University of Alberta · ABSTRACT The Mormon metalmark (Apodemia mormo) butterfly is widely...

UniversityofAlberta

Beyondthehostplant:Multi‐scalehabitatmodelsforanorthernperipheralpopulationofthebutterfly,Apodemiamormo(Lepidoptera:Riodinidae)

AshleyAnneWick

AthesissubmittedtotheFacultyofGraduateStudiesandResearchinpartialfulfillmentoftherequirementsforthedegreeof

MasterofScience

ConservationBiology

DepartmentofRenewableResources

©AshleyAnneWickSpring2013

Edmonton,Alberta

PermissionisherebygrantedtotheUniversityofAlbertaLibrariestoreproducesinglecopiesofthisthesisandtolendorsellsuchcopiesforprivate,scholarlyorscientificresearchpurposesonly.Wherethethesisisconvertedto,orotherwisemadeavailableindigitalform,theUniversityofAlbertawilladvisepotentialusersofthethesisof

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ABSTRACT

TheMormonmetalmark(Apodemiamormo)butterflyiswidelydistributed

throughoutwesternNorthAmerica.Itislistedasthreatened,however,in

Saskatchewan,Canadabecauseofasmallpopulationsizewithinarestrictedhabitat.

Tomosteffectivelymanageforthisspecies,landmanagersandconservationists

requireamorethoroughunderstandingofitsecologyandhabitat.Icompletedthree

studiesthatadvancetheunderstandingofthisthreatenedbutterfly.First,ina

microhabitatstudyIshowedthathostplantabundance,soilchemistry,and

microtopographyareimportantindeterminingwhetherabutterflyhabitatis

occupied.Second,Idevelopedandevaluatedthepredictionaccuracyofspecies

distributionmodelsusingtwomodelingtechniques,theresultsofwhichincreased

theknownA.mormocoloniesinSaskatchewanfrom37to88.Finally,Idocument

theovipositionofA.mormoinCanada,showingthatnorthernperipheral

populationsofthisbutterflyexhibitreproductivestrategiesdivergentfromthosein

itscentralrange.

Keywords:Apodemiamormo,Eriogonumpauciflorum,speciesdistribution

modeling,microhabitat,GrasslandsNationalPark,Saskatchewan,Conservation

ACKNOWLEDGEMENTS

Thisprojectwouldnothavebeenpossiblewithoutthekindnessofthepeopleofthe

villageofValMarie,SaskatchewanandthestaffatGrasslandsNationalPark.In

particular,HeatherSauderwasanoutstandingandintuitivefieldassistant,andan

excellentnaturalist.JohanneJanelleisatalentedwildlifephotographerwhohelped

documentnewbehaviorsoftheMormonmetalmark,PatFargeyofferedvaluable

ecologicalknowledgeofthelandscape,CarolandJimprovidedmewithalovely

homeinthefield.

ManypeopleintheDepartmentofRenewableResourcesattheUniversityofAlberta

offeredmeencouragement,supportandadvice.Dr.NadirErbilginhasbeena

constantsourceofpersonalandacademicencouragementasmyadvisor.Asidefrom

theinvaluableadvicefrommyadvisorycommittee,Drs.JohnSpence,ShelleyPruss,

andJensRoland,Ireceivedcounselingfromotherprofessorsandfellowgraduate

students.InparticularI’dliketothankmyfellowgraduatestudentsJenKlutschand

CrisiaTabacaruforhoursofguidanceandencouragement.Additionally,thankyou

toDevinGoodsman,DavidRoberts,KristaFink,SeanCoogan,JillianMartin,Kelsey

Jansen,Dr.EveylnMerrill,Dr.JamiePinzon,andDr.J.C.Cahillaswellastheentire

ErbilginandSpencelabs.

ThankstoParksCanada,theUniversityofAlberta,andtheFishandWildlifeFundof

Saskatchewanfortheirfundingandsupport.

Andfinally,mygratitudegoesouttothosewhohavesupportedandencouragedme

formanyyears,inparticularmyparents(ChristineBroeg,JeffBroeg,JeffWick,and

MarthaWick),grandparents,andbrothers(KyleandMorgan).Theyhavehelped

cultivateandsupportmyloveofnature.Aspecialthankstomylong‐timementor,

Dr.KeithSummervilleDrakeUniversity.

TABLEOFCONTENTS

CHAPTERI.TheecologyandnaturalhistoryoftheMormonmetalmark

Introduction 1

NaturalhistoryofA.mormo 1

Studysites 4

PreviousstudiesonApodemiamormo 5

Thesisoverview 7

LiteratureCited 9

CHAPTERII.Speciesdistributionmodelsrevealnewcoloniesofathreatened

butterfly,Apodemiamormo(Lepidoptera:Riodinidae)

Introduction 13

Materials&methods 14

Studyspecies:Apodemiamormo 14

Studyarea 15

Butterflypresence 16

Predictorvariables 16

Models 17

Results 18

Discussion 20

Environmentalvariablesthatpredictbutterflypresence 21

Groundvalidation&modelcomparison 21

GrazingandA.mormo 22

Implicationsfortheconservation&managementofA.mormo 23

Acknowledgements 25

Literaturecited 26

Tables&figures 32

CHAPTERIII.Beyondthehostplant:Additionalfactorsexplaining

microhabitatassociationsinanorthernperipheralpopulationofApodemia

mormo(Riodinidae)

Introduction 41

Materials&methods 42

Studysites 42

Apodemiamormoandhostplant 43

Studydesign 44

Dataanalysis 45

Results 45

Discussion 46

Acknowledgements 50

Literaturecited 51

Tables&figures 58

CHAPTERIV.FirstObservationsofMormonMetalmark(Apodemiamormo)

OvipositionBehaviorinCanada

Introduction 62

Observations 64

Discussion 64

Acknowledgements 66

Literaturecited 67

Tables&figures 70

CHAPTERV.

GeneralDiscussion:Summaryofnewinformationaboutthehabitatand

ecologyofApodemiamormo

Introduction 71

ConservationstatusofApodemiamormo 72

Grazing 73

Exoticspecies 74

Futuredirections 75

Literaturecited 77

APPENDIXI.

MarkreleaserecaptureoftheMormonmetalmark 80

LISTOFTABLES

Table2‐1.Predictorvariablesusedtobuildtheresourceselectionfunctionand

RandomForestsmodelspredictingnewcoloniesofApodemiamormoinsouthern

Saskatchewan.Candidatevariablesweretestedforassumptionspriortothefinal

models.

..................................................................................33

Table2‐2.GeneralizedlinearmodelcoefficientsofApodemiamormooccurrencesfor

thevariablesusingaresourceselectionfunction.P‐valuesillustratesignificancefor

eachvariable.Residualdevianceandnullshowthatdeviancedecreasedfromthe

nullmodel.

..................................................................................34

Table3‐1.Summarystatisticsaverages(withstandarderrorofthemean)ofhabitat

predictorsat102microhabitatplotsrepresentingoccupiedorunoccupiedsitesby

ApodemiamormoinsouthernSaskatchewan(Canada).

..................................................................................58

Table3‐2.Plantsurveyresultsareshownwiththescientificandcommonnamesof

theplantsaswellastheaverage%coverforeachspecies,dividedbyoccupiedand

unoccupiedsites.

..................................................................................59

Table3‐3.Coefficientsoflineardiscriminants(Scores)includedinthemodel.The

relationshipexplainswhethereachvariablewaspositivelyornegativelyassociated

withbutterflypresenceinhostplanthabitat.

..................................................................................60

TableAppendix‐1.MarkreleaserecaptureresultsfromAugust2011.G6‐BH1areplots

establishedinGrasslandsNationalPark,VM10‐VM14representplotsestablishedinthe

ValMarieCommunityPasture.Atotalof885butterflieswerecaughtwith142recaptures.

..................................................................................80

TableAppendix‐2.AnexplanationofthetermsshowninTable6‐1.Allresultswerebased

onanalysesfromtheProgramMARK(White2012;www.phidot.org).

..................................................................................81

LISTOFFIGURES

Figure2‐1.Twostudysiteswereusedintheseinvestigations,theValMarie

CommunityPastureandGrasslandsNationalPark’sEastandWestBlocks,locatedin

southernSaskatchewan,Canada.

..................................................................................35

Figure2‐2.VariableimportanceplotgeneratedbytheRandomForestsalgorithm.

Theplotshowsthevariablesimportancemeasuredasincreasednodepurity

(IncNodePurity)andtheincreaseofmeansquareerror(%IncMSE),which

representsthedeteriorationofthepredictiveabilityofthemodelwheneach

predictionisreplacedinturnbyrandomnoise.Highervaluesof%IncMSEand

IncNodePurityindicateahighervariableimportance.Thefullvariablenamesare

showninTable2‐1.

..................................................................................36

Figure2‐3.ThemapsshowtheEastBlockofGNP(a,b)theVMCP(c,d),andthe

WestBlockofGNP(e,f).TheRSFisshownontheright(b,d,f)andRandomForests

ontheleft(a,c,e).Themodelwasonlytargetedforareasthathavepreviouslybeen

delineatedasbadlandshabitatpriortothisstudy.

..................................................................................37

Figure2‐4.SitespredictedtohaveApodemiamormopresencesbyeachmodel

(probabilityof0.60orhigher)weresearchedinGrasslandsNationalParkandthe

neighboringValMarieCommunityPasture.Thefigureshowstheobserved(number

ofsitesinwhichApodemiamormowasfound)versusexpected(numberofsitesthat

werepredictedtocontainA.mormo).

..................................................................................38

Figure2‐5.SitespredictedtonothaveApodemiamormohabitatbytheRandom

ForestsmodelforboththeValMarieCommunityPasture(VMCP)andGrasslands

NationalPark(GNP).Thefigureshowstheobserved(numberofsitesinwhichthe

modelwascorrectinpredictingtheabsenceofA.mormo)versusexpected(number

ofsitesthatwerepredictedtonotcontainA.mormo).

..................................................................................39

Figure2‐6.Modelresultsseparatedbasedonwhetherornotanareawasungrazed,

grazedbybison,orgrazedbycattle.TheValMarieCommunityPasture(VMCP)is

grazedbycattle,theEastBlockofGrasslandsNationalPark(GNP)isnotgrazed,and

theWestBlockofGNPisgrazedbybison.Thefigureshowstheobserved(numberof

sitesinwhichApodemiamormowasfound)versusexpected(numberofsitesthat

werepredictedtocontainA.mormo).

..................................................................................40

Figure3‐1.Lineardiscriminantanalysisshowsthedistribution“discriminants”of

thetwodistributions,thefirstofwhichis“unoccupied”habitatandthesecond,

whichis“occupied”habitat.

..................................................................................61

Figure4‐1.FemaleMormonMetalmark,Apodemiamormo,ovipositingincracksin

thesoilinTimmonscoulee,GrasslandsNationalParkofCanada(WestBlock),near

ValMarie,SaskatchewanonAugust21,2011.Photo:JohaneJanelle.

..................................................................................70

CHAPTERI.

TheecologyandnaturalhistoryoftheMormonmetalmark

Introduction

Butterflies(Lepidoptera)aretaxonomicallywellknown,relativelyeasytofind,have

shortlifecyclesthatareeasytoworkwith,and,arecharismatic.Itisthereforeno

surprisethat,formodern‐dayresearch,butterfliesaretheinsectgroupmost

frequentlymodeledfordevelopmentandtestingofecologicaltheory(Thomas

2005).

Lifehistoryandbasicecologyofaspeciescansometimesbeoverlooked,especially

inthecaseofoutlier,orperipheral,populations.InthisintroductorychapterIfirst

presentanoverviewofthebiologyandecologyofmystudysubject,theMormon

metalmark(Lepidoptera:RiodinidaeApodemiamormoFelder&Felder1859),a

butterflylistedasthreatenedinthefarreachesofitsnorthernrangein

Saskatchewan,Canada(COSEWIC2003).Second,Idescribethestudysites,

GrasslandsNationalPark(GNP)andtheneighboringValMarieCommunityPasture

(VMCP),bothlocatedinextremesouthwesternSaskatchewan.Next,Isummarize

previousstudiesonA.mormoandrelatedspecies.Lastly,Ipresentmyobjectivesfor

thefollowingchaptersofthisthesisanddescribehowtheyrelatetothecurrent

conservationneedsofthespecies.

NaturalhistoryofA.mormo

Apodemiamormoisasmall‐bodied(wingspan25‐32mm)butterflyofthefamily

Riodinidae(Layberryetal.1998;COSEWIC2003).Themetalmarksbelongtothe

superfamilyPapilionidea,orthetruebutterflies.Riodinids,orthemetalmarks,are

so‐namedbecauseofmetallicmarksonthewingsofmanyspecies.Thename

“Mormonmetalmark”occasionallyelicitscuriosityastoitsetymology.Infact,Felder

&Felderweremistakenwhentheygavethebutterflyits“Mormon”moniker.The

Felderswerestay‐at‐hometaxonomistswhowouldreceivespecimensfromothers

onfieldexpeditions.WhentheygavethespeciesnametothisApodemiathey

thoughtthatthetypespecimenshadcomefromUtah,wheninfacttheywere

mistakenandthespecimenswerefromNevada.Intheend,thisbutterflywasleft

withaname,albeitinaccurate,thatenduresinnaturalists’memories.

Worldwidethereareover1400speciesofmetalmarks,only24ofwhicharefound

inNorthAmerica(Scott1986;Layberryetal.1998;Pelham2008).Accordingto

Opler&Powell(1962),A.mormowasoriginally5allopatricspecies,butthesehave

allbeenreducedtosubspeciesintaxonomicrevisions.Apodemiamormoisthemost

widespreadriodinidinNorthAmerica,witharangethatextendsfromnorthwest

MexicothroughoutthewesternUnitedStatesandupintosouthernCanada(Scott

1986;Layberryetal.1998).ItistheonlyriodinidfoundinCanada(Layberryetal.

1998)withadistributionthatbecomesmoreisolatedanddisjunctinitsnorthern

range(COSEWIC2003).ItispolytypicwithrespecttothearrayofEriogonumhost

speciesituses,andinthediversityofadultsizeandcoloracrossitsrange(Opler&

Powell1962;Powell1975).

