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Sex and hibernaculum temperature predict survivorship in white-nose syndrome affected little brown myotis (Myotis lucifugus)

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Title:
Sex and hibernaculum temperature predict survivorship in white-nose syndrome affected little brown myotis (Myotis lucifugus)
Series Title:
Royal society open science
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Grieneisen, Laura E.
Brownlee-Bouboulis, Sarah A.
Johnson, Joseph S.
Reeder, DeeAnn M.
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Royal Society Publishing
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English
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Myotis ( lcsh )
White-nose syndrome ( lcsh )
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serial ( sobekcm )

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Abstract:
White-nose syndrome (WNS), an emerging infectious disease caused by the novel fungus Pseudogymnoascus destructans, has devastated North American bat populations since its discovery in 2006. The little brown myotis, Myotis lucifugus, has been especially affected. The goal of this 2-year captive study was to determine the impact of hibernacula temperature and sex on WNS survivorship in little brown myotis that displayed visible fungal infection when collected from affected hibernacula. In study 1, we found that WNS-affected male bats had increased survival over females and that bats housed at a colder temperature survived longer than those housed at warmer temperatures. In study 2, we found that WNS-affected bats housed at a colder temperature fared worse than unaffected bats. Our results demonstrate that WNS mortality varies among individuals, and that colder hibernacula are more favourable for survival. They also suggest that female bats may be more negatively affected by WNS than male bats, which has important implications for the long-term survival of the little brown myotis in eastern North America.

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K26-05072 ( USFLDC DOI )
K26.5072 ( USFLDC handle )

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rsos.royalsocietypublishing.orgResearch Citethisarticle: GrieneisenLE, Brownlee-BouboulisSA,JohnsonJS, ReederDM.Sexandhibernaculum temperaturepredictsurvivorshipin white-nosesyndromeaectedlittlebrown myotis(Myotislucifugus). R.Soc.opensci. :b. http://dx.doi.org/.tn/rsos.b Received:December Accepted:nJanuary SubjectCategory: Biology(wholeorganism) SubjectAreas: ecology/healthanddiseaseand epidemiology/behaviour Keywords: Chiroptera,diseaseecology, Pseudogymnoascusdestructans Authorforcorrespondence: DeeAnnM.Reeder e-mail: dreeder@bucknell.edu Sexandhibernaculum temperaturepredict survivorshipinwhite-nose syndromeaectedlittle brownmyotis(Myotis lucifugus) LauraE.Grieneisen,SarahA.Brownlee-Bouboulis, JosephS.JohnsonandDeeAnnM.Reeder DepartmentofBiology,BucknellUniversity,Lewisburg,PAbnb,USA LEG, --bnf;DMR,--nf-.SummaryWhite-nosesyndrome(WNS),anemerginginfectiousdisease causedbythenovelfungus Pseudogymnoascusdestructans ,has devastatedNorthAmericanbatpopulationssinceitsdiscoveryin 2006.Thelittlebrownmyotis, Myotislucifugus ,hasbeenespecially affected.Thegoalofthis2-yearcaptivestudywastodeterminethe impactofhibernaculatemperatureandsexonWNSsurvivorship inlittlebrownmyotisthatdisplayedvisiblefungalinfectionwhen collectedfromaffectedhibernacula.Instudy1,wefoundthat WNS-affectedmalebatshadincreasedsurvivaloverfemalesand thatbatshousedatacoldertemperaturesurvivedlongerthan thosehousedatwarmertemperatures.Instudy2,wefoundthat WNS-affectedbatshousedatacoldertemperaturefaredworse thanunaffectedbats.OurresultsdemonstratethatWNSmortality variesamongindividuals,andthatcolderhibernaculaaremore favourableforsurvival.Theyalsosuggestthatfemalebatsmay bemorenegativelyaffectedbyWNSthanmalebats,whichhas importantimplicationsforthelong-termsurvivalofthelittle brownmyotisineasternNorthAmerica..IntroductionWhite-nosesyndrome(WNS)isanemerginginfectiousdisease estimatedtohavekilledover5.7millionNorthAmerican bats[1]inthe4yearsfollowingitsdiscovery.Itiscaused bythepsychrophilicfungus, Pseudogymnoascusdestructans (Pd) [2]thatgrowsintheskinofaffectedbatsduringperiodsof prolongedtorpor,orhibernation[3].Atleastsevenspecies TheAuthors.PublishedbytheRoyalSocietyunderthetermsoftheCreativeCommons AttributionLicensehttp://creativecommons.org/licenses/by/./,whichpermitsunrestricted use,providedtheoriginalauthorandsourcearecredited. on April 21, 2017 http://rsos.royalsocietypublishing.org/ Downloaded from

