UniversityofAlberta

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

by

AshleyAnneWick

AthesissubmittedtotheFacultyofGraduateStudiesandResearchinpartialfulfillmentoftherequirementsforthedegreeof

MasterofScience

in

ConservationBiology

DepartmentofRenewableResources

©AshleyAnneWickSpring2013

Edmonton,Alberta

PermissionisherebygrantedtotheUniversityofAlbertaLibrariestoreproducesinglecopiesofthisthesisandtolendorsellsuchcopiesforprivate,scholarlyorscientificresearchpurposesonly.Wherethethesisisconvertedto,orotherwisemadeavailableindigitalform,theUniversityofAlbertawilladvisepotentialusersofthethesisof

theseterms.

Theauthorreservesallotherpublicationandotherrightsinassociationwiththecopyrightinthethesisand,exceptashereinbeforeprovided,neitherthethesisnoranysubstantialportionthereofmaybeprintedorotherwise

reproducedinanymaterialformwhatsoeverwithouttheauthor'spriorwrittenpermission.

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

1

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

2

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

3

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.

4

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).

5

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

6

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.

7

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

8

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.

9

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

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

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Henderson,A.,P.Fargey,S.Pruss.F.Sperling.2008.Earlysightingofararebutterfly,

MormonMetalmark,inGrasslandsNationalPark,SK.BlueJay66:105‐106.

Hooper,R.R.1973.ButterfliesofSaskatchewan.SaskatchewanDepartmentof

NaturalResources,p.201.

Howe,W.E.1975.TheButterfliesofNorthAmerica.Doubleday,N.Y.,p268.

Layberry,R.A.,P.W.Hall&J.D.Lafontaine.1998.TheButterfliesofCanada.

UniversityofTorontoPress:280.

Michalsky,S.J.&R.A.Ellis.1994.VegetationofGrasslandsNationalPark.Grasslands

NationalPark,ValMarie,Saskatchewan.April1994.

Morse,S.2009.BringingabutterflybackfromtheBrink.U.S.Fish&WildlifeService

EndangeredSpeciesBulletin34:6‐7.

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

ConservationManagement.AnnualReviewofEntomology40:57‐83.

Opler,P.A.&J.A.Powell.1962.Taxonomicanddistributionstudiesofwestern

componentsoftheApodemiamormocomplex(Riodinidae).Journalofthe

Lepidopterists’Society15:145‐171.

Opler,P.A.1999.Afieldguidetowesternbutterflies.2nded.HoughtonMifflin

Company,Boston.540pp.260.

Pelham,J.P.2008.AcatalogueofthebutterfliesoftheUnitedStatesandCanada

withacompletebibliographyofthedescriptiveandsystematicliterature.Journalof

ResearchonLepidoptera40:1‐652.

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

oftheMormonmetalmark(Apodemiamormo)(Lepidoptera:Riodinidae)butterflyin

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

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ofNorthAmerica.Doubleday,GardenCity,NewYork,pp.259‐272.

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Sperling.2013.ApodemiamormoinCanada:populationgeneticdatasupportprior

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StanfordUniversityPress,Stanford.

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Thomas,J.A.2005.Monitoringchangeintheabundanceanddistributionofinsects

usingbutterfliesandotherindicatorgroups.PhilosophicalTransactionsoftheRoyal

SocietyB360:339‐357.

ThorpeJ.&N.Nicolichuk.2009.PredictivevegetationmappingatGrasslands

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metalmark(Apodemiamormomormo)ovipositionbehaviourinCanada.Canadian

FieldNaturalist:126:34‐37.

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Anweiler,G.G.2007.2007SearchfortheMormonMetalmark(Apodemiamormo)in

Alberta.UnpublishedReport.

Cannings,R.J.,E.Durance&L.K.Scott.1998.SouthOkanaganecosystemrecovery

plan:Scientificassessment.UnpublishedReport,BritishColombiaMinistryof

Environment.Penticton108.

COSEWIC.2003.COSEWICassessmentandupdatestatusreportontheMormon

MetalmarkApodemiamormoinCanada.CommitteeontheStatusofEndangered

WildlifeinCanada.Ottawa.vii+22pp.

Hooper,V.L.2002.StatusReportofApodemiaMormo(C.R.Felder)inSaskatchewan.

4pp.

Johnson,J.J.,Jones,J.,Baudour,M.,Wagner,M.,Flannery,D.,Werner,D.,Holden,D.,

Virun,K.,Wilson,T.,Delijani,J.,Delijani,J.,Newton,B.,Gundell,D.,Thummar,B.,

Blakey,C.,Walsh,K.,Sweeney,Q.,Ware,T.,Adams,A.,Taylor,C.,andLongcore,T.