TheCanadianpopulationsrepresentlessthan1%ofA.mormo’stotalpopulation

andglobalrange(Canningsetal.1998).TwoA.mormopopulationshavebeen

documentedinCanada:anendangeredpopulationintheSimilkameenRiverValley

ofBritishColumbiaandathreatenedpopulationinthemixedgrassprairiesof

southernSaskatchewan(COSEWIC2003).Thetwolistingsweregivenoriginallyby

theCommitteeontheStatusofEndangeredWildlifeinCanadaundertheFederal

SpeciesAtRiskAct(SARA;COSEWIC2003).TheprairiepopulationinSaskatchewan

ispartofanorthernoutlierpopulationthatcontinuesintotheUnitedStatesalong

theMilkandMissouriRiversandtheirtributaries,butisdisjunct(orspatially

separate)fromthemainpartofthedistributioninthesouthwesternUSA(Opler

1999;Prussetal.2008).ThelistinginSaskatchewanisbasedontheextreme

fluctuationsofpopulationsfromyeartoyearanditsoccurrenceinahighly

restrictedhabitat(COSEWIC2003;Prussetal.2008).Habitatlossand

fragmentation,exoticspecies,pollutionfromagrochemicals,andclimatechange

couldposeadditionalthreatstothealreadyrarebutterflyinSaskatchewan(Prusset

al.2008).Apodemiamormo(includingsubspecies)remainstheonlymemberofits

familyintheworldtobelisted,althoughthismaybeattributedtoalackof

informationaboutmanyriodinidsthatresideinthehyper‐biodiverseandloosely

regulatedneotropicalforests.

ApodemiamormoutilizesvariousspeciesofEriogonum,abuckwheatofthefamily

Polygonaceae,asitslarvalfoodplantthroughoutitsrange.TheSaskatchewan

populationusesthebranchedumbrellaplant(Polygonaceae:Eriogonum

pauciflorum)asitshostplantandnectarsource(Prussetal.2008).AlthoughE.

pauciflorumisabundantinandaroundGNP,itappearsthatA.mormodoesnot

occupymanyoftheseeminglysuitablehostplanthabitatpatches.Itispossiblethat

theremaybeotherbioticand/orabioticvariablesaffectingwhetherornotagiven

patchofhabitattobesuitableA.mormo.Theadultbutterflyusesrubberrabbitbrush

(Asteraceae:Ericamerianauseosa)asasecondarynectarsourceparticularlyduring

thelatterpartoftheflightseasonwhenEriogonumspp.maybedeteriorating

(Hooper2002).Thebutterflyalsotendstoperchonothersubstratesfoundin

badlandhabitatwhereEriogonumspp.isplentiful,suchasrocksandbaresoil,

creepingjuniper(JuniperushorizontalisMoench),sagebrush(ArtemisiacanaPursh),

wolf‐willow(ElaeagnuscommutateBernh.ExRydb)andbuffaloberryshrubs

(SheperdiaargenteaNutt.;Hooper2002;Wickpers.obs.).

TheSaskatchewanpopulationismarkedlyunivoltinewithastaggeredemergenceof

adultsatthebeginningofAugustthatdiminishestowardstheendofthemonth.

However,dependingonweather,itmayflyfrommid‐JulyintoSeptember

(Hendersonetal.2008;Wickpers.obs.).Itsflightandbreedingaremostly

synchronizedwiththefloweringofE.pauciflorum.Withinitslifespan,whichis

about10days,thebutterflyemerges,mates,andfemalesoviposittheireggs(Pruss

etal.2008;Wickpers.obs.).ThepeakadultpopulationinSaskatchewanisreached

inthesecondandthirdweeksoftheflightperiodinmid‐August,butmayvary

dependingontemperatureanddroughtconditions.AccordingtoHowe(1975),in

otherpartsofitsrangethebutterflyoverwintersasyounglarvaeinthestemsof

Eriogonumspp.orunderlitter.Itiscurrentlyunverifiedatwhatstagethebutterfly

overwintersinSaskatchewan.

Studysites

InthisresearchIattempttoexamineanddefinetheoccupiedhabitatconditionsof

A.mormoinitsnorthernperipheralrangeinSaskatchewan.Iconductedthese

studiesinGrasslandsNationalPark(GNP;49°15’N,107°0’W),whichislocatedin

extremesouthernSaskatchewan,inMayandJuneof2011.Long,cold,drywinters

andshort,hot,humidsummerscharacterizeitssub‐humidclimate.Meandaily

temperaturesforValMarie,Saskatchewanrangefrom‐12.4°CinJanuaryto18.3°C

inJulywithtemperatureextremesinJanuaryof‐49.4CandJuly41.1C

(EnvironmentCanada2012).Thereare170daysbetweenkillingfrosts(Frid&

Wilmhurst2009).

Ofthe52,700hathatcompriseGNP,approximately29,000haaresparsely

vegetatedbadlandhabitat,alsoreferredtoaserodedcommunities,andthe

remainingisclassifiedasuplandandlowlandgrasslands(Michalsky&Ellis1994).

Basedondatafromfieldsurveyssummarizedinthisthesis,Iestimatethatthereare

approximately4017hawithsignificantE.paciflorumcoveragewithinGNP(Wick

unpublisheddata).Theparkisdividedintotwomainareas,theEastBlockandthe

WestBlock,separatedbyapproximately40kmofpastureandfarmland.TheEast

Block,surroundedbytheWoodMountainplateau,includesmuchofRockCreek;the

WestBlocksurroundstheFrenchmanRivervalleysoutheastofthevillageofVal

Marie(SaskatchewanInstituteofPedology1992).TheWestBlockoftheParkis

hometoaherdofplainsbison(Bisonbison)whicharestockedatarateof1bison

per55ha(W.Olsonpers.com.2012),aswellasmanySARAlistedspecies,suchas

thegreatersagegrouse(Centrocercusurophasianus),greatershorthornedlizard

(Phynosomahernandesi),andSprague’spippet(Anthusspragueii).Avarietyof

wildlifeuniquetoCanada,GNPisecologicallyvaluable.Threatstowildlifewithinthe

Parkincludeinvasionbyexoticspeciessuchasyellowsweetclover(Melilotus

officinalis),crestedwheatgrass(Agropyroncristatum),andsmoothbrome(Bromus

inermis)(Prussetal.2003).

BecauseseveralA.mormocolonieshavebeendocumentedthere,IincludedtheVal

MarieCommunityPasture(VMCP;49°41’N,107°92’W),whichislocatedseveral

kilometersnorthwestofGNP.TheVMCPwasoriginallycreatedaspartofthePrairie

FarmRehabilitationAdministrationbuthassincebeenintegratedintotheAgri‐

EnvironmentServicesBranch.Thepastureismadeofup40,649haofvaried

topographymuchlikeGNP,andisgrazedbyatarateof1cowper35hectares(T.

Dyckpers.com.2012).TheVMCPiscommunallygrazedbycattlefromaroundthe

1stofMayuntiltheendofOctoberandpatronsarecharged43centsperdaypercow

plus$20/yearfeeforeachcalf.

PreviousstudiesonApodemiamormo

Apodemiamormoconsistsofacomplexofallopatricallyseparatedpopulationsin

NorthAmerica,with17recognizedsubspecies,15ofwhicharefoundinCalifornia

(Pelham2008;USFWS2001).Onesubspecies,Lange’smetalmark(Apodemiamormo

langei),wasdescribedin1938andisnoteworthybecauseithasbeenclosely

studiedafterbeingoneofthefirstinsectstobelistedasendangeredunderthe

EndangeredSpeciesActoftheUnitedStates(ESA;Arnold&Powell1983;Morse

2009;Johnsonetal.2011).Lange’smetalmarkisendemictotheAntiochDunesof

ContraCostaCounty,California(USFWS2001).In1983,Arnold&Powell(1983)

reporteddensitiesof28.7males/acreand41.8females/acreandfoundthatadults

ofbothsexeslivedaboutoneweek.Therearetwopopulationswithinthedunes

withonlyminimalgeneflowbetweenthem,eventhoughlessthanamileseparates

them.AccordingtoArnold&Powell(1983),onlyonemarkedindividual(amale)

traversedthegapinseveralyearsofstudy.Lange’smetalmarkemploysnakedstem

buckwheat(Eriognoumnudumssp.auriculatum)asahostplant.In2006,thepeak

countofLange’smetalmarkwasjust45(USFWS2001).

The2002designationofA.mormoasendangeredinBritishColumbiahasprompted

furtherresearchinCanada.SurveysinBritishColumbiahaveshownthattheadult

population,oncethoughttobelimitedtoafewhundredindividuals,mayactuallybe

upwardsof2000(S.Desjardinspers.com.2011).Arecentinvestigationfoundthat

theBritishColumbiaandSaskatchewanpopulationsarenotcloselyrelated

genetically,suggestingthattheyhavearrivedasaseparatenortherlycolonization

andhavenotmovedwesttoeast(Prosheketal.2013).Prosheketal.(2013)also

revealedthattheSaskatchewanpopulationismoregeneticallydiversethanthe

BritishColumbiapopulation,andconcludedthattheconservationlistingsas

threatenedandendangeredareaccurate.

AlldocumentedA.mormosightingsinSaskatchewanhaveoccurredinthecurrentor

proposedboundariesofGNPortheVMCP(COSEWIC2003).RonaldHooperwasthe

firsttodocumentA.mormoinSaskatchewanintheKilldeerBadlandsofwhatisnow

theEastBlockofGNPin1974,andmaderepeatedtripstonewlocationsinGNPin

1983andthenagainin2002,inwhichhediscoveredseveralmorecolonies(Hooper

2002).By2002therewere8knowncoloniesofthebutterflyinSaskatchewanbut

Hooperclaimsthatmorehadnotbeendocumentedbecauselepidopteristswerenot

lookingattherighttime,intherightplaces(Hooper2002).Searchesafterthe

specieswaslistedin2002yieldedadditionalcoloniessothat,whenIbeganthis

researchin2011,therewere37knowncoloniesofA.mormoinSaskatchewan.

ThelistingofA.mormoinCanadaledtoadditionalstudiesonthebutterflyinits

northernrange.Forinstance,in2010Petersonetal.recordedthefirstcaterpillar

observationsofthemetalmarkinCanada,inGNP.Thisstudyprovidednovel

informationonthebehaviorandphenologyofthespecies’caterpillarstagesinthe

metalmark’snorthernrange.TheyobservedcaterpillarsinJuneandJulyintheWest

BlockofGNPandfoundthatthecaterpillarsarecrepuscularandfeedforshort

periodsintheeveningsarounddusk,andlongerintheearlypre‐dawnhours.The

authorsdidnotobservethecaterpillarsfeedingwhenthetemperaturewaslower

than8C.Apodemiamormoisasolitaryforager,butonemaysometimesobserve

severalindividualsonthesameplant(Petersonetal.2010).

ApodemiamormohasneverbeenreportedinAlbertaeventhoughthereissuitable

habitatwherethehostplantoccurs.In2007,asurveyconductedforthebutterflyin

suitablehostplanthabitatinAlbertafailedtoyieldanyevidenceofA.mormo

(Anweiler2007).Repeatedsurveysareneededtoconfirmtheabsenceofthe

butterflyinAlberta.

RecentresearchshowsthatA.mormoovipositioninSaskatchewandiffersfromthat

inotherpartsoftherange;thisworkispresentedinChapterIVofthisthesis.This

studyshowedthatfemalesovipositedonsoilandunderrocks,notonthehostplant,

andthatthefemaleslaidtheireggssingly,notinclusters.Wicketal.(2012)

suggestedthattheSaskatchewanpopulationislikelyunderdifferentadaptive

pressuresthathavepotentiallypromptedthischangeinreproductivestrategy.

Thesisoverview

InthisthesisIpresentmyworkonA.mormo,andtheautecologyofthiselusive

butterflyinitsnorthernrange.Combiningtoolsandtheoryfromlandscapeecology,

populationandcommunityecology,andobservationalstudiesofnaturalhistory,I

havedefinedandcharacterizedhabitatforA.mormoatthemicroandlandscape

level.Asidefromtheprimaryresearchgoalsofthisproject,itismyhopethatthese

resultswillhelpParksCanadatobetterprotecthabitatthatiscriticalforalllife

stagesofthebutterfly.MyresultsformetalmarkpopulationsintheAgri‐

EnvironmentServicesBranch(formerlythePFRA)communitypasturespresented

inthisthesishavealreadybeenusedintheinter‐jurisdictionalSouthoftheDivide

Multi‐SpeciesActionPlan(inpress)inwhichspeciesdistributionmodelsforat‐risk

specieswerecombinedtodevelopregionalgoalsforconservationinsouthern

Saskatchewan.ApodemiamormoistheonlyinsectincludedinthisActionPlan,

whichhasbeenusedtoidentifybiodiversityhostspotsforatriskspeciesin

southernSaskatchewan.Additionally,datafrommyresultsisbeingincorporated

intoamulti‐speciesmanagementplanforGNP.

ToidentifynewcoloniesofA.mormo,inChapterIIIconstructedalandscape‐level

habitatsuabilitymodelacrossthestudyareabasedonspatialenvironmental

variables.Iusedtwocontrastingmodelingtechniques,aresourceselectionfunction,

whichisbasedonageneralizedlinearmodel,andRandomForests,whichisan

algorithmicmachine‐learningtechnique.Whiletheresultingmodelspredicted

differentbutterflylocalitieswithinthestudyarea,fieldassessmentsinpotential

habitatsshowedthatbothmodelsperformedmoderatelywellinpredicting

butterflypresencewithindependentsamples(verifiedwithground‐validation).

Predictionsuccesswas76%forRandomForestsand64%fortheResource

SelectionFunction.OverallpredictionsuccesswashigherwhenIremoveddatafrom

theVMCP,assuccesswasonly19%inthehabitatgrazedbycattle.Applicationand

fieldverificationofthesemodelsincreasedtheknownlocationsofA.mormo

coloniesinSaskatchewanfrom37to88.