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rsos.royalsocietypublishing.org R.Soc.opensci. :b................................................ofbatsareaffected,butlittlebrownmyotis( Myotislucifugus )areespeciallysusceptible,withanaverage 91%declineinnortheasternNorthAmerica[ 4].WNSaffectshibernationbehaviourandsurvivalin bats[2,5];theroleofmicroclimatefactors,suchashibernaculumtemperature,onthisrelationshipis mostlyunknown,althoughpopulationsoflittlebrownmyotisinwarmerWNS-affectedhibernacula suffergreaterdeclinesthanthoseincolderhibernacula[6].Inhealthybats,hibernaculumtemperature playsasignicantroleinenergybalanceandsurvivorship[ 7].Whenabatenterstorporitsmetabolism isdepressedandbodytemperaturedropstowithin1Cofambienttemperature[8].Hibernatorsuse upto90%oftheirstoredenergyforarousalbouts[9],inwhichtheybrieywarmuptoeuthermic bodytemperature.Arousingfromawarmertemperatureuseslessenergythanarousingfromacolder temperature,thusbatsroostingatwarmertemperaturescanarousemorefrequently[ 7].Sexalsoplays aroleinhibernationenergetics;femalebatsfaceselectivepressuretoretainenoughfatattheendof hibernationtoovulateuponspringemergence[10,11]. Thegoalofthisstudywastodeterminetheimpactofhibernaculumtemperatureandothercovariates suchassexandbodymassindex(BMI)onWNSsurvivorshipinlittlebrownmyotisincontrolled laboratorystudies.Because Pd growsoptimallybetween12.5Cand15.8C,andmuchmoreslowlyat lowertemperatures[12]wehypothesizedthatWNS-affectedbatshousedatcoldertemperatureswould exhibitincreasedsurvivaloverthosehousedatwarmertemperatures.Wefurtherhypothesizedthat femalesinfectedwith Pd wouldhaveincreasedsurvivalratesovermales,ashealthyfemalelittlebrown myotisstartandendhibernationwithgreaterfatreserves[ 11],andmaybebetterequippedtodealwith anenergeticallycostlydiseaseduringhibernation..Materialandmethods..StudyLittlebrownmyotiswerecollectedfromWNS-unaffected( n = 58)andWNS-affected(n = 49)hibernacula inPennsylvaniaon13and15January2010,respectively,andtransportedtoBucknellUniversitysbat vivarium.EverybatfromtheWNSsitehadvisiblefungalgrowthatcollectionandwasthuspresumed tobeWNS-affected.Uponarrivalatthelaboratory,dataonweight,sexandforearmlengthwererecorded foreachbatandBMI(massing /lengthofforearminmm)wascalculated.Batswerehousedfollowing [13]andinducedtohibernatebyplacingtheminadarkenedenvironmentalchamber.Batswereevenly distributedbetweensixnon-adjacentwiremeshcages(46 46 61cm);oneforunaffectedbats,one forWNS-affectedbatsineachofthreeenvironmentalchambers(setto4C,7Cor10C; table1).Lorch etal.[ 14],undernearlyidenticalhousingconditions,demonstratedthat Pd transmissiondoesnotoccur betweencages,and,asvisiblefungalgrowthwasnotobservedontheunaffectedbatsatanytimein ourstudyitisunlikelytransmissionoccurredbetweenthegroups.Chamberswerecheckedweekly formortalityanddeadbatswereremoved.Thisprovidedarelativedeathdate,sosurvivalcouldbe calculatedviaCoxregressiononthefollowingvariables:WNSstatusofasite,chambertemperature,sex andinitialBMI.Additionally,asubsetofanimalsfromeachgroupwereremovedonceforexperimental testingofthermalpreferencesforaseparatestudy.Whilethesetrialswereconductedwithaffectedand unaffectedbatsequally,thisdisturbance,combinedwithweeklyremovalofdeadanimals,verylikely affectedoverallsurvivorshipforallbats(seeResults)...StudyTofollowupontheresultsofstudy1,littlebrownmyotiswerecollectedfromapresumedunaffected hibernaculuminKentucky( n = 40)andfromaWNS-affectedhibernaculum( n = 40)inPennsylvania on15and21December2010,respectively.CollectionofanimalsandresearchinPennsylvaniawas conductedunderaPennsylvaniaGameCommissionpermittoDMR(no.183).Collectionofanimals fromKentuckywasconductedbyastatewildlifeofcial(BrookeHines)onnon-endangeredbats;thus anumberedpermitwasnotrequiredorissued.Althoughthepresenceof Pd wasdetectedonabatfrom theunaffectedsiteinApril2011,nomortalitywasdetectedatthesitethroughspring2012,strongly suggestingthebatsusedinthisstudywereunaffected[ 13].Inordertoavoiddisturbingmorebatsthan necessary,batswerecollectedregardlessofsex,resultinginanaldatasetthatwastooheavilymalebiased(77%)toincludesexintheanalysis.Batswereprocessedandhousedasdescribedinstudy1,and evenlydistributedbetween4Cand10Cchambers( table2 ).Eachbatwasttedwithatemperaturesensitivedataloggertorecorditsskintemperature( Tskin;[ 5])at30minintervals.Becauseofhigh on April 21, 2017 http://rsos.royalsocietypublishing.org/ Downloaded from

