoseRonceRwidespreadR distributionsRhaveRbeenRgraduallyRrestrictedRbyRwarmingRtemperaturesRandRtheR accompanyingRlossRofRsuitableRhabitatRfollowingRtheRlastRglacialRperiodRdHabelRetRalkR omnmekROnRtheRotherRhandhRspeciesRwhichRwereRuntilRrelativelyRrecentlyRquiteRprevalentRandR widespreadRmayRhaveRbecomeRspatiallyRrestrictedRtoRremnantRrelictRpopulationsRasRresultR ofRhumaniinducedRhabitatRlossRandRfragmentationRdekgkhRNorthRAmericanRbrownRbearhR Ursus arctos, Paetkau et al. 1998; regal fritillary butterfly, Speyeria idalia, KeyghobadiRetR alkRomnoekRInRsomeRcaseshRtheRsmallhRisolatedRnatureRofRnaturallyRformedRrelictRhabitatR 3R patchesRmayRbeRfurtherRexacerbatedRbyRcontemporaryRfragmentationRprocessesRRRRRRRRRRRR dekgkhRinlandRhypersalineRenvironmentsRinRWesternRMediterraneanhRGómezRetRalkRommrekR DueRtoRtheirRlimitedRdistributionsRandRincreasingRrarityhRmanyRrelictRspeciesRandR populationsRareRnowRofRhighRconservationRconcernRdHabelRetRalkRomnmekRAsRtheRoverallRrateR andRmannerRinRwhichRhabitatsRbecomeRfragmentedRdikekhRanthropogenicRvskRnaturaleRcanR causeRdivergentRevolutionaryRandRgeneticRtrajectoriesRforRpopulationsRdMacDougalli ShackletonRetRalkRomnnyRHabelRandRZachosRomnoehRitRisRvitalRthatRmanagementRstrategiesRforR relictRspeciesRandRpopulationsRconsiderRtheRhistoryRofRtheRlandscapeRandRtheRprocessesRbyR whichRpopulationRisolationRhasRarisenkR