2011.CaptiveRearingofLange’sMetalmarkButterfly,2010‐2011.FinalReportto

NationalFishandWildlifeFoundation.TheUrbanWildlandsGroup.

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Pruss,S.D.,A.Henderson,P.Fargey,andJ.Tuckwell.2008.RecoveryStrategyfor

theMormonMetalmark(Apodemiamormo)PrairiePopulation,inCanada.Speciesat

RiskActRecoveryStrategySeries.ParksCanadaAgency.Ottawa.vi+23pp.

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Olson,W.2012.Warden,GrasslandsNationalPark.

13

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).

14

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;

15

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).

16

AllknownA.mormocoloniesinSaskatchewanarelocatedwithinGNPandthe

VMCP.

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

17

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

18

(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:

19

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;

20

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

21

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

22

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).

23

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

24

associations.ToolssuchasSDMsmayhelpconservationistsandlandmanagers

predictandunderstanddistributions,assistinmakingdecisionsregarding

reintroduction,restoration,andmeasurestoreduceprobabilityoflocalextirpation

(Maneletal.2001;Bartel&Sexton2009).Inthefaceoftheperniciouseffectsof

climatechangeandhabitatalteration,habitatmodelingmethodsmaysavevaluable

timeandenergyindocumentingspecies’distributionsaswellasuncovering

unknownpopulations.

25

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.

26

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33

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

DEM SurfaceAnalyst(Spatial

Analyst,ArcGIS10)

Slope

(SLOPE)

Degrees 20m

DEM SurfaceAnalyst(Spatial

Analyst,ArcGIS10)

Aspect

(ASPECT)

Categorial

(N,NE,E,

etc.)

20m

DEM SurfaceAnalyst(Spatial

Analyst,ArcGIS10)

Hillshade

(HLSHD)

NA* 20m

DEM SurfaceAnalyst(Spatial

Analyst,ArcGIS10)

AreaSolar

Radiation

(SOLAR202)

NA* 20m

SRCPredictive

vegetation

model

NA* Vegetation

(SRCE2)

Index 1:20,000

polygon

DEM Rastercalculator

Compound

Topographic

Index(CTI)

Index 20m

SPOT5(2007)

SatelliteImagery

Rastercalculator

Normalized

Difference

Vegetation

Index

Index 10m

34

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

35

Figure2‐1.ThestudyareconsistingoftheValMarieCommunityPastureand

GrasslandsNationalPark’sEastandWestBlocks,insouthernSaskatchewan,

Canada.

36

Figure2‐2.VariableimportanceplotgeneratedbytheRandomForestsalgorithm.

Theplotshowsthevariableimportancemeasuredasincreasednodepurity

(IncNodePurity)andtheincreaseofmeansquareerror(%IncMSE),which

representsthedeteriorationofthepredictiveabilityofthemodelwheneach

predictionisreplacedinturnbyrandomnoise.Ahighervalueof%IncMSEindicates

ahighervariableimportance.ThefullvariablenamesareshowninTable1.

37

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.

38

Figure2‐4.Numberofobserved(siteswhereApodemiamormowasactuallyfound)

versusexpected(sitespredictedtohaveApodemiamormopresences)byeach

model(probabilityof0.60orhigher)weresearchedinGrasslandsNationalPark

andtheneighboringValMarieCommunityPastureinsouthernSaskatchewan,

Canada.

27

45

73

42

59

101

0

20

40

60

80

100

120

RSF RandomForests

RSF&RandomForests

Numberofsites

Model

Observed

Expected

39

Figure2‐5.Numberofobserved(siteswhereApodemiamormowasnotfound)

versusexpected(sitespredictedtonothaveApodemiamormo)bytheRandom

ForestsmodelforboththeValMarieCommunityPasture(VMCP)andGrasslands

NationalPark(GNP)insouthernSaskatchewan,Canada.

24

46

70

25

51

76

0

10

20

30

40

50

60

70

80

VMCP GNP VMCP&GNP

Numberofsites

Location

Observed

Expected

40

Figure2‐6.Numberofobserved(siteswhereApodemiamormowasactuallyfound)

versusexpected(sitespredictedtohaveApodemiamormopresences)modelresults

separatedbasedonwhetherornotanareawasungrazed,grazedbybison,or

grazedbycattle.TheValMarieCommunityPasture(VMCP)isgrazedbycattle,the

EastBlockofGrasslandsNationalPark(GNP)isnotgrazed,andtheWestBlockof

GNPisgrazedbybison,allsitesarelocatedinsouthernSaskatchewan,Canada.

