american bison...american bison status survey and conservation guidelines 2010 edited by c. cormack...

InternatIonal UnIon for ConservatIon of natUre

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American Bison Status Survey and Conservation Guidelines 2010 Edited by C. Cormack Gates, Curtis H. Freese, Peter J.P. Gogan, and Mandy Kotzman

IUCn/ssCamerican BisonspecialistGroup

Am

erican Bison S

tatus Survey and

Conservation G

uidelines 2010

Bison_Cover_SPREAD.indd 1 2/10/10 8:27:05 AM

iv AmericanBison:StatusSurveyandConservationGuidelines2010

Thedesignationofgeographicalentitiesinthisreport,andthepresentationofthematerial,donotimplytheexpressionofanyopinion

whatsoeveronthepartofIUCNconcerningthelegalstatusofanycountry,territory,orarea,orofitsauthorities,orconcerningthe

delimitationofitsfrontiersorboundaries.

TheviewsexpressedinthispublicationdonotnecessarilyreflectthoseofIUCN.

Publishedby: IUCN,Gland,Switzerland

Copyright: ©2010InternationalUnionforConservationofNatureandNaturalResources

Reproductionofthispublicationforeducationalorothernon-commercialpurposesis

authorizedwithoutpriorwrittenpermissionfromthecopyrightholderprovidedthesourceisfully

acknowledged.

Reproductionofthispublicationforresaleorothercommercialpurposesisprohibitedwithout

priorwrittenpermissionofthecopyrightholder.

Citation: Gates,C.C.,Freese,C.H.,Gogan,P.J.P.andKotzman,M.(eds.andcomps.)(2010).American

Bison: Status Survey and Conservation Guidelines 2010.Gland,Switzerland:IUCN.

ISBN: 978-2-8317-1149-2

Coverdesignby: C.CormackGates

Frontcoverphoto: Plainsbisonbulltendingacow(photoDianeHargreaves/Hargreavesphoto.com)

Backcoverphoto: Woodbisoncowwithcalf(photoDougLindstrand)

Layoutby: AmyKelley

Producedby: IUCN-SSC-AmericanBisonSpecialistGroup

Printedby: InstyPrints,Bozeman,Montana

Availablefrom: ThewebsitesofIUCNPublicationsServices,IUCN/SpeciesSurvivalCommission,WorldWildlife

Fund,AmericanBisonSociety,andWildlifeConservationSociety

RevisedJune2011.

AmericanBison:StatusSurveyandConservationGuidelines2010v

TableofContents

Acknowledgements ..................................................................................................................................xi

Authors, contributors and their affiliations .............................................................................................xii

Acronyms ................................................................................................................................................xiii

Executive Summary .................................................................................................................................xv

ChAptEr 1 Introduction: the Context ................................................................................................... 1

1.1 the Species Survival Commission and the American Bison Specialist Group ...................... 1

1.2 Context .................................................................................................................................. 1

1.3 Current Challenges for Conservation and Ecological restoration of Bison as Wildlife ......... 2

1.4 Large Wild populations ........................................................................................................... 2

1.5 Conserving the Wild Character and Genome of Bison............................................................ 3

1.6 reportable Diseases ............................................................................................................... 4

1.7 purpose of this Document ...................................................................................................... 4

ChAptEr 2 history of Bison in North America ...................................................................................... 5

2.1 palaeobiology and phylogeny ............................................................................................... 5

2.2 Original range ........................................................................................................................ 6

2.3 Abundance .............................................................................................................................. 7

2.4 Extirpation .............................................................................................................................. 8

2.5 Early recovery ........................................................................................................................ 8

2.6 Cultural Significance .............................................................................................................. 9

ChAptEr 3 taxonomy and Nomenclature ............................................................................................13

3.1 An historical Misnomer: Bison vs. Buffalo ............................................................................13

3.2 Genus: Bos vs. Bison .............................................................................................................13

3.3 Subspecies ...........................................................................................................................15

ChAptEr 4 Genetics ............................................................................................................................19

4.1 reduction of Genetic Diversity ..............................................................................................19

4.2 hybridisation .........................................................................................................................21

4.2.1 plains bison x wood bison ..........................................................................................21

4.2.2 Domestic cattle x bison ..............................................................................................22

4.3 Domestication ........................................................................................................................24

ChAptEr 5 reportable or Notifiable Diseases ....................................................................................27

5.1 Diseases of Conservation Concern .......................................................................................28

5.1.1 Anaplasmosis..............................................................................................................28