1.3RHabitatRfragmentationRversusRfragmentedRhabitatsRR

Species which occur in naturallyRinterconnectedRhabitatsRareRpredictedRtoRbeRnegativelyR affectedRbyRsuddenRhumaniinducedRhabitatRfragmentationRdHabelRandRZachosRomnoekRInR theoryhRsmallerRandRmoreRisolatedRhabitatRpatchesRwillRsupportRsmallRpopulationsRwhichR experienceRreducedRgeneRflowRandRincreasedRgeneticRdriftRasRaRresultRofRdecreasedR dispersalRandRsmallRlocalReffectiveRpopulationRsizeshRrespectivelyRdKeyghobadiRommtekR OverRtimehRgeneticRdiversityRwithinRpopulationsRwillRerodeRasRaRresultRofRincreasedRgeneticR driftRandRreducedRgeneRflowhRatRtheRsameRtimeRincreasingRgeneticRdifferentiationRamongR populationsRdTempletonRetRalkRnvvmyRFrankhamRetRalkRomnmekRAdditionallyhRintrapopulationR geneticRdiversityRmayRbeRfurtherRlostRandRgeneticRdifferentiationRincreasedRthroughR bottlenecksRandRlocalRextinctionsRwhichRoftenRaccompanyRpopulationRfragmentationR dGilpinRnvvnyRAndersenRetRalkRommqyRKellerRetRalkRommqyRBroquetRetRalkRomnmekRTheR disruptionsRtoRtheRdriftigeneRflowRequilibriumRwhichRoccurRinRrecentlyRfragmentedR 4R populationsRareRalsoRpredictedRtoRleadRtoRanRincreaseRinRtheRoccurrenceRofRinbreedingRandR inbreedingRdepressionhRasRwellRasRtheRrandomRfixationRofRdeleteriousRmutationsRandRaRlossR ofRadaptiveRpotentialRdLandeRnvvuyRKellerRandRWallerRommoyRFrankhamRetRalkRomnmekR ChangesRinRtheseRgeneticRprocessesRwillRcontributeRtoRanRoverallRdeclineRinRpopulationR viabilityRasRwellRasRtheRabilityRtoRrespondRtoRfutureRenvironmentalRchangehRandRthusR increaseRtheRprobabilityRofRextinctionRdReedRandRFrankhamRommpyRFrankhamRommraekR RInRcontrasthRorganismsRwhichRoccupyRnaturallyRfragmentedRhabitatsRareRtypicallyR specialistRspeciesRadaptedRtoRlivingRwithinRaRhabitatimatrixRmosaicRdHabelRandRZachosR omnoekRBecauseRofRtheirRecologicalRspecialisationhRpopulationsRareRgeographicallyRisolatedR inRdiscreteRhabitatRpatchesRandRexperienceRlimitedRgeneRflowkRAsRaRresulthRlocalR populationsRareRtypicallyRsmallRinRsizeRandRexhibitRlowerRgeneticRdiversityhRasRwellRasR increasedRinterpopulationRgeneticRdifferentiationRdHabelRandRSchmittRomnoekRInterestinglyhR howeverhRtheseRspecialistRspeciesRappearRtoRhaveRtheRcapacityRtoRpersistRoverRlongRperiodsR ofRtimeRdespiteRtheRgeneticRisolationRofRpopulationsRdHabelRandRSchmittRomnoekRItRisR theorizedRthatRdeleteriousRallelesRmayRhaveRbeenRpurgedRfromRtheseRpopulationsRoverR manyRgenerationsRdFrankhamRetRalkRommnyRHabelRandRZachosRomnoehRandRthatRtheirRcurrentR geneticRmakeiupRconsistsRofRaRsmallRnumberRofRallelesRwhichRareRhighlyRadaptedRtoRlocalR habitatRconditionsRdWattRetRalkRommpyRKarlRetRalkRommuekRThushRlowRgeneticRdiversityRinRtheseR populationsRisRactuallyRassociatedRwithRtheRabilityRtoRresistRtheRnegativeRconsequencesRofR geneticRbottleneckshRinbreedingRorRfluctuationsRinRlocalRpopulationRsizeRdCrnokrakRandR BarrettRommoyRReedRomnmeRandRmeansRthatRpopulationsRareRnotRdependentRonRgeneRflowRforR geneticRrefreshmentRdHabelRandRSchmittRomnoekRIndeedhRseveralRstudiesRhaveRnowR documentedRlongitermRpersistenceRofRgeneticallyRdepauperateRpopulationsRlivingRinR 5R geographicRisolationRdekgkhRblackRmangroveRtreehRAegiceras corniculatum; Ge and Sun nvvvyRSanRNicolasRIslandRfoxhRUrocyon littoralis dickeyi, Aguilar et al. 2004; ChillinghamR cattlehRBos taurus, Visscher et al. 2001; RedRApolloRbutterflyhRParnassius apollo, Habel et alkRomnoekRHoweverhRwhileRlowRgeneticRvariationRmayRnotRposeRanRimmediateRconcernRforR populationsRlivingRunderRisolatedRconditionshRtheRadaptiveRscopeRofRpopulationsRtoRrespondR toRfutureRenvironmentalRchangeRwillRsurelyRbeRlimitedkRR ClearlyRlandscapeRhistoryRisRimportantRtoRconsiderRwhenRstudyingRtheRecologyRandR geneticsRofRspeciesRinhabitingRdiscreteRhabitatRpatchesRdHabelRandRZachosRomnoekR AlthoughRmanyRnaturallyRpatchyRhabitatRtypesRareRnowRbecomingRfurtherRfragmentedRdueR toRcontemporaryRanthropogenicRactivitieshRtheRspecialistsRwhichRoccupyRtheseRhabitatsRmayR notRbeRasRnegativelyRaffectedRbyRincreasedRhabitatRisolationRasRotherRspeciesRbecauseRtheyR haveRessentiallyRadaptedRtoRpersistRunderRconditionsRofRlowRgeneRflowRandRgeneticR diversitykRForRsuchRspecialistshRlocalRhabitatRpatchRcharacteristicsRdekgkhRhabitatRsizeRandR qualityeRmayRbeRmoreRlikelyRtoRaffectRpopulationRpersistenceRthanRhabitatRisolationkRAR recentRtheoreticalRsimulationRsupportsRthisRhypothesisRdYeRetRalkRomnpehRhoweverhRempiricalR evidenceRofRthisRrelationshipRisRlimitedkRR