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rsos.royalsocietypublishing.org R.Soc.opensci. :b................................................Table. FactorsincludedinthestudyrnalCoxsurvivalregressionmodel.(Ahazardratioof.indicatesa.%lowerriskofdeath.Asterisksindicates ignircanceof p-values.) eect samplesize, n p-valueparameter groupwithhigherriskofdeathhazardratio(t%CI).......................................... ........................................... ........................................................................................................................................................................................................................................................................................siteWNSsite:n .b <. WNS-aected unaectedsite:t.......................................... ........................................... ........................................................................................................................................................................................................................................................................................sex female: ..b .......................................... ........................................... ........................................................................................................................................................................................................ .........................male:.......................................... ........................................... ........................................................................................................................................................................................................................................................................................sex site WNSfemale:.f.*WNSmaleversusWNSfemaleWNSfemale.fn(..nff) unaectedfemale: WNSmale: unaectedmale:n.......................................... ............................................ .............................................................................................................unaectedmaleversusunaectedfemalen.s. .t(.fb.nb).......................................... ............................................ .............................................................................................................unaectedfemaleversusWNSfemaleWNSfemale .(.n.nb).......................................... ............................................ .............................................................................................................unaectedmaleversusWNSmale WNSmale .(.t.f).......................................... ........................................... ........................................................................................................................................................................................................................................................................................housingtemperature*C:n .b.*CversusbCbC .tnf(..b) bC: C:.......................................... ............................................ .............................................................................................................CversusC C .bf(.f.n).......................................... ............................................ .............................................................................................................bCversusC n.s. .nb(.b.f).......................................... ........................................... ........................................................................................................................................................................................................................................................................................BMIinitial b .b.n .......................................... ........................................... ........................................................................................................................................................................................................................................................................................ on April 21, 2017 http://rsos.royalsocietypublishing.org/ Downloaded from