5.1.2 Anthrax .......................................................................................................................28

5.1.3 Bluetongue ..................................................................................................................29

vi AmericanBison:StatusSurveyandConservationGuidelines2010

5.1.4 Bovine spongiform encephalopathy ...........................................................................30

5.1.5 Bovine brucellosis ......................................................................................................30

5.1.6 Bovine tuberculosis ....................................................................................................31

5.1.7 Bovine viral diarrhoea .................................................................................................31

5.1.8 Johne’s disease ..........................................................................................................32

5.1.9 Malignant catarrhal fever (sheep associated) ............................................................32

5.2 Episodes of reportable Diseases in plains Bison .................................................................33

5.2.1 Yellowstone National park ..........................................................................................33

5.2.2 Grand teton National park/National Elk refuge (Jackson herd) ................................34

5.3 An Occurrence of reportable Diseases in Wood Bison .........................................................35

5.4 Disease Management in perspective .....................................................................................36

ChAptEr 6 General Biology, Ecology and Demographics ....................................................................39

6.1 General Biology .....................................................................................................................39

6.1.1 physiology ..................................................................................................................39

6.1.1.1 Metabolism ..................................................................................................39

6.1.1.2 Growth .........................................................................................................39

6.1.2 Behaviour ....................................................................................................................40

6.1.2.1 Social structure ..........................................................................................40

6.1.2.2 reproductive behaviour ...............................................................................40

6.1.2.3 Cow-calf behaviour .....................................................................................40

6.1.2.4 horning and wallowing ................................................................................41

6.1.2.5 Movements ..................................................................................................41

6.2 Ecology ..................................................................................................................................42

6.2.1 plains bison ................................................................................................................42

6.2.1.1 Ecological role .............................................................................................42

6.2.1.2 Contemporary habitat use, nutrition, and foraging .....................................43

6.2.1.2.1 Northern mixed grasslands ..............................................................45

6.2.1.2.2 Central shortgrass prairie ................................................................45

6.2.1.2.3 tall grasslands prairie and southern shortgrass prairie ..................45

6.2.1.2.4 Northern fescue grasslands ............................................................45

6.2.1.2.5 rocky Mountain forest .....................................................................45

6.2.1.2.6 Northern forests ...............................................................................46

6.2.1.2.7 Arctic lowland taiga ..........................................................................46

6.2.1.3 habitat and dietary overlap .............................................................................46

6.2.2 Wood bison .................................................................................................................46

6.2.2.1 Original distribution and ecoregions occupied ...........................................46

6.2.2.2 Contemporary habitat relationships, nutrition, and foraging ......................47

6.2.2.2.1 Northern forests ...............................................................................47

6.2.2.2.2 Subarctic boreal forests ..................................................................47

6.2.2.3 habitat and dietary overlap ........................................................................47

AmericanBison:StatusSurveyandConservationGuidelines2010vii

6.3 Demographics ........................................................................................................................47

6.3.1 population structure ..................................................................................................48

6.3.2 reproduction ..............................................................................................................49

6.3.3 Mortality factors and survival ....................................................................................49

6.3.4 population growth rates ............................................................................................53

ChAptEr 7 Numerical and Geographic Status .....................................................................................55

7.1 Introduction ...........................................................................................................................55

7.2 Numerical Status ..................................................................................................................56

7.3 Geographic Status .................................................................................................................57

7.4 population Size Distribution ..................................................................................................59

7.5 Mate Competition .................................................................................................................60

7.6 presence of Wolves ...............................................................................................................60

7.7 presence of reportable Diseases ..........................................................................................60

7.8 Cattle Gene Introgression ......................................................................................................61

7.9 Conclusions ...........................................................................................................................61

ChAptEr 8 Legal Status, policy Issues and Listings ...........................................................................63

8.1 Introduction ...........................................................................................................................63

8.2 history of protection and Conservation .........................................................................63

8.2.1 Early legal and policy efforts by governments to protect plains and wood bison ........... 63

8.2.1.1 Early policy development in the United States ............................................63

8.2.1.2 Early policy development in Canada............................................................63

8.2.1.3 policy development in Mexico .....................................................................64

8.2.2 plains bison conservation by the private sector ........................................................64

8.2.3 Conservation efforts by tribes and First Nations ...............................................................................64

8.3 Important policy and regulatory Considerations .................................................................65

8.3.1 Legal status and listings of bison ...............................................................................65

8.3.1.1 International and global status ....................................................................65

8.3.1.2 Status in North America ..............................................................................65

8.3.2 Disease status ...........................................................................................................73