1.4RRoleRofRhabitatRqualityRinRfragmentedRlandscapesR

In the context of habitat fragmentation and species conservation, most theoretical and empiricalRresearchRtoRdateRhasRfocusedRonRunderstandingRhowRhabitatRpatchRareaRandR isolationRinfluenceRdispersalhRpopulationRsizeRandRtheRpatch areai HanskiRandRGaggiottiRommqyRPrughRetRalkRommueRwithR 6R considerablyRlessRemphasisRonRtheRroleRofRwithinihabitatRpatchRqualityRdYeRetRalkRomnpekR HoweverhRthereRisRanRincreasingRbodyRofRevidenceRwhichRindicatesRthatRheterogeneityRinR qualityRamongRhabitatRpatchesRcanRbeRanRextremelyRimportantRdriverRofRlocalRpopulationR dynamicsRandRlongitermRpersistenceRinRfragmentedRlandscapesRdThomasRetRalkRommnyR BaguetteRetRalkRomnnyRYeRetRalkRomnpeRandRthusRshouldRbeRincludedRinRecologicalRmodelsR dFleishmanRetRalkRommoyRSchooleyRandRBranchRommtekRSeveralRstudiesRhaveRnowR demonstratedRtheRnegativeRimpactsRofRdecreasedRhabitatRqualityRonRtheRabundanceRandR distributionRofRpopulationsRoccupyingRfragmentedRlandscapesRdekgkhRDennisRandREalesR nvvtyRThomasRetRalkRommnyRFleishmanRetRalkRommoyRKraussRetRalkRommqekRInRcontrasthRthereR haveRbeenRveryRfewRstudiesRtoRdateRwhichRhaveRsoughtRtoRevaluateRtheReffectsRofRspatialR heterogeneityRinRhabitatRpatchRqualityRonRmeasuresRofRpopulationRgeneticRstructureRandR diversityRdbutRseeRPorlierRetRalkRommvyRPitraRetRalkRomnnyRAldaRetRalkRomnpekRInRtheoryhRhigherR qualityRhabitatRpatchesRshouldRsupportRlargerRpopulationsRwhichRareRmoreRgeneticallyR diverseyRhoweverhRempiricalRevidenceRofRthisRrelationshipRisRstillRlargelyRlackingRddeRVereR etRalkRommvyRPitraRetRalkRomnnekRUnderstandingRofRtheRgeneticRconsequencesRofRchangesRinR habitatRqualityRinRadditionRtoRhabitatRpatchRsizeRandRisolationRwillRbeRenableRusRtoRmoreR accuratelyRpredictRtheRdynamicsRandRlongitermRpersistenceRofRpopulationsRinRfragmentedR landscapeskRR R

