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OUPCORRECTEDPROOF–FINAL,6/12/2011,SPi Carnivore Ecology and Conservation OUPCORRECTEDPROOF–FINAL,6/12/2011,SPi Techniques in Ecology and Conservation Series SeriesEditor:WilliamJ.Sutherland BirdEcologyandConservation:AHandbookofTechniques WilliamJ.Sutherland,IanNewton,andRhysE.Green ConservationEducationandOutreachTechniques SusanK.Jacobson,MalloryD.McDuff,andMarthaC.Monroe ForestEcologyandConservation:AHandbookofTechniques AdrianC.Newton HabitatManagementforConservation:AHandbookofTechniques MalcolmAusden ConservationandSustainableUse:AHandbookofTechniques E.J.Milner-GullandandJ.MarcusRowcliffe InvasiveSpeciesManagement:AHandbookofPrinciplesandTechniques MickN.CloutandPeterA.Williams AmphibianEcologyandConservation:AHandbookofTechniques C.KennethDodd,Jr. InsectConservation:AHandbookofApproachesandMethods MichaelJ.Samways,MelodieA.McGeoch,andTimR.New RemoteSensingforEcologyandConservation:AHandbookofTechniques NedHorning,JulieA.Robinson,EleanorJ.Sterling,WoodyTurner,andSachaSpector MarineMammalEcologyandConservation:AHandbookofTechniques IanL.Boyd,W.DonBowen,andSaraJ.Iverson CarnivoreEcologyandConservation:AHandbookofTechniques LuigiBoitaniandRogerA.Powell OUPCORRECTEDPROOF–FINAL,6/12/2011,SPi Carnivore Ecology and Conservation A Handbook of Techniques Edited by Luigi Boitani and Roger A. Powell 1 OUPCORRECTEDPROOF–FINAL,6/12/2011,SPi 3 GreatClarendonStreet,OxfordOX26DP OxfordUniversityPressisadepartmentoftheUniversityofOxford. ItfurtherstheUniversity’sobjectiveofexcellenceinresearch,scholarship, andeducationbypublishingworldwidein Oxford NewYork Auckland CapeTown DaresSalaam HongKong Karachi KualaLumpur Madrid Melbourne MexicoCity Nairobi NewDelhi Shanghai Taipei Toronto Withofficesin Argentina Austria Brazil Chile CzechRepublic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore SouthKorea Switzerland Thailand Turkey Ukraine Vietnam OxfordisaregisteredtrademarkofOxfordUniversityPress intheUKandincertainothercountries PublishedintheUnitedStates byOxfordUniversityPressInc.,NewYork #OxfordUniversityPress2012 Themoralrightsoftheauthorhavebeenasserted DatabaserightOxfordUniversityPress(maker) Firstpublished2012 Allrightsreserved.Nopartofthispublicationmaybereproduced, storedinaretrievalsystem,ortransmitted,inanyformorbyanymeans, withoutthepriorpermissioninwritingofOxfordUniversityPress, orasexpresslypermittedbylaw,orundertermsagreedwiththeappropriate reprographicsrightsorganization.Enquiriesconcerningreproduction outsidethescopeoftheaboveshouldbesenttotheRightsDepartment, OxfordUniversityPress,attheaddressabove Youmustnotcirculatethisbookinanyotherbindingorcover andyoumustimposethesameconditiononanyacquirer BritishLibraryCataloguinginPublicationData Dataavailable LibraryofCongressCataloginginPublicationData LibraryofCongressControlNumber:2011938082 TypesetbySPIPublisherServices,Pondicherry,India Printedandboundby CPIGroup(UK)Ltd,Croydon,CR04YY ISBN: 978–0–19–955852–0(Hbk.) 978–0–19–955853–7(Pbk.) 1 3 5 7 9 10 8 6 4 2 OUPCORRECTEDPROOF–FINAL,6/12/2011,SPi Foreword Animals that must hunt and kill for at least part of their living are inherently interesting to many people. Perhaps that is because humans evolved to make our living that way as well, and carnivores often compete with us to this very day. Wolves, bears, lions, tigers, leopards, lynx, mink, weasels, and foxes, and a wide variety of their relatives, have long grabbed the human imagination. In any case, carnivorescompriseaverysignificantcontingentoftheworld’swildlife,andmany books havebeen writtenabout them. This book is distinct from its predecessors primarily through its emphasis on techniquesfordealingwithcarnivores:howtosamplethem,capturethemforstudy, handlethem,monitorthem,andevenhowtohelpminimizetheircompetitionwith us.Itisaveryhelpfulbookthatfillsanimportantnicheandcomesattherighttime. In many parts of the world carnivores are persecuted, while in other parts they arebeingrestored.Thussocietiesremaininterestedincarnivoresforonereasonor another, and science serves society’s interest through numerous carnivore studies. The authors of this book’s chapters have conducted a significant proportion of thosestudies formany