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rsos.royalsocietypublishing.org R.Soc.opensci. :b................................................Table. FactorsincludedinthestudyrnalCoxsurvivalregressionmodel.(Asterisksindicatesignircanceof p-values.) groupwithhigherhazard eectsamplesize, n p-valueparameterriskofdeathratio(t%CI) site housingCunaectedsite:b..*CunaectedversusCWNS-aected.(.f.nf) temperature*CWNS-aectedsite:f CWNS-aected Cunaectedsite:t CWNS-aectedsite:n Cunaected versusC unaected n.s..t(.n.) Cunaected versusC WNS-aected n.s..nt(.ff.f) CWNS-aected versusC WNS-aected n.s..(.f.ft)............................................................................................................................................................................. ............................................mortalityduringstudy1,batsremainedundisturbeduntil23March2011.Deadbatswereremovedand livebatscontinuedtohibernateuntil4April2011.Asweeklymortalitythereforecouldnotbecalculated, approximatedateofdeath,denedasdateoflastarousalboutbasedon Tskindata,wascalculatedfor thosebatswhosedataloggersdidnotmalfunction( n = 70; gure1 c ).SurvivalwascalculatedviaCox regressiononWNSstatusofasiteandchambertemperature.Additionally,aPearsons 2wasusedto analysehowsiteandtemperaturepredictedmortality,denedaswhetherornotabatsurvived,forthe fulldataset(n = 80)..Results..StudyThefactorsincludedinthenalsurvivalmodelwereWNSstatusofasite,sex,aWNSstatusofasite sexinteractionterm,housingtemperatureandinitialBMI.InitialBMIdidnothaveasignicantimpact onsurvival,nordiditdifferbetweenbatsfromWNS-affectedandunaffectedsites(table1 ).Temperature hadasignicanteffectonsurvival;batshousedat4Chadasignicantlylowerriskofdeaththanthose housedat7Cor10C( table1 and gure1 a).WNSstatusofasiteandtheinteractionbetweensiteand sexalsoweresignicant;batsfromtheWNSsitehadsignicantlylowersurvivalthanthosefromthe unaffectedsite,andWNSsitefemalebatshadahigherriskofdeaththanWNSsitemalebats( table1 and gure1 b),eventhoughfemaleshadhigherinitialBMIthanmales(two-sample t-test; t105= 2.32, p = 0.023)...StudyAhibernationchambertemperature WNSstatusofasiteinteractiontermwastheonlyfactorinthe Coxsurvivalmodel.WNS-affectedbatshousedat4Chadasignicantlyhigherriskofdeaththan unaffectedbats(table2 and gure1 c ).Likewise,batsfromtheWNSsitehadhighermortalitythan thosefromtheunaffectedsite(40%mortalityinunaffectedbatsand65%mortalityinWNS-affected bats; 2 1= 5.01, p = 0.025; table3),whichwasprimarilydrivenbydifferencesinmortalityinbatshoused at4C(30%mortalityinunaffectedbatsand65%mortalityinWNS-affectedbats; 2 1= 4.91, p = 0.027; table3)..DiscussionThisis,toourknowledge,therstcaptivestudytotesthowhibernaculumtemperatureandsexaffect hibernationsurvivalinnaturallyWNS-affectedversusunaffectedbats.WNSstatusofasite,housing temperatureandsexallsignicantlyinuencedsurvivalduringhibernation.TheroleofWNSstatusin survivalwasconsistentinbothstudies;WNS-affectedbatsfaredworsethanunaffectedbats.However, on April 21, 2017 http://rsos.royalsocietypublishing.org/ Downloaded from

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rsos.royalsocietypublishing.org R.Soc.opensci. :b................................................ 0 0 10 20 30 40 50 60 70 80 90 100 10C unaffected (19) 7C unaffected (19) 4C unaffected (21) 10C WNS (16) 7C WNS (16) 4C WNS (17) 1020304050 days into the study (a)cumulative survival (%)60708090100 0 0 10 20 30 40 50 60 70 80 90 100 4C unaffected (17) 10C unaffected (19) 10C WNS (18) 4C WNS (16) 1020304050 da y s into the stud y (c)cumulative survival (%)60708090110100 0 0 10 20 30 40 50 60 70 80 90 100 unaffected females (31) unaffected males (28) WNS males (24) WNS females (24) 1020304050 days into the study (b)cumulative survival (%)60708090100 Figure. CumulativesurvivalbyWNSstatus,temperatureandsex.