8.4 Legal and policy Obstacles hindering Conservation of Bison...............................................75

8.4.1.1.1 Confusing legal classification and status .........................................75

8.4.1.1.2 historical management policies .......................................................75

8.4.1.1.3 Complex partnerships needed to manage large landscapes ............75

8.4.1.1.4 Defining the social and economic value of wild bison .....................76

8.4.1.1.5 Coordination of policies, rules, and regulations by government ......76

8.4.1.1.6 Agricultural conflicts among mixed land ownership ........................76

8.5 Overcoming Obstacles to the Ecological restoration of Bison .............................................77

8.5.1 Disease management considerations .........................................................................77

8.5.2 Legal status and policy considerations ......................................................................77

viii AmericanBison:StatusSurveyandConservationGuidelines2010

8.5.2.1 role of the non-governmental organisations ..............................................77

8.5.2.2 State/provincial and federal governance .....................................................78

8.5.2.3 the private sector........................................................................................78

8.5.2.4 Indigenous peoples .....................................................................................78

8.5.2.5 Local communities and economies .............................................................79

8.5.3 Coordination of agency missions, goals, regulations, and policies affecting bison conservation and restoration ........................................80

8.5.4 recommendations ......................................................................................................80

8.5.5 recent initiatives to conserve and restore bison ..............................................................................81

8.5.5.1 United States ...............................................................................................81

8.5.5.2 Canada .........................................................................................................82

8.5.5.3 Mexico .........................................................................................................83

8.5.5.4 Non-governmental organisations ................................................................83

8.5.5.5 tribal initiatives ...........................................................................................84

ChAptEr 9 Conservation Guidelines for population, Genetic, and Disease Management .......................85

9.1 Introduction and principles ...................................................................................................85

9.2 Guidelines for population and Genetic Management ............................................................86

9.2.1 Guidelines that apply to most conservation herds .....................................................87

9.2.2 herd-level population and genetic management ........................................................88

9.2.2.1 Soft release procedures ..............................................................................88

9.2.3 Establishing a new herd ..............................................................................................88

9.2.4 Maintaining or manipulating existing herd size ..........................................................89

9.2.5 transferring bison between herds ..............................................................................90

9.2.6 recovering small or threatened herds ........................................................................91

9.2.7 recovering herds from germplasm introgression .....................................................92

9.2.8 herd size reduction .....................................................................................................92

9.3 Behaviour: Mating System, Social Structure, and Movements .............................................92

9.3.1 Social structure and spacing ......................................................................................93

9.3.2 Foraging and movements ...........................................................................................93

9.3.3 Mating behaviour ........................................................................................................94

9.3.4 Limiting factors and natural selection ........................................................................94

9.4 habitat and Biodiversity Management ..................................................................................94

9.5 Disease Guidelines: Considerations for Infected and Uninfected herds .............................95

9.5.1 prevention ...................................................................................................................96

9.5.2 Surveillance ................................................................................................................96

9.5.3 Management ...............................................................................................................97

9.5.4 research .....................................................................................................................97

9.5.5 Stakeholder involvement ............................................................................................97

AmericanBison:StatusSurveyandConservationGuidelines2010ix

9.6 Active Management: handling, herding, Infrastructure .......................................................98

9.6.1 handling ......................................................................................................................98

9.6.2 Fencing .......................................................................................................................99

9.6.3 Corrals, pens, and chutes ...........................................................................................99

9.7 Modelling to Assess Bison populations and habitat ..........................................................100

9.7.1 Guidelines for using computer simulations .............................................................100

9.8 Conclusions .........................................................................................................................101

ChAptEr 10 Guidelines for Ecological restoration of Bison .............................................................103

10.1 Introduction .........................................................................................................................103

10.2 Ecological restoration ........................................................................................................104

10.2.1 Geographic potential for ecological restoration ..................................................104

10.2.2 principles for ecological restoration applicable to bison .....................................105

10.3 the “Ecosystem Approach” for Designing Ecological restoration of Bison .......................107

10.3.1 Defining the biological landscape and objectives ................................................107

10.3.2 Defining the social landscape, the main stakeholders, and cultivating partnerships ................................................................................107

10.4 Guidelines for planning and Implementing Ecological restoration projects for Bison .......109

10.4.1.1 Feasibility assessment ..............................................................................109

10.4.1.2 Suitable release stock ...............................................................................110

10.4.1.3 preparation and release ............................................................................110