years, andthe editorsforeven longer. Botheditorsarewellqualifiedtoproducethisbook,havingstudiedandworked withcarnivoresandtheirconservation fordecades.Ihadthegreatopportunityof partneringwithLuigiBoitaniin1974,earlyinhiscareer,whenwespentamonthin Italy’sAbruzzoMountainslive-trapping,radio-collaring,andtrackingwolves.Ihad presented my paper “Current Techniques in the Study of Elusive Wilderness Carnivores” at the Eleventh International Congress of Game Biologists in Stock- holm in September 1973. It covered my experiences live-trapping and radio- trackingwolves,fishers,martens,andlynxesaswellasaliteraturereviewofcurrent techniquesusedtostudyothercarnivores.Iliketothinkofthatpaperasagermthat helpedspawnthepresentbook.LuigiattendedtheStockholmmeeting,soughtto apply my techniques with wolves in Italy, and asked me to join him there to get started.Ieagerlyagreed.LittledidIrealizethenthat40yearslater,LuigiandRoger Powellwoulddevoteawholebooktotechniquesforstudyingcarnivores. During the same general period when I met Luigi, I also met Roger Powell. Roger hadjoinedmyresearch team as a summerintern on awolf–deer projectin theSuperiorNationalForestofMinnesota,wherewehadalsobeenradio-tracking lynx,martens,andfishersontheside.Thedutiesclearlyagreedwithhim,forafew OUPCORRECTEDPROOF–FINAL,6/12/2011,SPi vi | Foreword yearslaterhebeganhisowncarnivorestudy,thisoneinvolvingfishers.Thatstudy became hisdissertationtopic, andI becameone ofhisadvisors. That was all long ago, and the field has advanced greatly and blossomed. Now instead of merely locating an animal via telemetry (a feat in itself years ago), one searches the profuse literature, decides on study objectives, carefully plans the study’s design, and chooses from any of the many high-tech radio-collars on the commercialmarketthat will bestserve the objectives. However, dealing with the most appropriate technology to study carnivores is onlyasmallpartofcarnivoreinvestigationsnow.Thedatacurrentlyobtainablehas opened many new carnivore research vistas, and Boitani and Powell and their collaborators have assembled a set of chapters that nicely address that array. An early chapter on carnivore surveys, for example, is basic, for such surveys are of special importance, both spatially and temporally. In some areas and with some species, just obtaining a general idea of numbers and distribution can be very important. Mapping such distributions plays a major role in these studies, and non-invasive sampling is particularly valuable, especially with endangered or rare species andininaccessible areas. These subjects are well coveredinthis book. In some areas of the world and with certain carnivores, detailed counts are required annually. Sometimes with such counts it is valuable to estimate various demographic parameters, and radio-telemetry often facilitates those estimates. To collarcarnivores,itisnecessarytocaptureandhandlethem,allowingconsiderable amounts of valuable data to be collected at that time. Once a carnivore is radio- collared, data can be obtained about its movements, activity, home range or territory,anddispersal.Oftendataaboutthecreature’spredationandfoodhabits canalsobecollected,aswellasinformationaboutitsreproductivebehavior.Several chapters ofthisbook deal withthesesubjects. A subsidiary type of information, not directly related to a collared carnivore’s movements, involves cause-specific mortality, including that from intraspecific strife and diseases. Learning all this basic ecological, physiological, and behavioral information then greatly aids in deriving mitigation measures for minimizing depredation on livestock and other conflicts with humans, as well as facilitating methods of restoring carnivores, monitoring the results, and furthering conserva- tion efforts.Addressing thoseissues furtherroundsout thisfinecompendium. Thus all in all, this book, edited by Luigi Boitani and Roger Powell, will be of greatusenotonlytocarnivoreresearchers,butalsotowildlifebiologiststhrough- out the world who deal with carnivores, and it should stand as a milestone in the carnivore-ecology andtechniquesliterature formany years tocome. L.DavidMech USGeological SurveyandUniversity ofMinnesota, USA