( a)Instudy,batsfromaWNSsiteandbatshousedatbCorC hadthehighestriskofdeath.(b)Instudy,femalesfromaWNSsitehadahigherriskofdeaththanWNSsitemales.(c )Instudy,bats fromaWNSsitehousedatChadahigherriskofdeaththanbatsfromanunaectedsite.Samplesize, n,inparentheses.theimpactofhousingtemperaturewasmorenuanced.Study1foundthatbatshousedatacolder temperaturefaredbetterthanthosehousedatwarmertemperatures,whereasstudy2foundunaffected batsfaredsignicantlybetterthanWNS-affectedbatsatacoldertemperaturebutnotatawarmer temperature(atwhich Pd exhibitsfastergrowth[12]).TheseresultssuggestthatWNSmayalterthe importanceofhibernaculumtemperatureinsurvivinghibernation.Thiscouldbeduetochangesin on April 21, 2017 http://rsos.royalsocietypublishing.org/ Downloaded from

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rsos.royalsocietypublishing.org R.Soc.opensci. :b................................................Table. Studymortalitybytreatmentgroup. group samplesize, n mortality(%, n)............................................................................................................................................................................. ............................................Cunaected %(b)............................................................................................................................................................................. ............................................CWNS f%()............................................................................................................................................................................. ............................................Cunaected %()............................................................................................................................................................................. ............................................CWNS f%()............................................................................................................................................................................. ............................................energeticexpenditurebyWNS-affectedbats,perhapsbyincreasingtheamountoftimespentgrooming andinotherexpensivebehaviours[ 13],aswellasadisruptioninphysiology[ 15].Thedifferences betweenstudy1andstudy2areprobablyowingto:(i)theinherentvariationinthephysiologyof capturedfree-rangingbats,and(ii)thefactwhileweonlycollectedbatsfromtheaffectedsitethathad visiblefungalgrowthontheirmuzzlesandwings,wecouldnotinanywaycontrolforinfectionload ordiseasestage.Owingtothisvariation,ourresultsshouldbeinterpretedwithcaution.Trulyteasing aparttheinteractionbetween Pd growthdynamicsandtemperatureonWNSsurvivorshiprequires experimentalstudiesinwhichinitialfungalloadiscontrolled.Indeed,we[ 16]performedtheseexact experimentsincaptivitybyinoculatingnaivebatswithknownfungalloadsandhousingthemattwo hibernationtemperatures.AsinthisstudyinwhichbatswerenaturallyWNS-affected,asignicanteffect ofhibernationtemperatureonsurvivorshipinaffectedbatswasfound,evenaftercontrollingforthe overallbenetsofcolderhibernaculaonsurvivorship.Thatwehavenowdemonstratedthiseffectin atleastsomenaturallyaffectedbatsandinexperimentallyinoculatedbatshighlightstheimportance ofhibernaculatemperatureonsurvivalinthefaceofWNS.Thendingthatcoldertemperatures areprotectiveforbatssuggeststhathibernaculatemperaturemodication(especiallyofmines)may beaviablemitigationstrategyforcurbingWNSandthatprotectionofcolderhibernaculashould beprioritized. Thisis,toourknowledge,therststudytoshowthattherearesexdifferencesinWNSsurvivalin naturallyaffectedbats,withfemalebatsatgreaterriskofdyingthanmales.However,theseresultsare notcongruentwithwhatwouldbeexpectedifsexdifferencesinsurvivalwerebasedondifferencesin energydistributionbetweenmalesandfemales.Ourstudyandpaststudieshavefoundthatfemales starthibernationwithhigherBMIthanmales[ 11,17],whichshouldbiasfemalestowardsincreased survivability.Indeed,oursubsequentstudyundercontrolledexperimentalconditionswithknown Pd inoculationdosesfoundsignicantlygreatersurvivorshipinfemales[ 16].Clearly,additionalstudiesof sexbiasesintheresponseto Pd infectionareneeded,includingthosethatgobeyondmortalityestimates andfocusonconsequencessuchasmetabolicandtnesscostsinbothmalesandfemalesthatsurvive