Table Of ContentOUPCORRECTEDPROOF–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
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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
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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