OUPCORRECTEDPROOF–FINAL,6/12/2011,SPi Contents Listofcontributors xv 1. Introduction:researchandconservationofcarnivores 1 LuigiBoitaniandRogerA.Powell 2. Designingcarnivoresurveys 8 LuigiBoitani,PaoloCiucci,andAlessioMortelliti 2.1 Challengesofsurveyingcarnivores 10 2.2 Planningasurvey 10 2.2.1 Fundamentalsofsurveydesign:establishinggoalsandobjectives 11 2.2.2 Fundamentalsofsurveydesign:carnivoresurveydata 12 2.2.3 Fundamentalsofsurveydesign:samplingdesign,methods, andprotocols 13 2.2.4 Fundamentalsofsurveydesign:statisticallyformalizing surveyobjectives 14 2.3 Dealingwithfalseabsence 15 2.3.1 Thefastgrowingfamilyofoccupancymodels 16 2.3.2 Assumptionsofoccupancymodels:theimportanceofa prioriplanning 16 2.3.3 Somepracticalissues 18 2.3.4 Designinganoccupancystudy 18 2.4 Keyissuesfordevelopingasurveydesign 19 2.4.1 Targetpopulationandspatialextentofthesurvey 19 2.4.2 Attributetomeasure 20 2.4.3 Samplingdesign 21 2.4.4 Samplingeffort 25 2.4.5 Tacklingsystemvariability:measuresofprecisionand theirmeaning 26 2.4.6 Fieldmethods 29 3. Mindthemap:tripsandpitfallsinmakingandreading mapsofcarnivoredistribution 31 CarloRondininiandLuigiBoitani 3.1 Mapsbasedonexpertknowledge 32 3.1.1 Geographicrangemaps 32 3.1.2 Deductivehabitatsuitabilitymodels(HSM) 33 OUPCORRECTEDPROOF–FINAL,6/12/2011,SPi viii | Contents 3.2 Mapsbasedonspecies’occurrencesurveys 34 3.2.1 Typesofdata 34 3.2.2 Biologicalsignificanceandtimerelevance 36 3.2.3 Extrapolatingpointstomapthedistributionofapopulation 38 3.2.4 InductiveHSM 42 3.2.5 CaveatsandlimitationsofdeductiveandinductiveHSM 45 4. Noninvasivesamplingforcarnivores 47 MarcellaJ.Kelly,JulieBetsch,ClaudiaWultsch,BernardoMesa,andL.ScottMills 4.1 Methodsofnoninvasivesampling 48 4.1.1 Signsurveys 48 4.1.2 Geneticsampling 49 4.1.3 Camera-trapsampling 54 4.1.4 Endocrine/hormonesampling 55 4.2 Recenttoolsandadvancesinnoninvasivesampling 56 4.2.1 NoninvasiveDNAtechniques 56 4.2.2 UsingnoninvasiveDNAdata 59 4.2.3 Datacollection,handling,andanalyseswithremotecameras 62 4.2.4 Datacollection,handling,andanalysesforendocrinestudies 65 4.3 Combiningnoninvasiveandtraditionalapproaches 67 4.3.1 Comparativeapproachesamongnoninvasivetechniques 67 4.3.2 Combiningtraditionalwithnoninvasiveapproaches 68 4.3.3 Dataqualityandintegrityinnoninvasivesurveys 69 5. Humaneandefficientcaptureandhandlingmethodsforcarnivores 70 GilbertProulx,MarcR.L.Cattet,andRogerA.Powell 5.1 Mechanicalcapturemethods 72 5.1.1 Trapsandsets 72 5.1.2 Trappingefficiency 73 5.1.3 Humaneness 74 5.1.4 Trapsandsetsforspecificcarnivores 75 5.2 Useofdrugsforcaptureandrestraintofcarnivores 78 5.2.1 Drugaccess,storage,andhandling 78 5.2.2 Selectionofdrugsforuseincarnivores 79 5.2.3 Methodstoadministerdrugs 81 5.2.4 Thevalueofknowledgeandexperience 81 5.3 Identification,prevention,andtreatmentofmedicalemergencies associatedwithcapture 84 5.3.1 Homeostasis,stress,distress,andtreatmentofmedicalemergencies 84 5.3.2 Necropsy 86 5.4 Euthanasia 86 5.5 Restrainingandmarkingtechniques 88 OUPCORRECTEDPROOF–FINAL,6/12/2011,SPi Contents | ix 5.6 Designingeffectivetrappingprogramsforcarnivores 89 5.7 Animalwelfare 89 Appendices 92 6. Carnivoresinhand 130 KerryR.Foresman 6.1 Aging 130 6.2 Standardbodymeasurements 132 6.2.1 Bodymass 133 6.2.2 Lengthmeasurements 133 6.2.3 Additionalbodymeasurements 136 6.2.4 Additionalmeasurements,sometoestimateage 137 6.2.5 Footpadpatterns 137 6.3 Tootheruptionandmeasurements 137 6.3.1 Tootheruption,wear,andage 138 6.3.2 Pulpcavitymeasurementsandage 139 6.3.3 Cementumannuliandage 140 6.4 Skullandskeletalmeasurements 142 6.4.1 Skullmeasurements 142 6.4.2 Skullfusionandage 144 6.4.3 Skeletalmorphologyandage 144 6.4.4 Eyelensandage 144 6.5 Pelageandage 145 6.6 Sexandreproduction 145 6.7 Injuries 148 6.8 Physiologicalparameters 148 6.8.1 Blood 149 6.8.2 Tissuesamples 149 6.8.3 Othersamples 150 6.9 Bioelectricalimpedance 150 6.10 Asymmetry 151 7. Radio-telemetryequipmentandapplicationsforcarnivores 152 MarkR.FullerandToddK.Fuller 7.1 Generalbackground 152 7.2 Basictelemetrysystem 156 7.3 Radio-trackingfieldprocedures 163 7.4 Satellitetelemetrysystems 163 7.5 Radio-telemetryapplicationsforcarnivores 166

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