1.5REvaluatingRtheRevolutionaryRpotentialRofRfragmentedR

populationsR Given the low levels of genetic diversity characteristicRofRfragmentedRpopulationshRbothR anthropogenicallyRandRnaturallyRinducedhRaRmajorRchallengeRinRspeciesRconservationRisR 7R howRtoRmaintainRtheRevolutionaryRpotentialRofRpopulationsRsoRthatRtheyRhaveRtheRabilityRtoR respondRtoRfutureRenvironmentalRstressorsRsuchRasRclimateRchangehRintroducedRdiseasesRorR parasitesRdFrankhamRommrbyRAllendorfRetRalkRomnmekRMostRstudiesRofRtheRgeneticReffectsRofR habitatRfragmentationRtoRdateRhaveRfocusedRonRneutralRgeneticRvariationRthroughRwhichRtheR relativeRinfluencesRofRgeneticRdriftRandRgeneRflowRonRfragmentedRpopulationsRcanRbeR inferredRdKeyghobadiRommtekRHoweverRneutralRmarkersRdoRnotRevolveRdirectlyRinRresponseR toRselectionhRandRwhetherRestimatesRofRneutralRgeneticRdiversityRaccuratelyRreflectRadaptiveR potentialRisRdebatedRdLuikartRetRalkRommpyRHoldereggerRetRalkRommuyRFrankhamRomnmekR ThereforehRinRorderRtoRfullyRunderstandRtheReffectsRofRhabitatRfragmentationRonRpatternsR andRlevelsRofRgeneticRdiversityhRadaptiveRgeneticRvariationRshouldRalsoRbeRexaminedR dHoldereggerRetRalkRomnmekRThisRwillRprovideRforRmoreRaccurateRassessmentsRofRlocalR populationRviabilityRandRpredictionsRonRwhetherRpopulationsRhaveRtheRcapacityRtoRrespondR toRchangingRenvironmentalRconditionsRdAllendorfRetRalkRomnmekRThisRisRanRareaRofRresearchR whichRhasRupRuntilRrecentlyRbeenRrelativelyRunderexploredkRHoweverhRnewRgenomicR methodsRandRanalyticalRtoolsRprovidedRthroughRtheRemergingRdisciplinesRofRpopulationRandR landscapeRgenomicsRdLuikartRetRalkRommpyRStorzRommryRJoostRetRalkRommtyRManelRetRalkRomnmeR nowRmakeRitRpossibleRtoRmoreRreadilyRstudyRpatternsRofRadaptiveRgeneticRvariationRinR naturalRpopulationsRofRnonimodelRorganismsRdHoldereggerRetRalkRommuekRR R

1.6RTheRroleRofRmovementRinRfragmentedRlandscapesR

Particularly for small, isolated populations inhabiting fragmented landscapes, dispersal is aRkeyRlifeihistoryRtraitRwhichRinfluencesRlocalRandRregionalRpopulationRdynamicshR populationRgeneticsRandRadaptiveRevolutionRdHanskiRnvvvyRRonceRommtyRNitepõldRetRalkR 8R ommvyRBentonRandRBowlerRomnoekRDispersalRabilityRorRtendencyRmayRitselfRbecomeRalteredR dueRtoRchangesRinRlandscapeRstructureRhoweverhRtherebyRaffectingRmanyRaspectsRofRtheR ecologyRandRevolutionRofRpopulationskRThereforehRunderstandingRhowRmovementRabilityRorR propensityRisRinfluencedRbyRhabitatRfragmentationRmayRallowRusRtoRbetterRpredictRhowRtheR dynamicsRandRgeneticsRofRlocalRpopulationsRwillRrespondRtoRlandscapeRchangeRdStevensR andRCoulonRomnoekRIncreasingRisolationRofRhabitatRpatchesRcouldRpotentiallyRselectRforR individualsRwithRmoreRmobileRphenotypesRwhoRareRableRtoRtraverseRlongerRdistancesRdekgkhR TaylorRandRMerriamRnvvrekRConverselyhRbecauseRmovementRbetweenRhabitatRpatchesRisR physicallyRcostlyRandRassociatedRwithRmanyRriskshRincreasedRisolationRofRpatchesRmayR selectRforRaRdecreasedRdispersalRpropensityRdekgkhRDempsterRnvvnyRSchtickzelleRetRalkRommsekR SuchRchangesRtoRtheRmovementRabilitiesRofRindividualsRamongRlocalRhabitatRpatchesRhaveR theRpotentialRtoReitherRexacerbateRorRameliorateRtheReffectsRofRhabitatRfragmentationRonR populationskRR RInRadditionhRitRhasRbeenRrecentlyRsuggestedRthatRspatialRheterogeneityRdekgkhRinR resourceRavailabilityeRwithinRhabitatRpatchesRcanRimposeRsignificantRselectiveRpressuresRonR routineRmovementsRwhichRcanRaffectRoverallRdispersalRabilityRandRmorphologyRdBaguetteR andRVanDyckRommtekR the most part occur within a habitat patch, for example mateilocationRandRforagingR activitiesRdVanRDyckRandRBaguetteRommrekRLocalRconditionsRwithinRhabitatRpatchesRmayR affectRmobilityhRpotentiallyRinRcontrastingRdirectionsRtoRtheReffectsRofRsurroundingR landscapeRstructureRdVanRDyckRandRBaguetteRommryRTurlureRetRalkRomnmekRWhileRtheReffectsR ofRhabitatRfragmentationRonRmovementRandRdispersalRabilityRhaveRbeenRrelativelyRwellR studiedRdekgkhRThomasRetRalkRnvvuyRHillRetRalkRnvvvahRbyRNorbergRandRLeimarRommoyRMerckxR 9R etRalkRommpyRVandewoestijneRandRVanRDyckRomnnehRweRknowRcomparativelyRveryRlittleR regardingRtheRinfluenceRofRwithinipatchRhabitatRheterogeneityRdekgkhRspatialRvariationRinR resourceRavailabilityeRonRtheRevolutionRofRmovementRabilityRdTurlureRetRalkRomnmyRYeRetRalkR omnpekRThereforehRfutureRstudiesRwhichRexamineRmovementRabilityRinRrelationRtoRbothR landscapeRstructureRandRlocalRhabitatRcharacteristicsRwillRprovideRnewRinsightsRintoRtheR factorsRshapingRdispersalRpropensityRofRpopulationskR