10.4.1.4 Socio-economic and legal requirements ...................................................111

10.4.1.5 Monitoring, evaluation, and adaptation .....................................................111

10.5 Summary ..............................................................................................................................112

Literature Cited .....................................................................................................................................113

AppENDIx A North American conservation herds of bison and their managing authorities ................131

x AmericanBison:StatusSurveyandConservationGuidelines2010

AmericanBison:StatusSurveyandConservationGuidelines2010xi

Acknowledgements

Thismanuscriptistheproductofmorethanthreeyearsof

cooperativeeffortbynumerouscontributors,manyofwhom

arelistedasauthors.Theirknowledgeandparticularlytheir

persistencewereinstrumentalinseeingthismajorundertaking

throughtosuccessfulcompletion.Theeditorsexpress

theirappreciationtoJoeTruettwiththeTurnerEndangered

SpeciesFundforhisadviceoncompilingthisdocument.We

acknowledgethesupportofinstitutionsandorganisationsthat

authorisedmembersoftheBisonSpecialistGroupandothers

tocontributetotheproject.Theyincludethefollowinginno

particularorderofpriority:U.S.NationalParkService;U.S.

FishandWildlifeService;U.S.GeologicalSurveyBiological

ResourcesDivision;ParksCanadaAgency;CanadianWildlife

Service;DepartmentofNationalDefenseinCanada;Comisión

NacionaldeAreasNaturalesProtegidas,Mexico;Universidad

NacionalAutónomadeMéxico,InstitutodeEcología;Stateof

MontanaFish,WildlifeandParks;SouthDakotaGameFishand

Parks;AlaskaDepartmentofFishandGame;YukonDepartment

oftheEnvironment;NorthwestTerritoriesEnvironmentand

NaturalResources;NorthernGreatPlainsOfficeoftheWorld

WildlifeFund;WildlifeConservationSociety;TheAmericanBison

Society;TheNatureConservancy;TurnerEndangeredSpecies

Fund;TurnerEnterprises;Inter-TribalBisonCooperative;Council

ofAthabascanTribalGovernments;FacultyofEnvironmental

Design,andtheDepartmentofArchaeologyintheFaculty

ofSocialSciencesattheUniversityofCalgary;Collegeof

VeterinaryMedicineandBiomedicalSciences,TexasA&M

University;DepartmentofAnthropology,UniversityofAlaska,

Fairbanks;DepartmentofZoology,UniversityofOklahoma;the

CanadianBisonAssociation;andtheNationalBisonAssociation.

Wealsowishtoacknowledgelogisticalsupportprovidedby

VermejoParkRanch,andparticularlythegenerosityofMarv

JensenandTedTurner,whoco-hostedameetingoftheBison

SpecialistGroupin2005toorganizethewritingproject.The

WildlifeConservationSocietysubsequentlyhostedtwomeetings

todevelopavisionforbisonrestorationinNorthAmericain

whichmanymembersoftheAmericanBisonSpecialistGroup

participated.Theseworkshopswereinstrumentalinbuilding

workingrelationships,sharingknowledge,anddevelopinga

senseofmission,whichcontributedtotheproject’ssuccess.

TheU.S.GeologicalSurveyandWildlifeConservation

Societyprovidedsupportfortechnicalediting,formattingand

compilationofthedocument.

Finally,theWorldWildlifeFundNorthernGreatPlains

Program—particularlystaffmembersSteveForrestandPeder

Groseth—wasinstrumentalindevelopingaframeworkforbison

conservation,addingcontentandguidingtheproductionof

thisdocumentbyprovidingfinancialandtechnicalsupportfor

thefirstVermejomeetingandsubsequentmeetingsandfor

technicalediting,layout,andpublicationofthefinaldocument.

xii AmericanBison:StatusSurveyandConservationGuidelines2010

Authors,contributorsandtheiraffiliations

Aune,Keith WildlifeConservationSociety,Bozeman,Montana,USA

Berger.Joel WildlifeConservationSociety,Bozeman,Montana,USA

Boyd,DelaneyP. DepartmentofNationalDefence,CanadianForcesBaseSuffield,MedicineHat,Alberta,Canada

Derr,JamesN. DepartmentofVeterinaryPathobiology,TexasA&MUniversity,CollegeStation,Texas,USA

Elkin,BrettT. GovernmentoftheNorthwestTerritories,DepartmentofEnvironmentandNaturalResources,