TodeterminepotentialmicrohabitatrequirementsofA.mormoinChapterIII,I

asked:“Ishostplanthabitatoccupiedbythebutterflyfundamentallydifferentfrom

habitatthatisunoccupied?”Iansweredthequestionbymeasuringhabitatvariables

onsoil,vegetation,andmicrotopographyin102quadrats,roughlyhalfofwhich

wereunoccupiedandtheotherhalfoccupiedbythebutterfly.Theresultsindicated

thatbutterfliesselectedhostplantsgrowinginhabitatwithgreater%hostplant

cover,greater%baregroundcover,asteeperslope,asouth‐westerlyaspect,lower

soilnitrogen,andhighersoilaciditythaninunoccupiedEriogonumcolonies.Thus,

unoccupiedhabitatdiffersdifferentfromhabitatcurrentlyoccupiedbythebutterfly.

Conservationeffortsshouldfocusonthoseareaswherethebutterflyiscurrently

found.

InChapterIV,isaversionofapaperpublishedinTheCanadianFieldNaturalistin

2012.ThispaperprovidesthefirstdocumentationoftheovipositionofA.mormoin

Canadaandshowsthatthisbehaviordivergesfromthatseeninotherpartsofthe

butterfly’srange.

Finally,inChapterV,thethesisdiscussion,Isummarizethemainfindingsofthe

researchIconducted,revisitkeyissuesthatrelatetothenaturalhistoryand

conservationofA.mormoinSaskatchewan,andconcludebypresenting

managementandconservationrecommendations.

Literaturecited

Arnold,R.A.&J.A.Powell.1983.Apodemiamormolangei.In:Arnold,R.A.(Ed.),

EcologicalStudiesofEndangeredButterflies(Lepidoptera:Lycaenidae):Island

Biogeography,PatchDynamics,andtheDesignofHabitatPreserves.Universityof

CaliforniaPublicationsinEntomology,pp.98‐128.

EnvironnentCanada.2012.http://www.climate.weatheroffice.gc.ca

ESRI.2011.ArcGIS:Versionrelease10.EnvironmentalSystemsResearchInstitute,

Redlands,California,USA.

Frid,L.&J.F.Wilmhurst.2009.Decisionanalysistoevaluatecontrolstrategiesfor

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CHAPTERII.

Speciesdistributionmodelsrevealnewcoloniesofathreatenedbutterfly,

Apodemiamormo(Lepidoptera:Riodinidae)

Introduction

Developingmodelstoeffectivelypredicthabitatconditionsrequiredbyrarespecies

isacentralpursuitofconservationbiologyandessentialtoconservationplanning

(Thompson2004;Guisanetal.2006;Turlureetal.2009;Lawsonetal.2012;Oliver

etal.2012).Forrarespecies,andparticularlyperipheralpopulations,

understandingecologicalrequirementsisespeciallyimportantfordefininghabitat

ordistribution.Speciesdistributionmodels(SDMs)supportconservationinitiatives

byrelatingaspecies’distributiontothedistributionofenvironmentalfactorsthat

predicthabitatuse(Guisan&Zimmermann2000;Soberón&Peterson2005;Araujo

&Guisan2006;Oliveretal.2012).Suchtoolsareparticularlyimportantto

maximizeeffectiveuseoflimitedresourcesforhabitatprotectionandconservation

(Cabeza&Moilanen2001;Araujoetal.2002;Wilseyetal.2012).Predictionsfrom

SDMscanassistconservationistsinprioritizingandfacilitatinghabitatprotection

forthreatenedspeciesbyidentifyingwheresuccessismostlikelytooccur(Manelet

al.1999;Jaberg&Guisan2001;Araujoetal.2002;Cabeza&Moilanen2003;Wilson

etal.2011).

PopularityofSDMshasincreasedinrecentyearsinpartbecauseofthedevelopment

ofavarietyofmappingandstatisticaltechniquesthateffectivelycorrelatespecies

distributionstoenvironmentalcharacteristics(Pulliam2000;Soberón&Peterson

2005;Araujo&Guisan2006).Suchmodelsofferausefulalternativetomonitoring

speciesontheground,whileaidingconservationistsinefficientlyassessingand

projectingimpactsofland‐usechangesorclimatechangeonthedistributionof

organisms,aswellasfacilitatinghabitatprotectionforthreatenedspecies(Kienast

etal.1996;Lischkeetal.1998;Maneletal.1999;Jaberg&Guisan2001;Rebelo&

Jones2010).

NorthernperipheralpopulationsofawidevarietyofspeciesresideinCanada,

includingthatoftheMormonmetalmark(Lepidoptera:Riodinidae,Apodemia

mormoFelder&Felder1859).Therangeofthisbutterflyextendsfromnorthwest

MexicothroughmuchofthewesternUSandintoextremesouthernCanada(Scott

1986;Layberryetal.1998).Apodemiamormoisofparticularecologicalinterest

becauseitsCanadianrangeisdisjunctfromcentralpartsofitsrangeand

populationsofthespeciesresidinginSaskatchewanhavedisparatereproductive

adaptations(Wicketal.2012).

TheCommitteeontheStatusofEndangeredWildlifeinCanada(COSEWIC)recently

assessedthetwopopulationsofA.mormoinCanada.Themixed‐grass“prairie

population”insouthernSaskatchewanwasdesignatedas‘threatened,’andasmall

populationlocatedintheSimilkameenRiverValleyofsouthernBritishColumbia

waslistedas‘endangered’(COSEWIC2003).

InthisstudyIusedSDMstodefinethebutterfly’shabitatbyidentifying

environmentalfactorsassociatedwithhabitatoccupancyatthelandscapelevel.In

particular,Iwasinterestedinwhythebutterflydoesnotoccupymanyareasin

whichitshostplant,EriogonumpauciflorumPursh,isabundant.Ihypothesizedthat

landscape‐levelenvironmentalfeaturesinfluencethesuitabilityofhabitatforA.

mormoandaccountforitsabsenceinsomehostplantpatches.Iusedthese

environmentalfeaturestomodelhabitatoccupancyforthebadlandsofsouthern

Saskatchewanthroughtheuseoftwomodelingapproaches:aresourceselection

function(RSF;Manlyetal.2002);andRandomForests(Breiman2001).Ithen

testedmodelpredictionswithgroundvalidationbysearchingforbutterfliesin

previouslyunsurveyedareasduringthebutterfly’sbriefadultflightperiod.

Materials&methods

Studyspecies:Apodemiamormo

TheprairiepopulationofA.mormoinSaskatchewanisstrictlyunivoltine,with

adultsemerginginearlyAugustandflyingfor2‐3weeks(Hendersonetal.2008;

Petersonetal.2010).Althoughthisbutterflyspeciesusesseveralspeciesof

Eriogonumthroughoutitsrange,theprairiepopulationreliessolelyonE.

pauciflorumasitsobligatorylarvalfoodplantandprimarynectarsource(Scott

1986;Prussetal.2008).InSaskatchewan,A.mormoalsousesrabbitbrush

(EricamerianauseosaPall.ExPursh)asasecondarynectarsource.Thebutterflyand

itshostplantsareassociatedwitherodedhillsidesandembankmentsinthe

badlandsofsouthernSaskatchewan(Prussetal.2008).

Studyarea

ThisworkwasconductedinGrasslandsNationalPark(GNP;Figure2‐1;49°06’N,

107°25’W)islocatedinextremesouthernSaskatchewanandcomprises52,700ha

inthemixedgrassecoregion(Thorpe&Nicolichuk2009).TheParkcontains29,000

haofbadlands,terraintooruggedandunproductiveforhumancultivation

(SaskatchewanInstituteofPedology1992;Prussetal.2008).Muchofthebadlands

arearemainsunaffectedbyhumanactivity,apartfromsomehistoricalcattle

grazing.Thebutterfly’sprimaryhostplant,E.pauciflorum,occursonlyinthe

badlands.Iestimatethatthereareapproximately4017haofE.paucifloruminthe

badlandsofGNPbasedoncalculationsfromvegetationtransectscompletedin2011

(Wickunpublisheddata).TheParkismadeupoftwoparcelsseparatedbyabout40

km:1)theWestBlock,intheFrenchmanRiverValley,whichcontainsaherdof

Bisonbisonand2)theEastBlock,locatedaroundRockCreek(Figure2‐1).TheWest

Blockisgrazedatarateofonebisonper55ha(Olsonpers.com.2012)andtheEast

Blockisungrazedwiththeexceptionofasmallexperimentalareathatdoesnot

includebadlandswhichisgrazedbycattle.

LocatedseveralkilometersnorthwestofGNP,theValMarieCommunityPasture

(VMCP;Figure2‐1;49°41’N,107°92’W)comprises40,629hathatwassetaside

duringtheDustBowlunderthePrairieFarmRehabilitationAdministration,which

wasrecentlychangedtotheAgri‐EnvironmentalServicesBranch.TheVMCPis

grazedatarateofonecowper35ha(T.Dyckpers.com.2012).

AllknownA.mormocoloniesinSaskatchewanarelocatedwithinGNPandthe

Butterflypresence

MultipleA.mormo surveysconducted inGNPandtheVMCPfrom1974until2011

provided301geo‐referenced locationswhere thebutterflyhadbeendocumented.

Usinga222mbuffer radius todefinecolonies theserecordssuggesta totalof37

known colonies. The 222 m buffer was calculated by walking the perimeters of

knowncoloniesandaveragingthesizeofthesecolonies(S.Prusspers.com.2012).

Theseknownbutterflylocationsweretreatedas“used”pointsduringthemodeling

described below. Absence datawere not systematically documented prior to this

studyso Igeneratedasetofunused,oravailablehabitat locationswherethehost

plant occurred but A. mormo presence was unknown (Manly et al. 2002). I

generated this list of available habitat areas as a random sample of landscape

locationsbyinGeographicInformationSystem(GIS;Boyceetal.2002;ArcMap10;

ESRI2011).Thesepresence/availablelocationscomprisedtheresponsevariablefor

subsequentA.mormomodels.

Predictorvariables

Allvariablesusedtomodelandpredictbutterflyhabitat(forafulllistseeTable2‐1)

weregeospatialandavailableinGIS(ArcMap10;ESRI2011).Iemployeda20x20

m cell size to characterize A.mormohabitat in the study area. This resolution is

compatiblewiththebiologyoftherathersessilebutterflyandallpredictorvariables

wereavailableat thisscale (orsmaller).Becauseof theaprioriknowledge thatA.

mormo’s host plant is restricted to badlands, I also restricted the study area to

badlandshabitat.Otherhabitats,suchasrivervalleybottomsorhilltopprairies,do

not supportpopulationsof thehostplant, and thuswere considered tobematrix

habitat.

I derived six variables as environmental predictors (Table 2‐1) from a digital

elevationmodel (DEM) of the study area, and developed a normalized difference

vegetationindex(NDVI)inGISfromSPOT‐5(2007)satelliteimagery.TheNDVIisa

measureof ‘greenness’, ordensityofplantgrowth, inagivenareausingremotely

senseddata.IusedacorrelationmatrixwithintheProgramR(RDevelopmentCore

Team2009) todeterminewhetherpredictorvariableswerehighlycorrelated(r>

0.70;seeManlyetal.2002)forfurtheranalysesIidentifiedthemoreparsimonious

model in relation to Akaike’s Information Criterion (AIC) which increases when

including a variable adds more variance than it explains (Burnham & Anderson

2002;Manlyetal.2002).TopographicRuggednessIndex(TRI)wasremovedfrom

further analyses because it was correlated with slope (SLOPE). Aspect (ASPECT)

was strongly correlated with solar radiation (SOL202) and hillshade (HLSHD), a

measurementofshadedrelief,soIdidnot includeASPECTinthemodelingefforts

(Table 2‐1). HLSHDwas not shown to be significant in the RSF, but excluding it

increasedthemodelsAIC,soIincludedit.

Models

IusedtwomodelstostudytherelationshipbetweenA.mormopresenceandthe

environmentalpredictors.Inthefirstapproach,Iconstructedaresourceselection

function(RSF)usinganexponentialgeneralizedlinearmodelwiththelogitfunction

(Guisan&Zimmermann2000;Manlyetal.2002).ARSFisanyfunctionthatgivesa

resultproportionaltotheprobabilitythataresourceunitwillbeselectedbyan

animal(Manlyetal.2002).Resourceselectionfunctionsarebasedonthepremise

thatindividualsofagivenspeciesbehaviorallyselectparticularresources,andthese

functionshavebeenusedtodeveloppredictivemodelsinconservationplanningfor

avarietyofmammalandavianspecies,includingmountaincaribou,northern

goshawksandnorthernflyingsquirrels(Johnsonetal.2004;Carrolletal.2006;

Hough&Dieter2009).This‘use/availability’designisthemostcommonlyused

methodwhendealingwithdatacollectedinwildlifestudies(Johnsonetal.2006).I

usedthePackagelme4(Batesetal.2011)intheProgramRtoprovideGLM‐based

logisticregressionRSFtoinputintotheGIS.

Asasecondmodelingapproach,IemployedRandomForests,abootstrapped

regressiontreeanalysisinthemachine‐learningfamilyofmodelingtechniques

(Breiman2001;Breiman&Cutler2005).RandomForestshasseveraladvantages

overothermodelingmethods.Forinstance,itcanincludemanycorrelated

variables,itisthoughtnottooverfit,itassumesnonlinearinteractionsamong

predictorvariables,anditdeliversanoutputof‘variableimportance’byranking

predictorvariableswithrespecttohowoftentheycorrectlypredictanoutcomeof

presenceorabsence(Prasadetal.2006;Cutleretal.2007;Roberts&Hamann

2011).BecauseRandomForestsisanalgorithmicmodel,ithasnoassumptionsof

normalityorco‐linearity.Instead,themodellearnstherelationshipamongvariables

startingfromanuniformedstate,andgoesontoelucidatecomplexrelationships

amongvariables(Breiman2001).Iusedthepackage“RandomForests”inthe

ProgramRtobuildthismodel(Liaw&Wiener2002;Breiman&Cutler2005;R

DevelopmentCoreTeam2009).