6

23

77

31 27

106

0

20

40

60

80

100

120

Cattlegrazed Notgrazed Bisongrazed

VMCP EastBlock(GNP)

WestBlock(GNP)

Numberofsites

Locations

Observed

Expected

41

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

42

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).

43

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

44

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

45

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

46

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

47

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).

48

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.

49

Mostimportantly,theresultspresentedhereimproveunderstandingofhabitat

requirementsforA.mormoinitsnorthernrange,andcanbeusedbylandmanagers

toidentifyandconservehabitatforthebutterfly.Thefollowingcriteriashouldbe

helpfulfordefininganddesignatingcriticalhabitatforA.mormo:patcheswithhigh

%ofE.pauciflorumcover,growingonsoilswithhigherpHandlowavailable

nitrogen,onsteepslopesthataresouth‐westtosouthfacingatlowerthanaverage

elevation,inbadlandhabitatwithbaregroundcover,andparticularly,areasthatare

currentlyorhavebeenrecentlyoccupiedbythebutterfly.

50

Acknowledgements

ThisprojectwasfundedbyParksCanadaandtheUniversityofAlberta.This

researchwouldnothavebeenpossiblewithoutthehelpofmyfaithfulfield

assistant,HeatherSauder,aswellasindispensibleassistancefromthepeopleofVal

Marie,SaskatchewanandGrasslandsNationalPark.

51

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58

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.

59

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

60

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 +

61

Figure3‐1.Lineardiscriminantanalysisshowsthedistribution“discriminants”of

thetwodistributions,thefirstofwhichis“unoccupied”habitatandthesecond,

whichis“occupied”habitat.

62

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.

63

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.

64

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

65

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.

66

Acknowledgements

Manythankstoourfaithfulfull‐timefieldassistant/localguide,HeatherSauder,as

wellastovolunteersfromacrossNorthAmericaandtheeverhelpfulstaffat

GrasslandsNationalPark.Thanksalsotothereviewersfortheirhelpfulcomments.

ThisprojectwasfundedbytheParksCanadaAgencyandtheUniversityofAlberta.

67

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Arnold,R.A.,andJ.A.Powell.1983.Apodemiamormolangei.Chapter6inEcological

studiesofsixendangeredbutterflies(Lepidoptera,Lycaenidae):Island

biogeography,patchdynamicsanddesignofhabitatpreserves.Universityof

CaliforniaPublicationsinEntomology99:1‐161.

Berger,D.,R.Walters,andK.Gotthard.2008.Whatlimitsinsectfecundity?Bodysize

andtemperature‐dependenteggmaturationandovipositioninabutterfly.

FunctionalEcology22:523‐529.

Bonebrake,T.C.,C.L.Boggs,J.M.McNally,J.Ranganathan,andP.R.Ehrlich.2010.

Ovipositionbehaviourandoffspringperformanceinherbivorousinsects:con‐

sequencesofclimaticandhabitatheterogeneity.Oikos119:927‐934.

Bryant,S.R.,C.D.Thomas,andJ.S.Bale.2002.Theinfluenceofthermalecologyon

thedistributionofthreenymphalidbutterflies.JournalofAppliedEcology39:43‐

55.

CommitteeontheStatusofEndangeredWildlifeinCanada(COSEWIC).2003.

COSEWICassessmentandupdatestatusreportontheMormonMetalmark

ApodemiamormoinCanada.CommitteeontheStatusofEndangeredWildlifein

Canada,Ottawa,Ont.vii+22pages.

De‐Silva,D.L.,A.S.Vasquez,andJ.Mallet.2011.Selectionforenemy‐freespace:eggs

placedawayfromthehostplantincreasesurvivalofaneotropicalithomiine

butterfly.EcologicalEntomology36:667‐672.

Elrod,MortonJ.1970.TheButterfliesofMontanawithKeysforDeterminationof

Species.TheMidcontinentLepidopteraSeries,no.17.AssociationofMinnesota

Entomologists.(Reprintfrom1906UniversityofMontana,Missoula,Montana).

68

Fischer,K.,A.N.M.Bot,P.M.Brakefield,andB.J.Zwaan.2003a.Fitness

consequencesoftemperature‐mediatedeggsizeplasticityinabutterfly.Functional

Ecology17:803‐810.

Fischer,K.,E.Eenhoorn,A.N.M.Bot,P.M.Brakefield,andB.J.Zwaan.2003b.Cooler

butterflieslaylargereggs:developmentalplasticityversusacclimation.Proceedings

oftheRoyalSocietyofLondonB270:2051‐2056.