1.7RButterflieswRimportantRflagshipRspeciesRandRaRmodelR

systemRforRtheRstudyRofRhabitatRfragmentationR Butterflies (OrderwRLepidopteraeRareRoneRofRtheRmostRextensivelyRstudiedRandRwelli describedRgroupsRofRinvertebratesRdGastonRnvvnekRDueRtoRtheirRstrictRecologicalR requirementshRbutterfliesRareRrecognizedRasRpotentiallyRvaluableRindicatorsRofRecosystemR healthRdMcGeochRommteRandRclimateRchangeRdHellmanRommoyRParmesanRommpekRInRrecentR yearsRmanyRspeciesRhaveRexperiencedRmarkedRdeclinesRinRabundancehRandRinRsomeR regionshRsuchRasRtheRUkKkhRtheseRlossesRhaveRbeenRwellidocumentedRandRtheRcausesRwelli understoodkRConsequentlyhRbutterfliesRareRoftenRtheRfocusRofRconservationReffortsRasR protectionRofRtheirRhabitatRisRseenRasRaRproxyRforRmoreRgeneralRspeciesRconservationRdekgkhR FoxRetRalkRomnnekRInRadditionhRinRcomparisonRtoRotherRthreatenedRinvertebrateRtaxahR butterfliesRareRperceivedRasRcharismaticRandRaestheticallyRappealingRanimalshRandRhaveR thusRbecomeRanRimportantRflagshipRforRraisingRawarenessRandRdevelopingRnewR methodologiesRforRinvertebrateRspeciesRconservationRdSamwaysRnvvqyRNewRnvvtekR 10R ButterfliesRhaveRlongRbeenRusedRasRaRmodelRorganismRforRtheRstudyRofRpopulationR biologyRdGilpinRandRHanskiRnvvnyRHanskiRnvvvyREhrlichRandRHanskiRommqekRTheirRutilityRinR thisRfieldRstemsRfromRtheRrelativeReaseRwithRwhichRtheyRcanRbeRobservedRandRmanipulatedR inRbothRlaboratoryRandRfieldRexperimentsRandRtheirRwellRstudiedRlifeihistoriesRandRgeneralR ecologyRdBoggsRetRalkRommpekRAsRwellhRspeciesRwithinRthisRtaxonomicRgroupRareR ecologicallyRdiverseRandRexhibitRpopulationRs

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