IconstructedapredictivemapofA.mormohabitatinbothGNPandtheVMCPusing

eachmodel.InAugust2012Ivalidatedmodelperformancebymethodically

searchingforhostplantsandbutterfliesinthesitesthateachmodelpredicted

wouldhavebutterflypresence(>0.60relativeprobabilityofpresence).Additionally,

Isearchedforhostplanthabitat,inlieuofbutterfliesbecausetheflightseasonhad

ended,whereRandomForestspredictedhabitatwouldbeabsent(<0.40relative

probabilityofpresence).IvalidatedpredictionsofabsenceforonlyRandomForests

becauseitperformedbetterthantheRSFinthefirstportionofthestudy.

Results

Thefinalmodelsincludedthefollowingfivevariables:normalizeddifference

vegetationindex(NDVI),compoundtopographicindex(CTI),slope(SLOPE),

hillshade(HLSHD),andareasolarradiation(SOL202).AlthoughHLSHDdidnot

contributesignificantlytotheRSFmodel,excludingitresultedinaless

parsimoniousmodel,soitwasincluded.ThefollowingRSFcoefficientsareshownin

Table2‐2andtheequationwasusedintheGIStopredictA.mormohabitatacross

thestudyarea:

y=exp[4.390‐CTI(0.391)+HLSHD(0.012)‐SOL202(0.006)‐SLOPE(0.043)‐

NDVI(62.4)]

AlthoughRandomForestsrelaxesmanyassumptionsaboutpredictorvariables

requiredbyGLMs,Iusedthesamesetofpredictorvariablesforbothmodelsto

supportthebestpossiblecomparisonbetweenmodels.TheRandomForestsmodel

isbasedonamachine‐learningalgorithmandthereforecannotberepresentedby

anequation,butinsteadgivesoutputintermsoftherelativeimportanceof

variables(Figure2‐2).Outputsshowthatthevariableimportancemeasuredbythe

increasein%meansquareerror(Inc%MSE).GreaterInc%MSEindicateshigher

variableimportance;inotherwords,importancevaluesareassignedinrelationto

thenumberoftimesthatavariableisincludedinaniterationofthemodeland

correctlypredictspresenceinaknownarea.Inthefinalmodel,NDVIwasthemost

importantpredictoraccordingtothevariableimportanceplot(Figure2‐2).In

decreasingorderofimportancethepredictorsincludedintheRandomForests

modelwere:NDVI,SLOPE,SOL202,HLSHD,CTI.

Overall,findingsindicatedthatbutterflypresencewasassociatedwithmoderately

sloped,dryhabitatwithlittlevegetation,lowsolarradiationwithhighershaded

reliefvalues.Iusedeachmodeltopredictprobabilityofbutterflypresencebasedon

environmentalcharacteristicsofeach20x20mcellwithinthestudyarea(Figure2‐

3).TheRSFpredictedamuchhigherproportionofunsuitablehabitatintheVMCP

(Figure2‐4d)thaninGNP(Figure2‐4b,f)incomparisontotheresultsfromRandom

Forests(Figures2‐4a,b,c).

In2012,IsearchedpreviouslyunsurveyedareasintheVMCPandGNPwherethe

modelspredictedA.mormopresenceorabsence.Toselectsearchsites,Idividedthe

20x20pixelsintothefollowingtwocategoriesbasedontheoutputofeachmodel:

1)areaspredictedtohavebutterflies(>0.6probability)and2)areaswithlow

probability(<0.4)ofoccurrence.Withthesebinaryhigh/lowmapsIrandomly

selected200pointsfromtheoutputofeachmodelforeachcategoryinGIS.I

searchedforA.mormointhesesitesduringthepeakflightseasoninAugust2012.It

appearedthattheflightperiodendedearly,dueto2012beingadroughtyear;

therefore,Ididnotincludedatacollectedafter27August,inordertoavoid

detectionerrors.Nobutterflieswereseenafterthisdate,eveninseveralareaswith

knownA.mormocolonies.

Ivisited168sitesthatwerepredictedtohavebutterflies(Figure2‐4),and76sites

thatwerepredictednottohavebutterflies(Figure2‐5).Regardingpredictionof

presence,RSFwasaccurate64%ofthetime,whileRandomForestswascorrect

76%ofthetime(Figure2‐4).SinceRandomForestsconsistentlyoutperformedthe

RSFmodelwithrespecttobutterflyandhostplantpresence,IusedRandomForests

torandomlyselectsitespredictedtohavenobutterflies.RandomForestswas

accurateat92%inpredictionsofnohabitatsuitableforA.mormo(Figure2‐5).

ThemodelresultsapportionedbythetypeofgrazingareshowninFigure2‐6.The

modelswereonlyaccurateincorrectlypredictingA.mormopresence19%(N=31)

ofthetimeintheVMCP,whichisgrazedbycattleatarateofonecowper35ha.The

modelsweremoreaccurate73%(N=106)intheWestBlockofGNP,whichisgrazed

bybisonatarateofonebisonper55ha.ThemodelsweremostaccurateintheEast

BlockofGNP(85%,N=27)wherethereisnograzingbycattleorbisoninanyareas

studiedthatwereoccupiedbyA.mormo.

Discussion

Speciesdistributionmodelshavebeendevelopedformanyspecies,buttheir

applicationinconservationbiologyhasbeencompromisedbythelackof

independentassessmentsofmodelperformance(Greavesetal.2006).Thereare

severalstudiesthathavevalidatedsuchmodelsinthefield.Forexample,Newbold

etal.(2010)usedSDMstomodelthedistributionofbutterflies,mammals,reptiles,

andamphibiansandthentestedtheaccuracyoftheirmodelsusingground

validation.Groundvalidationuncovered15newlocalitiesoftherarebat,

Barbastellabarbastellus,inPortugal,extendingitsknownrangeby100km(Rebelo

&Jones2010).Inthepresentstudy,IdevelopedalternativeSDMstopredictA.

mormopresenceandabsence,andthenusedgroundvalidationtotesttheabilityof

themodels,tocorrectlypredictpresenceandabsenceandtodocumentnew

coloniesofthebutterfly,aswellascomparetwomodelingmethods.

TheSDMsIusedheresuccessfullypredictedsuitableandunsuitablehabitatforA.

mormoandprovideanimprovedunderstandingofwhatconstituteshabitatforthis

species.Giventhesesuccesses,thesemodelscanbeappliedintheconservationofA.

mormoandtheseapproachesmayhavegeneralutilityformodelingpopulationsof

threatenedspeciesthatusepatchyhabitats.

Environmentalvariablesthatpredictbutterflypresence

Priortothisinvestigation,knowledgeofthehabitatoftheA.mormoprairie

populationwaslimitedtothepresenceofitshostplant,E.pauciflorum,inbadlands.

Inthisstudy,Idemonstratedthatfiveenvironmentalvariablescanbeusedto

improvepredictionsofbutterflypresenceandabsencewithintheseconstraints.The

NDVIwasthemostimportantpredictorofbutterflypresenceinbothmodels.NDVI

hadanegativerelationshipwithprobabilityofbutterflyoccurrence;A.mormois

mostlikelytooccurinsparselyvegetatedareaswherehostplantsoccurinthe

badlands.IcouldnotidentifyhostplantatthescaleofSPOTsatelliteimages(10m).

However,IsuspectthatlowNDVIvaluesindicatethepresenceofE.pauciflorumand

E.nauseosabecausetheseplantsarelikelyfavoredbythelowlevelsofcompetition

inhighlyerodedhabitat.Inarelatedstudy(ChapterIIIofthisthesis),I

demonstratedthatA.mormooccursinareaswithlittlevegetationcoveraslongas

itshostplantsareabundant;infact,scarcityofotherplantspeciesmayencourage

butterflypresence.Ingeneral,A.mormohabitatcanbecharacterizedassiteswithE.

pauciflorumthatarenotsteeplysloped,withlittlevegetation,lowsolarradiation,

anddrierwithhighershadedreliefwithhostplantdensities.

Groundvalidation&modelcomparison

ValidationagainstnewdataisessentialtotestifSDMseffectivelypredict

occurrencesofspecies.Ingeneral,modelperformanceisbesttestedagainstnew

presence/absencedatafromthestudyarea,butthathavenotbeenusedtodevelop

themodel(Maneletal.2001;Wintleetal.2005).Validationofmodelusefulnessis

bestachievedbysearchingforthetargetspeciesinareaspredictedbythemodel

(Boyceetal.2002).Therefore,IsearchedforA.mormoduringtheAugust2012

flightperiodatsiteswheretheywerepredictedtobeeitherpresentorabsent.

Groundvalidationconfirmedtheusefulnessofbothmodels,andincreasedthe

numberofknownA.mormocoloniesfrom37to88inasingleflightseason.

InadditiontobetterpredictingnewsitesforA.mormo,RandomForestsmore

accuratelypredictedtheabsenceofbutterflyhabitat.Althoughtheevaluationof

modelsisinextricablyrelatedtotheirintendedpurpose(Araujo&Guisan2006),my

resultscorroboratetheearliersuggestionthatRandomForestsissuitablefor

uncoveringpatternsandrelationshipsinhighlycomplex,non‐linearecologicaldata

(Cutleretal.2007).Thisapproachiswellsuitedforusewiththreatened,

endangered,orperipheralspecies(Marmionetal.2009).

GrazingandA.mormo

WithintheVMCP,inwhichcattleweregrazed,themodelswerecorrectonly19%of

thetime.Modelpredictionsweresubstantiallymoreaccurateinareasungrazed

(85%)orgrazedbybison(73%)intheEastandWestBlocksofGNP,respectively.

TheseresultssuggestthatcattlegrazinghasadifferentialimpactontheabilityofA.

mormotouseavailablehostplantpatches,oratleastontheabilityofeither

modelingapproachtopredictbutterflypresenceinareasgrazedbycattle.However,

furtherworkisrequiredtoascertainifgrazingtypes(cattleorbison)differentially

affectA.mormo,butalsothestockingrates,asthesearehigherintheVMCP.

ManyofthesitesintheVMCPthatwerepredictedtohavebutterflieshadabundant

hostplants,butnobutterflies,evenduringthepeakoftheflightseason.Neither

cattlenorbisonappearedtotargetE.pauciflorumasafoodsource.Recentresearch

demonstratedthatovipositingfemalesofA.mormoplaceeggsdirectlyonsoilor

rocks,notonthehostplant,aswaspreviouslythought(Wicketal.2012).Thereis

limitedpeer‐revieweddatatocomparelifehistoryandtheecologicaleffectsof

managedorunmanagedgrazingbothin(Fuhlendorfetal.2010).

However,theeffectthatlivestockgrazingmayhaveonbutterflypopulationsis

complexandthenatureoftherelationshipdependsonmanyfactors,suchas

butterflylifehistorytraits,plantcommunitysuccession,grazingregimes,and

invasiveplantdynamics(Swengel2001;Vogeletal.2007;Prestonetal.2012).

Furtherwork,suchasacomparativestudyontheecosysteminteractionsofcattle

andbisoninbadlandshabitatontheSaskatchewanpopulationofA.mormowould

likelyclarifytheseissues.

Implicationsfortheconservation&managementofA.mormo

Resourcelimitationisapressingproblemforconservationandlandmanagement

planning.Thisstudyresultedinanadditional106observationsofindividualA.

mormolocalitiesinGNPandVMCP,simplybyusingwidelyavailablespatialdatato

predictbutterflydistributions.Whilethenumberofknowncoloniesin

Saskatchewanincreasedby237%(i.e.from37to88colonies),thisstudyalso

confirmedtheobservationthatA.mormoisnotfoundinmanyareasofhostplant

habitat(Prussetal.2008).

TheresultsIhavepresentedherehavesignificantimplicationsfortheconservation

ofA.mormoinitsnorthernperipheralrange.First,itappearsthatnewcoloniesof

thespeciescanbepredictedwithremotelysenseddata.ThehabitatofA.mormo

wasreasonablywellcharacterizedusingwidelyavailablepredictorenvironmental

variablesderivedfromadigitalelevationmodel(DEM)andsatelliteimagery(SPOT

2007).Second,usingthesameenvironmentalpredictors,thepossiblepresenceofA.

mormomaybeinvestigatedinotherareasofsouthernSaskatchewan.Themodel

couldalsobeappliedtosearchareasofAlbertawithsimilarbadlandsbutwhereno

A.mormocolonieshavebeenpreviouslyuncovered(Anweiler2007).

ThisstudywasfocusedontheprairiepopulationofA.mormoinSaskatchewan;

however,greaterknowledgeofthispopulationwilllikelycontributetoconservation

effortsacrossthespecies’northernrange,includingpopulationsinBritish

Columbia,Montana,WashingtonandUtah.Ourunderstandingoftheconservation

ofmostinvertebratespeciesoftendependsonanintimateunderstandingofhabitat

associations.ToolssuchasSDMsmayhelpconservationistsandlandmanagers

predictandunderstanddistributions,assistinmakingdecisionsregarding

reintroduction,restoration,andmeasurestoreduceprobabilityoflocalextirpation

(Maneletal.2001;Bartel&Sexton2009).Inthefaceoftheperniciouseffectsof

climatechangeandhabitatalteration,habitatmodelingmethodsmaysavevaluable

timeandenergyindocumentingspecies’distributionsaswellasuncovering

unknownpopulations.

Acknowledgements

ThisprojectwasfundedbyParksCanadaandtheUniversityofAlberta.Grasslands

NationalParkstaff,enthusiasticvolunteers,H.Sauder,K.Illerbrun,C.Tabacaru,J.

Klutsch,A.B.Sturm,andB.Smerekovawereinstrumentalinassistingintheground

validationsearches.ThankstoAgricultureCanada,pasturemanagerT.PlattandT.

DyckforpermitsandaccesstothePFRApastures.ThankstoA.Hamann,D.Roberts

andE.Merrillforstatisticalguidance.

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Tables&Figures

Table2‐1.Predictorvariablesusedtobuildtheresourceselectionfunctionand

RandomForestsmodelspredictingnewcoloniesofApodemiamormoinsouthern

Saskatchewan.Candidatevariablesweretestedforassumptionspriortothefinal

models.