Yellowknife,NorthwestTerritories,Canada

Ellison,Kevin WildlifeConservationSociety,Bozeman,Montana,USA

Freese,CurtisH. Bozeman,Montana,USA

Gates,C.Cormack FacultyofEnvironmentalDesign,UniversityofCalgary,Calgary,Canada

Gerlach,S.Craig DepartmentofCross-CulturalStudiesandResilienceandAdaptationProgram,UniversityofFairbanks,

Alaska,USA

Gogan,PeterJ.P. UnitedStatesGeologicalSurvey,NorthernRockyMountainScienceCenter,Bozeman,Montana,USA

Gross,JohnE. U.S.NationalParkService,FortCollins,Colorado,USA

Halbert,NatalieD. DepartmentofVeterinaryPathobiology,TexasA&MUniversity,CollegeStation,Texas,USA

Hugh-Jones,Martin DepartmentofEnvironmentalSciences,SchooloftheCoastandEnvironment,LouisianaState

University,BatonRouge,Louisiana,USA

Hunter,David TurnerEndangeredSpeciesFund,Bozeman,Montana,USA

Joly,DamienO. WildlifeConservationSociety,Nanaimo,BritishColumbia,Canada

Kotzman,Mandy CreativePursuitsLLC,LaPort,Colorado,USA

Kunkel,Kyran WorldWildlifeFund,Bozeman,Montana,USA

Lammers,DuaneJ. RapidCity,SouthDakota,USA

Larter,NicholasC. DepartmentofEnvironmentandNaturalResources,GovernmentoftheNorthwestTerritories,Fort

Simpson,Canada

Licht,Daniel U.S.NationalParkService,RapidCity,SouthDakota,USA

List,Rurik InstitutodeEcología,UniversidadNacionalAutónomadeMéxico,MexicoCity,Mexico

Nishi,John ALCESGroup,Calgary,Alberta,Canada

Oetelaar,GeraldA DepartmentofArchaeology,UniversityofCalgary,Calgary,Alberta,Canada

Paulson,RobertL. TheNatureConservancy,RapidCity,SouthDakota,USA

Potter,BenA. DepartmentofAnthropology,UniversityofAlaska,Fairbanks,USA

Powers,Jenny U.S.NationalParkService,FortCollins,Colorado,USA

Shaw,JamesH. NaturalResourceEcologyandManagement,OklahomaStateUniversity,Stillwater,Oklahoma,USA