WNS. Ethicsstatement.ThisstudywascarriedoutinstrictaccordancewiththerecommendationsintheGuidefortheCare andUseofLaboratoryAnimalsoftheNationalInstitutesofHealth.TheprotocolwasapprovedbytheInstitutional AnimalCareandUseCommitteeatBucknellUniversity(protocolnumberDMR-04).Researchwasconductedunder aPennsylvaniaGameCommissionpermittoDMR(no.70).Dataaccessibility.DataareopenlypublishedintheDryadRepositoryat: doi:10.5061/dryad.db1n0 [18].Acknowledgements.WethankScottBearer,TheNatureConservancy,BrookHines(KentuckyDepartmentofFishand WildlifeResources)andGregoryTurner(PennsylvaniaGameCommission)forassistinginanimalprocurement.We alsothankK.B.BoomerforstatisticalconsultingandMeganVodzakandChelseyMusanteforeldassistance.Authorcontributions.L.E.G.designedtheexperiment,carriedoutthestatisticalanalysisanddraftedthemanuscript; D.M.R.assistedinstudydesignandexecutionandhelpeddraftthemanuscript;S.A.B.helpedconducttheresearch andcontributedtothedraft;J.S.J.assistedwiththestatisticalanalysisandmanuscriptrevision.Allauthorsgavenal approvalforpublication.Fundingstatement.L.E.G.andS.A.B.weresupportedbyGraduateStudiesatBucknellUniversity.D.M.R.andJ.S.J.were supportedbytheWoodtigerFundandtheUSFish&WildlifeService.Competinginterests.Wehavenocompetinginterests.References.USFWS. 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brsos.royalsocietypublishing.org R.Soc.opensci. :b................................................USA tn,ftttb.( doi:.b/pnas. bt) .MeteyerCU,BucklesEL,BlehertDS,HicksAC, GreenDE,Shearn-BochslerV,ThomasNJ, GargasA,BehrMJ.tHistopathologic criteriatoconrrmwhite-nosesyndromein bats. J.Vet.Diagn.Invest. ,. (doi:.bb/fnbt ) .TurnerGG,ReederDM,ColemanJTH.A rve-yearassessmentofmortalityandgeographic spreadofwhite-nosesyndromeinNorthAmerican batsandalooktothefuture. BatRes.News b. .ReederDM etal. Frequentarousalfrom hibernationlinkedtoseverityofinfectionand mortalityinbatswithwhite-nosesyndrome. PLoS ONE b,ent.( doi:.b/journal.pone.nt) f.LangwigKE,FrickWF,BriedJT,HicksAC,KunzTH, KilpatrickAM.Sociality,density-dependence andmicroclimatesdeterminethepersistenceof populationssueringfromanovelfungaldisease, white-nosesyndrome. Ecol.Lett. ,b. (doi:./j.f-n..nt.x ) b.BoylesJG,DunbarMB,StormJJ,BrackV.b Energyavailabilityinuencesmicroclimate selectionofhibernatingbats. J.Exp.Biol. t .(doi:./jeb.bt ) n.GeiserF.Metabolicrateandbodytemperature reductionduringhibernationanddailytorpor. Annu.Rev.Physiol. ,tb.( doi:.f/ annurev.physiol.ff..) t.ThomasDW,DoraisM,BergeronJ-M.ttWinter energybudgetsandcostofarousalsforhibernating littlebrownbats, Myotislucifugus. J.Mammal. b bbt.(doi:.b/ntfb) .KunzTH,WrazenJA,BurnettCD.ttnChangesin bodymassandfatreservesinpre-hibernatinglittle brownbats(Myotislucifugus ). Ecoscience ,nb. .JonassonKA,WillisCKR.Changesinbody conditionofhibernatingbatssupportthethrifty femalehypothesisandpredictconsequencesfor populationswithwhite-nosesyndrome. PLoSONE ,ef.( doi:.b/journal.pone.f) .VerantML,BoylesJG,WaldrepJrW,WibbeltG, BlehertDS.Temperature-dependentgrowthof Geomycesdestructans,thefungusthatcausesbat white-nosesyndrome. PLoSONE b,efn. (doi:.b/journal.pone.fn ) .Brownlee-BouboulisSA,ReederDM. White-nosesyndrome-aectedlittlebrownmyotis (Myotislucifugus )increasegroomingandother activebehaviorsduringarousalsfromhibernation. J.Wildl.Dis. n ,nnt.( doi:.bnt/-) .LorchJM etal. Experimentalinfectionofbats with Geomycesdestructans causes white-nose syndrome. Nature t,bfbn.( doi:.n/ naturet) .CryanPM,MeteyerCU,BoylesJG,BlehertDS. Wingpathologyofwhite-nosesyndrome inbatssuggestslife-threateningdisruptionof physiology. BMCBiol. ,n.( doi:.nf/b-bbn-) f.JohnsonJS etal. Host,pathogen,and environmentalcharacteristicspredict white-nosesyndromemortalityincaptive littlebrownmyotis(Myotislucifugus ). PLoS ONE n,e.(doi:.b/journal.pone. ) b.StormJJ,BoylesJG.Bodytemperatureand bodymassofhibernatinglittlebrown batsMyotis lucifugus inhibernaculaaectedbywhite-nose syndrome. ActaTheriol. ,b. (doi:.b/sf--n) n.GrieneisenLE,Brownlee-BouboulisSA,JohnsonJS, ReederDM.Datafromsexandhibernaculum temperaturepredictsurvivorshipinwhite-nose syndrome(WNS)aectedlittlebrownmyotis (Myotislucifugus ). DryadRepository.( doi:.f/ dryad.dbn) on April 21, 2017 http://rsos.royalsocietypublishing.org/ Downloaded from