*NA=Notapplicable

Source Process Outcome

Variable

Units Resolution

Digitalelevation

model(DEM)

NA* Elevation Meters 20m

DEM SurfaceAnalyst(Spatial

Analyst,ArcGIS10)

Topographic

Ruggedness

Index(TRI)

Index 20m

Analyst,ArcGIS10)

(SLOPE)

Degrees 20m

Analyst,ArcGIS10)

Aspect

(ASPECT)

Categorial

(N,NE,E,

Analyst,ArcGIS10)

Hillshade

(HLSHD)

NA* 20m

Analyst,ArcGIS10)

AreaSolar

Radiation

(SOLAR202)

NA* 20m

SRCPredictive

vegetation

NA* Vegetation

(SRCE2)

Index 1:20,000

polygon

DEM Rastercalculator

Compound

Topographic

Index(CTI)

Index 20m

SPOT5(2007)

SatelliteImagery

Rastercalculator

Normalized

Difference

Vegetation

Index 10m

Table2‐2.GeneralizedlinearmodelcoefficientsofApodemiamormooccurrencesfor

thevariablesusingaresourceselectionfunction.P‐valuesillustratesignificancefor

eachvariable.AICmodelselectionwasusedtorankmodelsupportandfindthe

mostparsimoniousmodel.Residualdevianceandnulldeviancearealsoshown,

showingthatdeviancedecreasedfromthenullmodel.

ResourceSelectionFunctionmodelcreation

Nulldeviance: 1391.6

Residualdeviance: 1044.8

Variable1 RSFCoefficient RSFP‐value2

Intercept 10.49 0.1077

CTI ‐0.391 1.36e‐08*

HLSHD 0.012 0.1396

SOL202 ‐0.006 0.0074*

SLOPE ‐0.043 0.000122*

NDVI ‐62.372 <2e‐16*

1VariableswereexplainedinTable1.

2*Indicatessignificanceat0.01=alpha

Figure2‐1.ThestudyareconsistingoftheValMarieCommunityPastureand

GrasslandsNationalPark’sEastandWestBlocks,insouthernSaskatchewan,

Canada.

Figure2‐2.VariableimportanceplotgeneratedbytheRandomForestsalgorithm.

Theplotshowsthevariableimportancemeasuredasincreasednodepurity

(IncNodePurity)andtheincreaseofmeansquareerror(%IncMSE),which

representsthedeteriorationofthepredictiveabilityofthemodelwheneach

predictionisreplacedinturnbyrandomnoise.Ahighervalueof%IncMSEindicates

ahighervariableimportance.ThefullvariablenamesareshowninTable1.

Figure2‐3.ProbabilityoffindingApodemiamormoinGrasslandsNationalPark.

Mapsaredividedupbysection,theEastBlock(a,b)theVMCP(c,d),andtheWest

BlockofGNP(e,f).Theresourceselectionfunctionisshownontheright(b,d,f)and

RandomForestsontheleft(a,c,e).Themodelwasonlytargetedforareasthathave

previouslybeendelineatedasbadlandhabitatpriortothisstudy.

Figure2‐4.Numberofobserved(siteswhereApodemiamormowasactuallyfound)

versusexpected(sitespredictedtohaveApodemiamormopresences)byeach

model(probabilityof0.60orhigher)weresearchedinGrasslandsNationalPark

andtheneighboringValMarieCommunityPastureinsouthernSaskatchewan,

Canada.

RSF RandomForests

RSF&RandomForests

Numberofsites

Observed

Expected

Figure2‐5.Numberofobserved(siteswhereApodemiamormowasnotfound)

versusexpected(sitespredictedtonothaveApodemiamormo)bytheRandom

ForestsmodelforboththeValMarieCommunityPasture(VMCP)andGrasslands

NationalPark(GNP)insouthernSaskatchewan,Canada.

VMCP GNP VMCP&GNP

Numberofsites

Location

Observed

Expected

Figure2‐6.Numberofobserved(siteswhereApodemiamormowasactuallyfound)

versusexpected(sitespredictedtohaveApodemiamormopresences)modelresults

separatedbasedonwhetherornotanareawasungrazed,grazedbybison,or

grazedbycattle.TheValMarieCommunityPasture(VMCP)isgrazedbycattle,the

EastBlockofGrasslandsNationalPark(GNP)isnotgrazed,andtheWestBlockof

GNPisgrazedbybison,allsitesarelocatedinsouthernSaskatchewan,Canada.

Cattlegrazed Notgrazed Bisongrazed

VMCP EastBlock(GNP)

WestBlock(GNP)

Numberofsites

Locations

Observed

Expected

CHAPTERIII.

Beyondthehostplant:Additionalfactorsexplainingmicrohabitatassociations

inanorthernperipheralpopulationofApodemiamormo(Riodinidae)

Introduction

Understandingrelationshipsbetweenspeciesandtheirhabitatsisacentralaspect

ofecology(Grinnell1917;Elton1927).Definingandunderstandinghabitatisof

particularsignificanceforspeciesatriskandthoseattheextremeperipheryoftheir

ranges(Fraser2000).Accordingtoevolutionarytheory,individualsshouldbemore

concentratedatthecenteroftheirrangethantheperipheryandthusonecanexpect

gradualdeclinesinabundanceatrangeedges(Brown1984).However,these

featuresaredynamicandsomespeciesarepresentlyshiftingtheirdistributionsto

higherlatitudesinresponsetoclimatewarming(Rootetal.2003;Parmesan&Yohe

2003).Undersuchconditions,northernperipheralpopulationswilllikelybecome

increasinglyimportantforpersistenceofsuchtaxa.

Peripheralpopulationsareofevolutionarysignificanceastheyarethoughttobe

freertoevolveandthusmaybepremiersitesforevolutionarychange,and

ultimately,speciation(Mayr1940).Althoughmanyperipheralpopulationsaremore

geneticallyimpoverishedthancentralpopulations,theyalsomaybemore

geneticallydistinctandthereforeofinteresttostudentsofevolutionandecology

(Noss1994).Environmentalchangessuchasglobalwarmingcanaffectthe

distribution,phenology,abundance,anddiversityofspecies(Crick&Sparks1999;

Roy&Sparks2000;Parmesan&Yohe2003;Rootetal.2003),particularlyat

latitudinalandaltitudinalrangelimitsandthusperipheralpopulationsmaybe

importanttolong‐termpersistenceofmanyspecies(Hunter1991;Fraser2000;

Daviesetal.2006).Furthermore,phenologicalmismatchesbetweeninsectsand

theirhostplants,whichmaybemostcommonatrangelimits,canhavedire

consequencesinperipheralpopulations(Hunter1992;Quiring1993;Peterson

1997).InalandmarkpaperRosenzweig(1991)suggestedthatitiscrucialto

investigatemicrohabitatcharacteristicsofpopulationsatrangeperipheries,where

naturalselectionmaydrivethemtoselectthemostadvantageousmicrohabitats.

Lepidopterahavelongbeenusedasamodelspeciesforstudiesinecologyand

evolutionarybiologyandrecentliteraturesuggeststhatLepidopteradiscriminate

amonghabitatsbasedonenvironmentalvariablesbeyondsimplepresenceoftheir

hostplants(Papaj&Rausher1987;Lastraetal.2006;Ashtonetal.2009).For

example,severalstudiessuggestthatsoilnutrientsaffecthabitatpreferencesof

butterflyspecies(Ehrlich1965;Ravenscroft1994;Prudicetal.2005).Grassland

habitatsthatvaryinexposuretosolarradiationbecauseofslopeandaspectcreatea

varietyofthermalmicroenvironmentsthatdramaticallyaffectlarvalgrowthand

development,especiallyinthecooler,northernrangeofthebaycheckerspot

butterfly(Murphy&Weiss1988).

InthischapterIinvestigatemicrohabitatcharacteristicsforthedisjunctnorthern

peripheralpopulationoftheMormonmetalmark(Apodemiamormo)insouthern

Saskatchewan,Canada.Theconservationstatusofthispopulationislistedas

threatened(COSEWIC2003).Myobjectivewastodefinepossiblebutterfly

microhabitatusebyexaminingsoils,vegetation,andtopographyforhostplant

habitatswherethebutterflyispresent(occupiedhabitat)andthatwhereitisabsent

(unoccupiedhabitat).

Materials&methods

Studysites

GrasslandsNationalPark(GNP;49°15’N,107°09’W)wasestablishedin1984and

islocatedinthemixedgrassprairieregionofsouthernSaskatchewan,Canada.With

long,cold,drywintersandshort,hot,humidsummers,GNPcomprisesuplandand

lowlandgrasslandsthatareinterspersedwithsparselyvegetatedbadlandhabitat.

The52,700hathatcompriseGNPincludeapproximately29,000haofbadlands,

whichareerodedcommunitiescharacterizedbysparsevegetation(Michalsky&

Ellis1994).

GrasslandsNationalParkisapportionedintotwomainareas:theEastBlockandthe

WestBlock.Theseparcelsareseparatedbyapproximately40kmofprivatelyowned

pastureandfarmland.TheEastBlockincludesmuchofRockCreekandis

surroundedbytheWoodMountainplateau;theWestBlocksurroundsthe

FrenchmanRivervalleysoutheastofthevillageofValMarie,SK(Saskatchewan

InstituteofPedology1992).TheWestBlockcontainsaherdofbison(Bisonbison)

thatgrazetheareaatadensityofonebisonper55ha(W.Olsonpers.com.2012).

InadditiontoGNP,IalsostudiedbutterflyhabitatsintheValMarieCommunity

Pasture(VMCP;49°41’N,107°92’W),whichislocatedseveralkilometers

northwestofthepark.TheVMCPwasoriginallycreatedaspartofthePrairieFarm

RehabilitationActbutisnowmanagedbytheAgri‐EnvironmentServicesBranch.It

ismadeupof40,649haandisstockedatadensityofonecowper35haandcattle

grazefromApriluntiltheendofOctober(T.Dyckpers.com.2012).TheVMCP

containsseveralcoloniesofA.mormoaswellaslargeareasofhostplanthabitat

whererepeatedsurveyshavenotuncoveredthepresenceofthebutterfly.

Apodemiamormoandhostplant

Apodemiamormoisasmallbutterflyoftheprincipallyneotropicalfamily

Riodinidae.Themostwide‐rangingriodinidinNorthAmerica,itoccursfromMexico

toCanada,throughoutthewesternUnitedStates.Onlytwopopulationsarefoundin

southernCanadaandthesecomprisetheextremenorthernrangeofthespecies

(Scott1986;Layberryetal.1998):the“prairiepopulation”inSaskatchewanandthe

“mountainpopulation”intheSimilkameenRiverValleyinBritishColumbia

(COSEWIC2003;Prussetal.2008).Thesubjectofthisstudyistheprairie

population,locatedinthenorthernmixedgrassprairieecoregioninSaskatchewan.

WhileA.mormopopulationsinthesouthernpartofitsrangemayhavemultiple

flightperiods,theprairiepopulationisstrictlyunivoltinewithadultsgenerally

emergingatthebeginningofAugustandwaningtowardstheendofthemonth

(Arnold1980;Petersonetal.2010).However,dependingonweather,theflight

periodofA.mormocanrangefrommid‐JulyintoSeptember(Hendersonetal.2008).

Theadultpopulationdensitytypicallypeaksduringmid‐August.

Thebutterflyusesbranchedumbrellaplant(Eriogonumpauciflorum)asitssole

larvalhostplantandprimarynectarsource.Rabbitbrush(Ericamericanauseosa)is

usedasasecondarynectarsource,althoughitisnotalwayspresenttheareawhere

butterflycoloniesoccur.Adultsusebothplants,aswellascreepingjuniper

(Juniperushorizontalis),baresoil,androcks,forperchingandmating(Wickpers.

obs.).Femalesintheprairiepopulationovipositinsmallcrevicesinthesoilandon

rocksnearE.pauciflorum,notdirectlyonthehostplantaspreviouslythoughtfrom

studiesofotherpopulations(Wicketal.2012).

Studydesign

Historically,presenceofA.mormohasbeendocumentedthroughfieldsurveysby

ParksCanada,theUniversityofAlberta,andFishandWildlifestaff.LocationsofE.

pauciflorumpatchesunoccupiedbythebutterflyhavealsobeenrecorded(Parks

Canada,unpublisheddata2011).Usingthisinformation,Iestablished1025mx5m

quadratsinareaswhereE.pauciflorumwaspresentinthebadlandsofGNPand

VMCPin2011.Roughlyhalf(n=50)ofthesequadratswerelocatedinareaswhereA.

mormohadpreviouslybeendocumented;thesearehereafterreferredtoas

“occupiedhabitat”.Theremainingquadrats(n=52)in“unoccupiedhabitat”were

chosenaccordingtoarandomstratifieddesignandlocatedinknownE.pauciflorum

locations.ThesequadratswerevisitedfromMaytoJuneof2011andaseriesof

microhabitatmeasurementsweretakenineachquadrat.

Afterverifyingthathostplantswereindeedpresent,Irecordedtheelevationas

measuredatthecenterofeachquadratandnotedifthehabitatwasgrazed(by

bisoninGNPorcattleinVMCP).Icollectedasoilcorefromthecenterofeach

quadratusinganopen‐facedauger.Thesoilsamplesweredriedandsubsequently

analyzedfortotalnitrogencontent(TKN,mgL;anindicatorofsoilfertilitystatus),

acidity(pH)andsoilelectricalconductivity(EC;ameasurementofsoilsalinity)in

theHighVolumeLabattheUniversityofAlberta.OnsiteIalsomeasuredsoil

penetrabilityusingapentrometer(g/cm;E280DaytonPocketPentrometer).

AtthecenterofeachquadratImeasuredslopeandaspect.Tocharacterizethebiotic

communityIestimated%bareground,%hostplantcoverand%coverofall

vascularplantspeciestothenearest5%.Grassesandsedgesweredifficultto

reliablyidentifytospecies,soIestimatedsedgesandgrassesasatotalgrasscover

tocharacterizetherelationshipsbetweenthisvegetationtypeandthehostplant.