Fischer,K.,A.N.M.Bot,andP.M.Brakefield.2004.Geneticandenvironmental

sourcesofeggsizevariationinthebutterflyBicyclusanynana.Heredity92:163‐169.

Geister,T.L.,M.W.Lorenz,K.H.Hoffmann,andK.Fischer.2009.Energeticsof

embryonicdevelopment:effectsoftemperatureoneggandhatchlingcompositionin

abutterfly.JournalofComparativePhysiology179:87‐98.

Hooper,R.R.2002.StatusReportofApodemiamormo(C.R.Felder)in

Saskatchewan.4pages.

Johnson,J.J.,J.Jones,M.Baudour,M.Wagner,D.Flannery,D.Werner,D.Holden,K.

Virun,T.Wilson,J.Delijani,J.Delijani,B.Newton,D.Gundell,B.Thummar,C.Blakey,

K.Walsh,Q.Sweeney,T.Ware,A.Adams,C.Taylor,andT.Longcore.2011.Captive

rearingofLange’sMetalmarkbutterfly,2010‐2011.FinalReporttoNationalFish

andWildlifeFoundation.TheUrbanWildlandsGroup,LosAngeles,California.

Koda,K.,andH.Nakamura.2010.Effectsoftemperatureonthedevelopmentand

survivalofanendangeredbutterfly,Shijimiaeoidesdivinusbarine(Leech)

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

Kohler,Steve.1980.ChecklistofMontanabutterflies(Rhopalocera).Journalofthe

Lepidopterists’Society34:1‐19.

Layberry,RossA.,P.W.Hall,andJ.D.Lafontaine.1998.TheButterfliesofCanada.

UniversityofTorontoPress,Toronto,Ontario.280pages.

69

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

oftheMormonmetalmark(Apodemiamormo)(Lepidoptera:Riodinidae)butterfly

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

Proshek,B.,L.A.Crawford,C.S.Davis,S.Desjardins,A.E.Henderson&F.A.H.

Sperling.2013.ApodemiamormoinCanada:populationgeneticdatasupportprior

conservationranking.JournalofInsectConservation17:155‐170.

Pruss,S.D.,A.Henderson,P.Fargey,andJ.Tuckwell.2008.Recoverystrategyforthe

MormonMetalmark(Apodemiamormo)prairiepopulationinCanada.Speciesat

RiskActRecoveryStrategySeries.ParksCanadaAgency,Ottawa,Ontario.vi+23

pages.

Pyle,R.M.2002.TheButterfliesofCascadia:Afieldguidetoallthespeciesof

Washington,Oregon,andsurroundingterritories.SeattleAudubonSociety,Seattle.

420pages.

Scott,JamesA.1986.TheButterfliesofNorthAmerica,aNaturalHistoryandField

Guide.StanfordUniversityPress,PaloAlto,California.581pages.

Steigenga,M.J.,andK.Fischer.2007.Ovariandynamics,eggsize,andeggnumberin

relationtotemperatureandmatingstatusinabutterfly.Entomologia

ExperimentalisetApplicata125:195‐203.

70

Tables&Figures

Figure4‐1.FemaleMormonMetalmark,Apodemiamormo,ovipositingincracksin

thesoilinTimmonscoulee,GrasslandsNationalParkofCanada(westblock),near

ValMarieonAugust21,2011.Photo:JohaneJanelle.

71

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

72

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

73

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.

74

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

75

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

76

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.

77

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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

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SpeciesatRiskAct(SARA).2003.

http://www.sararegistry.gc.ca/the_act/HTML/Guide_e.cfm

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

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COSEWIC.2003.COSEWICassessmentandupdatestatusreportontheMormon

MetalmarkApodemiamormoinCanada.CommitteeontheStatusofEndangeredWildlife

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

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ManagingPartnersofPaloCoronaRegionalPark–MontereyPeninsulaRegionalPark

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

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80

APPENDIXI.

MarkreleaserecaptureoftheMormonmetalmark

Table.Appendix‐1.MarkreleaserecaptureresultsfromAugust2011.G6‐BH1areplots

establishedinGrasslandsNationalPark,VM10‐VM14representplotsestablishedinthe

ValMarieCommunityPasture.Atotalof885butterflieswerecaughtwith142recaptures.

N*HAT

Lower

95%CI

Upper

95%CI SE Ha

N*HAT/

m2

N*HAT

/Ha

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

1

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

VM1

0 99.14 26.45 171.82 37.08 0.27 0.04 361.38

VM1

2 88.29 48.87 127.72 20.12 0.41 0.02 215.86

VM1

4 56.26 9.68 102.85 23.77 0.32 0.02 177.47

81

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