DataAnalysis

InordertousethedatatodiscriminatebetweenoccupiedandunoccupiedhabitatsI

ranalineardiscriminantanalysis(LDA)usingtheRpackageMASS(Venables&

Ripley2002;RDevelopmentCoreTeam2009).Lineardiscriminantanalysisisa

classicparametricmethodofclassificationusedwithacategoricalresponsevariable

(Sherrod2012).AnLDAaimstominimizewithingroupvariancebyexplainingthe

varianceamonggroupsusingasetofpredictorvariablesandmaximizestheratioof

between‐classvariancetowithin‐classvariancebyfindingalineartransformation

ora“discriminantfunction”(Sherrod2012).TheLDAresultsyieldtwo

distributions:oneeachforoccupiedandunoccupiedhabitat.IusedaWelchtwo

samplet‐testtotestwhetherthetwodistributionsstatisticallydifferedfromone

another.

Results

AssummarizedinTable3‐1,averagemeasurementsfortheindependentvariables

differedbetweenunoccupiedandoccupiedhabitat.Soilpenetrability,availablesoil

nitrogen,%baregroundcover,andelevationwerealllowerinoccupiedhabitats,

whileunoccupiedhabitatshadlowersoilsalinityandhighersoilpH.Occupied

patcheswereonsteeperslopesandmoregenerallyhadasouth‐westerlyaspect.

Thevegetationcompositionandstructuresurveys(Table3‐2)weredoneinthe

earlypartofsummer(May/June),andtherefore,asidefromthehostplantand

primarynectarplant(E.nauseosa),themostcommonplantswerespringflowers.

Someplantswereplentifulintheplotssurveyedearlyintheseasonandallbut

absentfromplotsbythelatesummer,andbecauseofthisobviousphenological

effect,theseplantswereexcludedfromfurtheranalyses.However,yellowumbrella

plant,saltbush,pricklypearcactusandsagewereallpresentthroughoutthe

summer.Table3‐3showstheabundancesofplantsthatweresurveyedintheplots.

SinceIwasinterestedinwhethertheoverallcommunitiesdifferedbetween

occupiedandunoccupiedhabitat,IpresenttheresultsoftheLDA(Figure3‐1),

insteadofpresentingresultsforeachindividualvariable.TheLDAsuggestsdistinct

distributionsforoccupiedandunoccupiedhabitats.AWelchTwoSampleT‐test

showedasignificantdifference(df:133.9,T=2.347,P=0.02)betweenthe

distributions,indicatingthatE.pauciflorumpatchesthatareoccupiedorunoccupied

byA.mormodifferbasedonacombinationofthefollowingsevenvariables:%E.

pauciflorumcover,elevation,aspect,availablesoilnitrogen,slope,%bareground

cover,andsoilpH(Table3‐3).

Discussion

PresenceofA.mormoinE.pauciflorumhabitatpatchesreflectsacombinationof

topographic,soilandbioticvariables.Althoughhostplantpresenceisakey

predictorofbutterflypresence,itisinsufficienttofullycharacterizehabitat

occupancybythebutterfly.Apodemiamormowasfounddisproportionatelyin

patchesofE.pauciflorumthathadlowerelevationthanaverage,greaterslope,anda

south‐westerlyaspect,lowersoilnitrogen,higheracidity,andhigher%bareground

cover.

Clearly,theextentoflocalE.pauciflorumcoverincreasestheprobabilityofA.

mormopresence.Itislikelythathighhostplantdensitiesareimportantfor

developinglarvaeastheyundertakeshortdistancemigrationfromonehostplantto

anotherduringtheirlateinstardevelopment,asadultA.mormofemalesdonot

ovipositonhostplants(Petersonetal.2010;Wicketal.2012).Inanotherstudy,

densityofhostplantflowerheadswasthemostimportantfactorindetermining

presenceandabundanceoftheendangeredlargebluebutterfly,Maculineateleius

(Bataryetal.2007).

Otherstudieshaveshownthatmicrohabitatfactorsaffectwhichplantsinsects

chooseastheirhosts.Yamatotoetal.(2007),forexampledemonstratedthatthe

biomassofparticularhostplantspeciespositivelyaffectedtheabundanceof

butterflyspecies.Thisalignswithmyfindings:althoughhostplantpresenceis

insufficienttopredictA.mormopresence,althoughprobabilityofoccupancy

increaseswithabundanceofthehostplant.InastudyofthevulnerableBritish

silverspottedskipperbutterfly(Hesperiacomma)thebutterfly’sidealhabitat

consistedofbrokensouth‐facingterrainwith45%hostplantcoverand40%bare

ground(Thomasetal.1986).Interestingly,thebutterflyhadapparently

disappearedfromsitesafterthesehabitatcharacteristicshadchanged.

MyresultsarealsoconsistentwithfindingsabouttheQuinocheckerspotbutterfly

(Euphydryasedithaquino),forwhichpresencewaspositivelyassociatedwith

microhabitatfeaturessuchashostplantpresence,vegetationstructure,andareas

withhighsolarinsolation(Osborne&Redak2000).Theseauthorsalsofoundthat

highshadewasassociatedwithdelayedemergencefromdiapausewhereaslow

shadewasassociatedwithearlyemergenceandaccelerateddevelopment.

Furthermore,Dobkinetal.(1987)andWeissetal.(1988)bothfoundthat

topographicheterogeneitywithrespecttoslopeexposureinserpentinegrasslands,

contributedtothelongtermpersistenceofpopulationsofE.editha.Thisislikely

becausemicroclimate,whichdependsonmicrotopography,affectsthephenologyof

hostplantsoflarvaeandnectarsources(Weissetal.1988).Shadingand

microclimateeffectsonA.mormocouldbeinvestigatedtoidentifypotential

mechanismsthatexplainhabitatpreferencesdeterminedinthisstudy.

Similarmicrohabitatfeaturesareimportanttohabitatuseinotherspeciesof

conservationconcern.Forexample,astudyonthemicrohabitatselectionoffive‐

linedskinks(Eumecesfasciatus;listedasspecialconcernunderSARA)intheir

northernperipheralpopulationsinCanadafoundthatsitepreferencesweredriven

primarilybythermoregulationandprotectionfrompredators(Quirtetal.2006).

Similartothecurrentstudy,thefindingsofQuirtetal.(2006)increasedthe

understandingofhabitatselectioninthenorthernrangeofanorthernperipheral

population.

Presenceofexoticplantsinfluenceshabitatuseinmanybutterflyspecies,mainly

throughcompetitiveexclusionofhostplants(Proctor&Woodwell1975;Murphy&

Ehrlich1988;Murphy&Weiss1988;Mattonietal.1997;Osborne&Redak2000).

However,theonlyexoticspeciesIdocumentedinE.pauciflorumhabitatwas

Melilotusofficinalis.Althoughtheoccurrenceofthisspeciesinhostplanthabitatwas

verylowatanaverageof0.1%coverinoccupiedhabitatand0.9%coverin

unoccupiedhabitat,itmayhaveinfluencedhabitatselectionbyA.mormo.Infact,in

GNPtherearelargetractsoflandthathavebeendenselyoccupiedbythisexotic

plant,someofthemborderingsmallE.pauciflorumpatches,whichmayinfluenceA.

mormomovementbetweenlargerhabitatpatches.Additionally,dispersalwasnota

parameterthatweexaminedinthisstudy,butcouldlikelyinfluencewhetherornot

thebutterflycanreachagivenE.pauciflorumhabitat.

Conservationimplications

Inthefaceofglobalwarmingandothermajorenvironmentalandlandusechanges,

peripheralpopulationswilllikelybeimportantforthelong‐termpersistenceof

manyspecies(Hunter1991;Fraser2000).Itisvitaltounderstandtheecologyand

habitatrequirementsofthesepopulationsinordertoeffectivelymanagehabitats

forpersistenceofsuchperipheralpopulations.ThedistributionofA.mormointhe

prairiepopulationisrestrictedintwoways:butterfliesonlyoccurinbadland

habitatandinproximityofthelarvalhostplant,E.pauciflorum.Beyondthese

previouslyunderstoodhabitatrestrictions,Ihaveshownthatotherenvironmental

characteristicsinfluencebutterflyoccupancyinE.pauciflorumpatches.Isuggest

thattheprimaryconcernforconservationoftheprairiepopulationofA.mormoisto

preserveandmaintainthequalityofhabitatthatthebutterflycurrentlyoccupies

becauseitappearsthatA.mormoisselectingforasubsetofhostplanthabitat.

Mostimportantly,theresultspresentedhereimproveunderstandingofhabitat

requirementsforA.mormoinitsnorthernrange,andcanbeusedbylandmanagers

toidentifyandconservehabitatforthebutterfly.Thefollowingcriteriashouldbe

helpfulfordefininganddesignatingcriticalhabitatforA.mormo:patcheswithhigh

%ofE.pauciflorumcover,growingonsoilswithhigherpHandlowavailable

nitrogen,onsteepslopesthataresouth‐westtosouthfacingatlowerthanaverage

elevation,inbadlandhabitatwithbaregroundcover,andparticularly,areasthatare

currentlyorhavebeenrecentlyoccupiedbythebutterfly.

Acknowledgements

ThisprojectwasfundedbyParksCanadaandtheUniversityofAlberta.This

researchwouldnothavebeenpossiblewithoutthehelpofmyfaithfulfield

assistant,HeatherSauder,aswellasindispensibleassistancefromthepeopleofVal

Marie,SaskatchewanandGrasslandsNationalPark.

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Tables&Figures

Table3‐1.Summarystatisticsaverages(standarderrorofthemean)ofhabitat

predictorsat102microhabitatplotsrepresentingoccupiedorunoccupiedsitesby

ApodemiamormoinsouthernSaskatchewan,Canada.

Variable Unoccupied Occupied

Penetrabilitya

3.234(0.199)

2.95(0.188)

TKNmgLb 0.099(0.07) 0.089(0.005)

Soilacidity(pH) 5.858(1.49) 5.962(0.190)

Soilsalinity(EC)c 1055.263(169) 1159.043(198.081)

Bareground%cover 50.980(3.0) 51.270(3.101)

Elevation 841.10(5.905) 831.532(6.337)

Aspect 171.62(13.62) 201.04(15.12)

Slope 12.58(1.22) 17.766(1.754)

aPenetrabilityisameasureofthepenetrabilityofthesoil.

bTKNisavailablesoilnitrogen,anindicatorofsoilfertilitystatus.

cSoilelectricalconductivity(EC)isameasurementofsoilsalinity.

Table3‐2.Plantsurveyresultsexpressedastheaverage%coverforeachspeciesin

occupiedandunoccupiedsites,insouthernSaskatchewan,Canada

Latinname CommonName Occupied% Unoccupied%

Eriogonumpauciflorum Branchedumbrellaplant 21.5 16.0

Ericamericanauseosa Rabbitbrush 6.4 5.1

Juniperushorizontalis Creepingjuniper 3.7 2.8

Opuntiapolyacantha PricklyPearCactus 0.46 0.92

Poaceae&Cyperaceae Grassesandsedges 5.4 7.4

Hymenoxysrichardsonii ColoradoRubberweed 0.1 1.1

Eriogonumflavum YellowUmbrellaplant 0.7 0.6

Artemisiasp. Sage(prairieandpasture) 4.5 5.0

Atriplexnuttallii Saltbush 2.3 0.9

Melilotusofficinalis Yellowsweetclover 0.1 0.6

Rosasp. Wildrose 1.6 2.6

Table3‐3.Coefficientsoflineardiscriminants(Scores)includedinthemodel.

Explainingtherelationshipofeachvariablewithbutterflypresenceinhostplant

habitat.

Variable Scores Relationship

Elevation ‐0.0139 ‐

Aspect 0.0042 +

TKN ‐4.1195 ‐

Slope 0.0665 +

%Baregroundcover 0.0248 +

pH 0.2216 +

%Eriogonumcover 0.0627 +

Figure3‐1.Lineardiscriminantanalysisshowsthedistribution“discriminants”of

thetwodistributions,thefirstofwhichis“unoccupied”habitatandthesecond,

whichis“occupied”habitat.

CHAPTERIV.

FirstObservationsofMormonMetalmark(Apodemiamormo)Oviposition

BehaviourinCanada

ThischapterwaspublishedinTheCanadianFieldNaturalist.

Received11November2011,Accepted3June2012.

Wick,A.A.,Jannelle,J.,Pruss,S.&Erbilgin,N.2012.Firstobservationsofovipositionbehaviourof

theMormonmetalmark(Apodemiamormo)inSaskatchewan.TheCanadianFieldNaturalist.126:

34‐37.

Introduction

TheMormonMetalmark,Apodemiamormo(FelderandFelder1859)isabutterflyof

theprimarilyneotropicalfamilyRiodinidae.Itsrangeextendsfromnorthwestern

MexicothroughmuchofthewesternUnitedStates,butitsdistributionbecomes

patchyinthenorthwesternU.S.andsouthernCanada(Scott1986;Layberryetal.

1998).TheCanadianprairiepopulationsoftheMormonMetalmarkcomprisethe

mostnortherlydocumentedextentofthespecies’range.

ThespecieswasfirstobservedinCanadainAugust1974,whenlepidopteristRonald

HooperdocumentedtheMormonMetalmarkinwhatisnowtheeastblockof

GrasslandsNationalParkofCanada(GNP)insouthernSaskatchewan(Hooper

2002*).Surveysweresporadic,butin1983searcheffortsyieldedtwonewcolonies,

inthewestblockofGrasslandsNationalPark.In2002,sixadditionalcolonieswere

discovered(Hooper2002*).Thespecieswasassessedasthreatenedin

Saskatchewan(Hooper2002*),andthePrairiepopulationoftheMormon

MetalmarkwasassessedasthreatenedbytheCommitteeontheStatusof

EndangeredWildlifeinCanada(COSEWIC)(COSEWIC2003*).Withthese

assessments,increasedsearcheffortshaveledtothedocumentationof40colonies

inGNP,aswellasthefederalValMarieCommunityPasturemanagedbyAgriculture

andAgri‐FoodCanada,whichislocatednorthwestofthepark.

ThereisasecondpopulationinCanadalocatedintheSimilkameenRivervalleyin

southernBritishColumbia(theSouthernMountainpopulation);thispopulationwas

assessedasendangeredbyCOSEWIC(COSEWIC2003*).Bothpopulationsareonthe

ListofWildlifeSpeciesatRiskofthefederalSpeciesatRiskAct,thePrairie

populationasthreatenedandtheSouthernMountainpopulationasendangered.

RecentworkonthepopulationgeneticsoftheMormonMetalmarkinthenorthern

partofitsrangehasreinforcedthelistingsofendangeredandthreatenedunderthe

SpeciesatRiskActandhasuncoverednewinformationthattheBritishColumbia

andSaskatchewanpopulationsareonlydistantlyrelated(Prosheketal.2012).This

suggeststhatfurtherresearchshouldinvestigatewhetherthesepopulationsmay

warrantaseparatetaxonomicstatus(Prosheketal.2012).Littleisknownaboutthe

biologyandpopulationdynamicsofthenorthernprairiepopulations,andmuchof

theinformationcurrentlyavailablereferstoobservationsintheU.S.southwest.

MormonMetalmarklarvaeinSaskatchewanareknowntofeedontheBranched

UmbrellaPlant(alsoknownasFew‐floweredBuckwheat)(Eriogonumpauciflorum

Pursh),whichgrowsalmostexclusivelyonerodedorheavyclaysoils,foundon

hillsides,slopes,andembankments(COSEWIC2003*).TheBranchedUmbrellaPlant

iscommonundertheseconditionsinbadlandshabitat,ofwhichthereareroughly

290km2withinthecurrentandproposedboundariesofGrasslandsNationalPark

(Prussetal.2008*).RubberRabbitbrush,Ericamerianauseosa(Pall.exPursh)G.L.

Nesom&Baird,isalsousedbyMormonMetalmarkadults,whichfeedonthenectar

andperchontheplants.In2010,thefirstobservationsofMormonMetalmark

caterpillarsinGrasslandsNationalParkprovidedvaluableinformationaboutthe

earlylifehistoryofthisbutterflyinCanada(Petersonetal.2010).However,there

wasnodocumentedevidenceofovipositionbehavioroftheMormonMetalmarkin

Canada.GiventheimportanceofGrasslandsNationalParkinthisspecies’Canadian

rangeandthelackofbiologicalinformationspecifictonorthernpopulations,further

understandingofthelifehistoryandbehaviourofthisspeciesisvitalforeffective

conservationplanning.

Observations

OnseveraloccasionsinAugust2011,weobservedMormonMetalmarkfemales

ovipositinginTimmonscouleeinthewestblockofGrasslandsNationalPark,near

thetownofValMarie.OnAugust21,betweennoonand5:00P.M.,wefollowed

severalfemalesthatwouldfindanareaofexposedsoilorrockwithinthehost

plant’shabitat.Oneinparticularbegancurlingherabdomenunderneathherand

walkingforwarduntilshelocatedasuitablespotunderarock(Figure1).Allthe

individualsweobservedlaidasingleamber‐colouredeggapproximatelythesizeof

apinhead,eitherincracksinthesoilorundersmallrocks.Intheinstanceswhere

theeggswerelaidinsoilcracks,morethanoneeggmayhavebeenlaid,butwedid

notobservethis.Whiletheselocationswereallnear(<2m)BranchedUmbrella

Plants,wedidnotobserveanyeggsbeinglaidonthelowerleavesofthehostplant

ingroupsof2–4,aspreviouslydescribed(ArnoldandPowell1983;Scott1986;Pyle

2002).Theentireovipositionprocesstookanywherefrom5to30seconds.

Discussion

Evidencethatthisovipositionbehaviourdiffersfromthatwhichhasbeen

documentedinthesouthernportionofthespecies’rangesuggeststhatdifferent

ecologicaladaptationsmaybeatworkatthenorthernperipheryofitsrange.These

adaptationsmaybetheresultofalternatereproductivestrategies.Thephysiologyof

ectothermssuchasbutterfliesandotherarthropodsislargelydependentonoptimal

temperatureranges,andnaturalandexperimentalmanipulationsrevealthatthese

organismsarecapableofphenotypicplasticityinresponsetotemperature

differences(Fischeretal.2003a,2003b,2004;SteigengaandFischer2007;Berger

etal.2008).Specifically,incertainspeciesofbutterfly,femalesraisedincooler

temperaturestendtoproduceasmallernumberoflargereggsthanconspecifics

raisedunderwarmerconditions(Fischeretal.2003b;Geisteretal.2009).

AtypicalMormonMetalmarkeggdepositionnumbersandsitesmayalsoreflectthe

coldertemperaturesfoundinnorthernpartsofitsrange.Instudyingotherbutterfly

species,Bergeretal.(2008)suggestthatbotheggplacementandmaturationare

limitedbytemperature;eggdevelopmenttimesarealsoinfluencedbydirectsolar

radiation(Bryantetal.2002).Eggslaidinopenhabitatdevelopedmorequickly

(Bryantetal.2002),andthoseeggswithanorientationtomorningsunhadhigher

survivorshipinthecoldestyears(Bonebrakeetal.2010).Incoolerclimates,where

developmenttimeislimited,thepositionofaneggcanhaveaprofoundinfluenceon

survivalandmaturation(Bonebrakeetal.2010).

Experimentswithotherbutterflyspeciesalsoindicatesignificanttemperature

effectsoneggandlarvaldevelopmentaswellasonmortalityrates(Fischeretal.

2004;Koda&Nakamura2010).Thus,MormonMetalmarkeggslaidintheground

andcoveredwithsnowmaybesubjecttomoremoderatetemperaturerangesas

wellaslowerratesofdesiccationthaneggslaidabovegroundonthehostplant.

However,atthispoint,itisunclearwhethereggsorearlyinstarcaterpillars

overwinterintheSaskatchewanpopulation.

Otherstudieshaveshownthateggsplacedawayfromthehostplantonalternate

substratesmayalsobenefitfromdecreasedpredation.Forexample,inastudyofthe

neotropicalbuttterflyOleriaonega,significantincreasesinsurvivalwerefound

whentheeggsweretransferredtoalternatesubstrates(De‐Silvaetal.2011).

However,De‐Silvaetal.(2011)statethatthisanti‐predationstrategymust

outweighthecosttothelarvaeoffindingthehostplant.Additionally,theMormon

Metalmarkmaynotsufferfromextremelyhighpredationfromants,asitdoesinthe

southernpartsofitsrange,suchasCalifornia,wheretheendangeredsubspecies

Lange’sMetalmark(Apodemiamormolangei)resides(Johnsonetal.2011*).

Thedifferencesinovipositionbehaviourreportedhereaddimportantbiological

informationforthisspeciesofconservationconcernandmayaffecttheresidence

descriptionundertheSpeciesatRiskActandthewayinwhichcriticalhabitatis

designatedandprotectedunderthatlegislation.Thisreproductivestrategydiverges

fromwhatwecurrentlyknowaboutthespeciesinotherpartsofitsrange.Further

investigationwillelucidatethemechanismsdrivingthisbehavioraldifference

acrossthespecies’range.

Acknowledgements

Manythankstoourfaithfulfull‐timefieldassistant/localguide,HeatherSauder,as

wellastovolunteersfromacrossNorthAmericaandtheeverhelpfulstaffat

GrasslandsNationalPark.Thanksalsotothereviewersfortheirhelpfulcomments.

ThisprojectwasfundedbytheParksCanadaAgencyandtheUniversityofAlberta.

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rearingofLange’sMetalmarkbutterfly,2010‐2011.FinalReporttoNationalFish

andWildlifeFoundation.TheUrbanWildlandsGroup,LosAngeles,California.

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survivalofanendangeredbutterfly,Shijimiaeoidesdivinusbarine(Leech)

(Lepidoptera:Lycaenidae).EntomologicalScience13:29‐34.

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Lepidopterists’Society34:1‐19.

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UniversityofTorontoPress,Toronto,Ontario.280pages.

Peterson,K.,E.Amosa,S.Pruss,andN.Erbilgin.2010.Firstcaterpillarobservations

oftheMormonmetalmark(Apodemiamormo)(Lepidoptera:Riodinidae)butterfly

inGrasslandsNationalPark,Saskatchewan,Canada.BlueJay68:37‐40.

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Sperling.2013.ApodemiamormoinCanada:populationgeneticdatasupportprior

conservationranking.JournalofInsectConservation17:155‐170.

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MormonMetalmark(Apodemiamormo)prairiepopulationinCanada.Speciesat

RiskActRecoveryStrategySeries.ParksCanadaAgency,Ottawa,Ontario.vi+23

pages.

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Washington,Oregon,andsurroundingterritories.SeattleAudubonSociety,Seattle.

420pages.

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Guide.StanfordUniversityPress,PaloAlto,California.581pages.

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relationtotemperatureandmatingstatusinabutterfly.Entomologia

ExperimentalisetApplicata125:195‐203.

Tables&Figures

Figure4‐1.FemaleMormonMetalmark,Apodemiamormo,ovipositingincracksin

thesoilinTimmonscoulee,GrasslandsNationalParkofCanada(westblock),near

ValMarieonAugust21,2011.Photo:JohaneJanelle.

CHAPTERV.

GeneralDiscussion

SummaryofnewinformationaboutthehabitatandecologyofApodemiamormo

Introduction

TheaimofthisthesiswastoimproveunderstandingofthebiologyoftheMormon

metalmark(Lepidoptera:Apodemiamormo),aswellastheecologicalrequirements

ofthisthreatenedspeciesitsnorthernrangeinSaskatchewan,Canada.Ipresented

anup‐to‐datereviewofinformationaboutthebutterflyinbothCanadaandthe

UnitedStatesinChapterI.InChapterIIIcomparedtwolandscape‐levelpredictive

speciesdistributionmodelsforA.mormo.Idevelopedthesemodelswithgeospatial

environmentaldataandrecordsofA.mormooccupancyusingtwocontrasting

modelingtechniques:theGLM‐basedresourceselectionfunction,andRandom

Forests,whichisanonparametricbootstrappedregressiontreeanalysisrootedina

machine‐learningalgorithm.Ithenground‐validatedthesemodelsbyconducting

searchesfornewbutterflylocalitieswithintheknownrange.ThroughtheseeffortsI

increasedthenumberofknowncoloniesinSaskatchewanfrom37to88inonly

threeweeksofsearcheffort.InChapterIIIIinvestigatedthemicrohabitat

characteristicsthatwereassociatedwiththepresence/absenceofthebutterfly.In

thatstudyIdemonstratedthatmicrohabitatattributesarerelatedtoA.mormo

occupancy.Morespecifically,butterfliesselectedhabitatwithhigher%hostplant

cover,higher%baregroundcover,asteeperslope,asouth‐westerlyaspect,lower

soilnitrogen,andhighersoilpH.

A2012paperpublishedinTheCanadianFieldNaturalistandincludedinthisthesisas

ChapterIVreportsthefirstobservationsoftheovipositionbehaviorofA.mormoin

Canada,presentingnewinformationspecifictothespecies’northernrange(Wicketal.

2012).Thispaperdemonstratedthatthebutterflyovipositsdirectlyonsoilandrocks,not

justonthehostplant,Eriogonumpauciflorum,aswaspreviouslythoughtfromstudiesof

thespecieselsewhere.InthisfinalchapterIfurtherdiscussseveraltopicstouchedonin

thebodyofthethesis,includingthebutterfly’sconservationstatusinCanada,grazingand

itsrelationshiptothebutterfly,andthepotentialinteractionofA.mormowithexotic

species.Finally,Ioutlinepotentialfuturedirectionsforworkonthisspeciesinits

northernperipheralpopulations.

ConservationstatusofApodemiamormo

Butterfliescontinuetobeacrucialconservationtarget,notjustfortheirkeyroleinfood

websoraspollinatorsinmanyterrestrialecosystems,butalsobecausewecanusethem

asmodelspeciestoeffectivelyanswerecologicalandevolutionaryquestions.Asmany

havepointedout,(e.g.,Newetal.1995)habitatsecuritymustcontinuetobethemain

generalstrategyforbutterflyconservation,andthecaseofA.mormoisnoexceptionto

thisprinciple.AsIreflectonthestudiesIconductedaspartofmyMaster’sthesisatthe

UniversityofAlberta,itismyopinionthattheconservationstatusofA.mormoin

Saskatchewanisaccurate,andthatthecontinuedpersistenceofthebutterflywillbe

assuredgivencertaincircumstances.Primarily,landmanagersofGNPmustcontinuetobe

mindfulofthebutterfly’spresenceanditshabitatrequirements.Trails,campgrounds,and

touristfacilitiesshouldnotbeplacedinareaswherethehostplant,Eriogonum

pauciflorum,occurs,evenifthebutterflydoesnotcurrentlyoccupythishabitat.

Fortunately,thehostplant,E.pauciflorum,occursinbadlandshabitat,whichisnotoften

usedforotherpurposessuchasrecreationoragriculture.

PermanentprotectionofGNPiskeytothecontinuedsurvivalofA.mormoinCanada.The

“mountainpopulation”ofA.mormoinBritishColumbiaisnotlocatedinaprotectedarea,

andisthuscorrectlylistedas“endangered.”Unfortunately,thethirdlocationinCanada

thathousesA.mormoisintheValMarieCommunityPasture,partofthePrairieFarm

RehabilitationAdministration,whichhassincebeenintegratedintotheAgri‐Environment

ServicesBranch.Canada’scommunitypastureprogramhasrecentlybeendissolvedand

thefutureofthisland,andthewildlifethatusesit,remainuncertain.

Inadequateknowledgeofthisbutterfly’secologyandhabitatrequirementsinitsnorthern

rangehasimpededcriticalhabitatdesignationfortheprairiepopulationinsouthern

Saskatchewan.Priortothisstudy,habitatrequirementsfortheprairiepopulationwere

unclearandtheRecoveryStrategyforthespeciesstressedtheimportanceofdefiningsuch

habitatforthelong‐termpersistenceofA.mormo’sprairiepopulation(Prussetal.2008).

TheinformationIgatheredaspartofthisthesishasalreadybeencontributedtotheSouth

oftheDividemulti‐speciesactionplanforsouthwestSaskatchewan.Inthefollowing

sectionsIdiscusstwokeyinteractionsthatmayimpactthecontinuedpersistenceofA.

mormoinSaskatchewan:grazingandinvasivespecies.

Grazing

LivestockareubiquitousthroughoutwesternNorthAmerica.Althoughpresenceof

livestockcanbeeffectivelyintegratedwithhabitatmanagementandusedasa

managementtool,evenforbutterflycommunities,applicationstendtobelimited

(Severson1990;Cushman2009).Livestockareheavier,moreabundant,andmore

concentratedthannativemammalsandmoreover,tramplingbylivestockcancompact

soil,reducingitscapacityforwaterinfiltration.OnestudyonSmith’sbluebutterfly

(Euphilotesenoptessmithi)investigatedtheuseofgrazingtohelprestorenative

communitiesthathadbeeninvadedbyEurasianexotics(Cushman2009).WhileCushman

foundthatgrazingreducedtheoverallcoverofexoticgrasses,itincreasedthecoverof

exoticforbs.Further,Eriogonumparvifolium,whichisarelativeofthehostplantofA.

mormo,wasreducedbothinsizeandvolumebygrazing(Cushman2009).Grazing

practices,alongwithindustryandrecreation,aretoblamefortheextensivehabitatlossof

thedunehabitatneededbytheendangeredApodemiamormolangeiinCalifornia(USFWS

2001).

ThestudyareaofmyinvestigationconsistedofGrasslandsNationalPark(GNP),whichis

comprisedofaWestBlockandEastBlock,aswellastheValMarieCommunityPasture

(VMCP).TheVMCPisgrazedbyatarateofonecowper35hectares(T.Dyckpers.com.

2012).TheWestBlockofGNPisgrazedbyonebisonper55ha(W.Olsonpers.com.2012)

andtheEastBlockisungrazed,exceptforasmallexperimentalareathatdoesnotinclude

A.mormohabitat.ThespeciesdistributionmodelsIdevelopedinChapterIIcorrectly

predicted19%,73%and85%ofbutterflypresenceintheVMCP,theWestBlockofGNP

andtheEastBlock,respectively.

IsuspectthatdifferencesinaccuracyofthepredictivemodelsbetweentheVMCPandGNP

maybeinpartexplainedbydifferencesingrazingtype(cattle,bison,orungrazed).My

resultssuggestedthatcattlegrazingisassociatedwithreducedabilityofeithermodeling

approachtocorrectlypredictbutterflypresenceinbadlandshabitatandraisesthe

possibilitythatcattlegrazingoritsintensityisanegativefactorintheconservationofA.

mormo.

ManyofthesitesintheVMCPthatwerepredictedtohavebutterflieshadabundanthost

planthabitat,butnobutterflies,evenduringthepeakoftheflightseason.Thismaybedue

tothefactthattherequirementsofmanyinsectspeciesaredeterminedbylarvae,not

adultsovipositingfemalesofA.mormoplaceeggsdirectlyonsoilorrocks,notonthehost

plant,aswaspreviouslythought(Thomasetal.1992;Wicketal.2012).Onehypothesisis

thatcattlegrazingdiminishesthequalityofeggandlarvalhabitatsbycompactingsoil,and

thereforeovipositingfemalebutterfliesmightavoidhabitatsthathavebeendegradedby

grazing.

ThefederalgovernmentiscurrentlydivestingthePrairieFarmRehabilitation

Administration(PFRA)communitypastures(suchastheVMCP)fromfederaltoprovincial

control,aprocessthatbeganin2012.ThePFRAhasbeenheraldedbysomeasoneof

Canada’sgreatestsuccessstories,asitscreationin1935helpedquelltheeffectoftheDust

Bowl(Monk2012).Nowhereisthelossofthisprogramfeltmorestronglythanin

southernSaskatchewan–62outoftheprogram’s80communitypasturesarelocatedin

Saskatchewan,covering720,340hectares,andtheyhavebeenmanagedbythefederal

governmentforecologicalpurposesfor65years(Herriot2012;Wark2012).Theeffectsof

thedissolutionofthisprogramarealreadybeingseeninlossofjobsinSaskatchewan,

wheresomeschooldistrictsareaquarter‐filledwithchildrenofPFRAemployees(Monk

2012).Itcurrentlyremainsuncertainastowhatwillhappentotheselargetractsofland,

muchofwhichcontainsthreatenedandendangeredspecies,inadditiontoseveral

coloniesofA.mormo.

Exoticspecies

Otherstudieshavefoundthatnon‐hostplants,especiallyEurasianexotics,canindirectly

affectbutterflydistributionthroughcompetitiveexclusionofhostplants(Proctor&

Woodwell1975;Murphy&Ehrlich1988;Murphy&Weiss1988;Mattonietal.1997;

Osborne&Redak2000).InarecentreportonthestatusofA.mormolangei,invasive

plantswerelistedasoneoftheprimarythreats.ThehostplantofA.mormolangei,

Eriogonumparviifolium,likeE.pauciflorum,reliesonnaturaldisturbance.InthecaseofE.

parviifoliumtheconstanterosion,reformationanddestabilizationofthesanddunesin

Californiaisneededforittothrive.TheinvasivespeciesdocumentedintheAntioch

Dunes,whereA.mormolanegiresides,stabilizethesanddunesystem,eliminatingthe

disturbanceregimeonwhichE.parviifoliumrelies.

TheonlyexoticspeciesthatIdocumentedincoexistencewithA.mormo’shostplantinthis

studywasyellowsweetclover(Melilotusofficinalis).However,theoccurrenceofthis

speciesinhostplanthabitatispresentlylowatanaveragecoveroflessthan1%inhabitat

occupiedbythebutterfly.AlthoughthedegreeofinvasionofM.officinalisinE.pauciflorum

habitatwasmarginal,thereisreasonforlandmanagerstobecautiousofthisexotic

species.InGNPtherearesubstantialtractsoflandthathavebeenengulfedbythisplant,

someofthemborderingsmallE.pauciflorumhabitatpatches.ManyoftheseE.pauciflorum

patcheshavethepotentialtobeusedassteppingstonehabitatbyindividualbutterfliesas

theymovefromonecolonytoanother,movementsthatarelikelytobecriticalinthe

maintenanceofametapopulationofA.mormoinGNP.

Futuredirections

Althoughtheworkpresentedinthisthesishasmadesubstantialstridesinadvancing

understandingoftheecologyofnorthernperipheralpopulationsofA.mormo,likemost

research,italsohighlightstheneedforadditionalinvestigation.Althoughworkonother

populationsrevealsthatcaterpillarsaretypicallysolitaryforagersanduselitteratthe

baseofhostplantsasshelter,littleisknownaboutcaterpillarforagingandmovementin

thenorthernpopulation(Petersonetal.2010).Thus,futureworktobetterunderstandthe

movementanddispersalofthecaterpillarwouldbevaluable.InourpaperintheCanadian

FieldNaturalistwedocumentedthatA.mormoexhibitsdivergentovipositionchoicein

northernpopulationsandthusotherlifehistorymodificationsmayexist(ChapterIVof

thisthesis;Wicketal.2012).Thisnewknowledgeofthebutterflyinitsnorthernrange

leadsonetoaskhowelsethisbutterflymayhavechangeditslifehistorystrategytoadapt

tothefrigid,blusterywintersandhot,drysummersinsouthernSaskatchewan?

Myspeciesdistributionmodelsandassociatedmaps(ChapterII)successfullypredicted

locationsofmanynewcoloniesofA.mormoinGNPandtheVMCP.Datausedtodevelop

thesemodelsarewidelyavailableasdigitalelevationmodels(DEMs)andSPOTsatellite

imagery,bothofwhichhavecoveragethatextendsacrosstheentiretyofthebutterfly’s

knownandpotentialnorthernrange.Ichosetouseonlythesedatatoenhancepotential

transferabilityofthesemodelstootherareaswherethebutterflyisnotknowntooccur,

suchasthoseintheotherprairieregionsinsouthernSaskatchewan,Manitoba,orAlberta.

PrevioussearchesinAlbertahavenotyieldedA.mormo(Anweiler2007),however,with

thehelpofmymodels,areasworthyoffocuscouldbeidentifiedandspecificsitescouldbe

searched.Inthefuture,Iwouldrecommendthatmodelparameterizationincludesoils

data,asIshowedthatsoilsalinityandacidityarerelatedtobutterflypresenceonhost

planthabitatatthemicrohabitatlevelinChapterIII,andsuchdataarecommonly

available.

Itwasnotwithinthescopeofmystudiestoinvestigaterelationshipsbetweenplant

chemistryandhostplantselectionorlarvalsurvival.However,investigationofthis

relationshipmayhelpanswerthequestionofwhythebutterflyoccupiessomehostplants

andnotothers.Butterflyovipositionchoiceandlarvalfeedinghavebeendirectlylinkedto

plantquality,particularlyintermsofprimaryandsecondarychemistry(Carter&Feeney

1999).

Thus,therearemyriadveinsforfutureworkonA.mormoinitsCanadianrange,anditis

myhopethattheideaspresentedinthisthesiswillpromoteandprovidebaselinedatafor

futureinvestigations.

Literaturecited

Carter,M.&P.Feeney.1999.Host‐plantchemistryinfluencesovipositionchoiceofthe

spicebushswallowtailbutterfly.JournalofChemicalEcology.25:1999‐2009.

Mattoni,R.,G.F.Pratt,T.R.Longcore,J.F.Emmel,&J.N.George.1997.Theendangered

quinocheckerspotbutterfly,Euphydryasedithaquino(Lepidoptera:Nymphalidae).The

JournalofResearchontheLepidoptera.34:99‐118.

Monk,S.2012.TheDeathofthePFRA:DestroyingCanada’s“GreatestSuccessStory.”

CanadianCowboyMagazine2012:31‐35.

Murphy,D.D.&P.R.Ehrlich.1988.TheconservationbiologyofCalifornia’sremnant

nativegrasslands.InL.F.HuennekeandH.A.Mooney,Grasslandstructureandfunction:

Californiaannualgrassland.Junk,Dordrecht,TheNetherlands.201‐211.

Murphy,D.D.&S.B.Weiss.1988.Ecologicalstudiesandtheconservationofthebay

checkerspotbutterfly,Euphydryasedithabayensis.BiologicalConservation46:183‐200.

New,T.R.,R.M.Pyle,J.A.Thomas,C.D.Thomas,&P.C.Hammond.1995.Butterfly

ConservationManagement.AnnualReviewofEntomology40:57‐83.

Osborne,K.H.&R.A.Redak.2000.Microhabitatconditionsassociatedwiththe

distributionofpostdiapauselarvaeofEuphydryasedithaquino(Lepidoptera:

Nymphalidae).AnnalsoftheEntomologicalSocietyofAmerica93:110‐114

Peterson,K.,E.Amosa,S.D.Pruss&N.Erbilgin.2010.Firstcaterpillarobservationsofthe

Mormonmetalmark(Apodemiamormo)(Lepidoptera:Riodinidae)butterflyinGrasslands

NationalPark,Saskatchewan,Canada.BlueJay.68:37‐40.

Proctor,J.&S.R.J.Woodwell.1975.Theecologyofserpentinesoils.Advancesin

EcologicalResearch9:255‐366.

SpeciesatRiskAct(SARA).2003.

Thomas,C.D.,J.A.Thomas&M.S.Warren.1992.Distributionsofoccupiedandvacant

butterflyhabitatsinfragmentedlandscapes.Oecologia92:563‐567.

Wark,W.2012.PrairieFarmRehabilitationAdministration(PFRA).TheEncyclopediaof

Saskatchewan.Availableonline:http://esask.uregina.ca

Wick,A.A.,Jannelle,J.,Pruss,S.&Erbilgin,N.2012.Firstobservationsofoviposition

behaviouroftheMormonmetalmark(Apodemiamormo)inSaskatchewan.TheCanadian

FieldNaturalist.126:34‐37.

Documentscited

MetalmarkApodemiamormoinCanada.CommitteeontheStatusofEndangeredWildlife

inCanada.Ottawa.vii+22pp.

Cushman,J.H.2009.ImpactofcattlegrazingontheSmith’sBlueButterfly,itshostplant

andthesurroundingplantcommunity.PreparedforTheNatureConservancyandother

ManagingPartnersofPaloCoronaRegionalPark–MontereyPeninsulaRegionalPark

DistrictandBigSurLandTrust.

Pruss,S.D.,A.Henderson,P.Fargey&J.Tuckwell.2008.RecoveryStrategyforthe

MormonMetalmark(Apodemiamormo)PrairiePopulation,inCanada.SpeciesatRiskAct

RecoveryStrategySeries.ParksCanadaAgency.Ottawa.vi+23pp.

USFWSAntiochDunesDraftComprehensiveConservationPlan.2001.U.S.Fishand

WildlifeServiceCalifornia/NevadaRefugePlanningOffice.

Personalcommunications

Dyck,Trevor.2012.LandManager,AgricultureandAgrifoodCanada(AAFC)

Olson,Wes.2012.Warden,GrasslandsNationalPark.

APPENDIXI.

MarkreleaserecaptureoftheMormonmetalmark

Table.Appendix‐1.MarkreleaserecaptureresultsfromAugust2011.G6‐BH1areplots

establishedinGrasslandsNationalPark,VM10‐VM14representplotsestablishedinthe

ValMarieCommunityPasture.Atotalof885butterflieswerecaughtwith142recaptures.

95%CI SE Ha

N*HAT/

G6 1097.49 887.08 1307.90 107.35 2.54 0.04 431.59

G5 534.23 397.17 671.29 69.93 0.18 0.29

2918.0

G4 359.94 183.86 536.03 89.84 0.49 0.07 730.54

BH1 417.56 286.12 548.99 67.06 0.82 0.05 510.09

0 99.14 26.45 171.82 37.08 0.27 0.04 361.38

2 88.29 48.87 127.72 20.12 0.41 0.02 215.86

4 56.26 9.68 102.85 23.77 0.32 0.02 177.47

Table.Appendix‐2.AnexplanationofthetermsshowninTable6‐1.Allresultswerebased

onanalysesfromtheProgramMARK(White2012;www.phidot.org).

TerminTable6‐1 ExplanationofTerm

N*HAT PopulationestimationfromJolly‐SeberMethod

Lower95%CI ConfidentintervalforN*HAT

Upper95%CI ConfidentintervalforN*HAT

SE StandarderrorforN*HAT

Ha Hectareofeachplot

N*HAT/m2 Populationestimation(N*HAT)perm2

N*HAT/Ha Populationestimation(N*HAT)perhectare

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