Stephenson,RobertO. AlaskaDepartmentofFishandGame,Fairbanks,Alaska,USA

Truett,Joe TurnerEndangeredSpeciesFund,Glenwood,NewMexico,USA

Wallen,Rick U.S.NationalParkService,YellowstoneNationalPark,Mammoth,Wyoming,USA

Wild,Margaret U.S.NationalParkService,FortCollins,Colorado,USA

Wilson,GregoryA. CanadianWildlifeService,Edmonton,Alberta,Canada

AmericanBison:StatusSurveyandConservationGuidelines2010xiii

Acronyms

ABS AmericanBisonSociety

ABSG AmericanBisonSpecialistGroup,adivisionoftheIUCNBSG

ADFG AlaskaDepartmentofFishandGame

AGFD ArizonaGameandFishDepartment

ALCES® ALandscapeCumulativeEffectsSimulator,FOREMTechnologies

ANpp Herbaceousabovegroundnetprimaryproductivity

ApF AmericanPrairieFoundation

ApFrAN AnimalPlantandFoodRiskAssessmentNetwork,Canada

AphIS U.S.DepartmentofAgricultureAnimalandPlantHealthInspectionService

BLU Bluetongue

BNp BadlandsNationalPark,SouthDakota

BrCp BisonResearchandContainmentProgram,NorthwestTerritories

BSE Bovinespongiformencephalopathy

BSG IUCNBisonSpecialistGroup

BtB Bovinetuberculosis

BVD Bovineviraldiarrhoea

CAMp ConservationActionManagementPlanprocess,IUCNCaptiveBreedingSpecialistGroup

CAtG CouncilofAthabascanTribalGovernments,Alaska

CBA CanadianBisonAssociation

CBD InternationalConventiononBiologicalDiversity

CBSG IUCN/SSCConservationBreedingSpecialistGroup

CDOJ CanadianDepartmentofJustice

CFIA CanadianFoodInspectionAgency

CItES ConventiononInternationalTradeinEndangeredSpeciesofWildFaunaandFlora

CMN CanadianMuseumofNature

CONANp ComisionNacionalDeAreasNaturalesProtegidas,Mexico

COSEWIC CommitteeontheStatusofEndangeredWildlifeinCanada

CSp CusterStatePark,SouthDakota

CWS CanadianWildlifeService

CWCS ComprehensiveWildlifeConservationStrategy

DNDC TheDepartmentofNationalDefence,Canada

EhD Epizootichemorrhagicdisease

EINp ElkIslandNationalPark,Alberta

EIS EnvironmentalImpactStatement

ESA U.S.EndangeredSpeciesAct

ESU Evolutionarilysignificantunit

FAD ForeignAnimalDisease

FEArp FederalEnvironmentalAssessmentReviewPanel,Canada

FMD Foot-and-mouthdisease,orheartwater

FNNWr FortNiobraraNationalWildlifeRefuge,Nebraska

GEU Geminateevolutionaryunit

GtNp GrandTetonNationalPark,Wyoming

GWBE GreaterWoodBuffaloEcosystem,Canada

GWBNp GreaterWoodBuffaloNationalPark,Canada

GYA GreaterYellowstoneArea

hMSp HenryMountainsStatePark,Utah

hOAA HealthofAnimalsAct,Canada

InVESt IntegratedValuationofEcosystemServicesandTradeoffs

ItBC IntertribalBisonCooperative

IUCN SSC IUCNSpeciesSurvivalCommission

IUCN SUSG IUCNSustainableUseSpecialistGroup

JD Johne’sdisease

MBS MackenzieBisonSanctuary,NorthwestTerritories

MCA MontanaCodeAnnotated

MCF Malignantcatarrhalfever

MDOL StateofMontanaDepartmentofLivestock

MFWp StateofMontanaDepartmentofFish,WildlifeandParks

MLVA Multiplelocus,variablenumber,tandemrepeatanalysis

MtDNA Mitochondrialdeoxyribonucleicacid

N Populationsize

xiv AmericanBison:StatusSurveyandConservationGuidelines2010

NBA NationalBisonAssociation,U.S.A

NBMB NorthernBuffaloManagementBoard,Canada

NBr NationalBisonRange,Montana

NCC NatureConservancyofCanada

Ne Effectivepopulationsize

NEp Nonessentialexperimentalpopulation

NEpA NationalEnvironmentalPolicyAct,U.S.A

NEr NationalElkRefuge,Wyoming

NGO Non-governmentalorganisation

NpS U.S.NationalParkService

NrCS NaturalResourceConservationService,U.S.A

NWt NorthwestTerritories,Canada

NtENr NorthwestTerritoriesEnvironmentandNaturalResources

OIE WorldOrganizationforAnimalHealth

pANp PrinceAlbertNationalPark,Saskatchewan

pCA ParksCanadaAgency

pES Pay-for-EnvironmentalServices

phVA PopulationandHabitatViabilityAssessment

ppAs Privateprotectedareas

pVA Populationviabilityanalysis

rAC ResearchAdvisoryCommitteeforbisondiseaseresearchinWBNP

rDr ReportableDiseasesRegulations

r̂ Observedexponentialrateofpopulationincrease

rm Maximumexponentialrateofpopulationincrease

rMEF RockyMountainElkFoundation

SAGArpA SecretaryofAgriculture,LivestockProduction,RuralDevelopment,FisheryandFood,Mexico

SCBD SecretariatoftheConventiononBiologicalDiversity

SDGFp SouthDakotaGame,FishandParks

SEMArNAt SecretaríadeMedioAmbienteyRecursosNaturales,México

SENASICA ServicioNacionaldeSanidad,InocuidadyCalidadAgroalimentaria,Mexico

SErI SocietyforEcologicalRestorationInternational

SErS SocietyforEcologicalRestorationScience

ShNGp Sully’sHillNationalGamePreserve,NorthDakota

SrL SlaveRiverLowlands,NorthwestTerritories,Canada

SNMNh SmithsonianNationalMuseumofNaturalHistory

SWAp StateWildlifeActionPlan(namevariesbystate)

tB Tuberculosis

tGpp TallgrassPrairiePreserve,Oklahoma

tNC TheNatureConservancy

trNp TheodoreRooseveltNationalPark,NorthDakota

tSE Transmissiblespongiformencephalopathies

tESF TurnerEndangeredSpeciesFund

USNArA U.S.NationalArchivesandRecordsAdministration

USDA U.S.DepartmentofAgriculture

USDOI U.S.DepartmentoftheInterior

USFS U.S.ForestService

USFWS U.S.FishandWildlifeService

USGSBrD U.S.GeologicalSurveyBiologicalResourcesDivision

VJDhSp VoluntaryJohne’sDiseaseHerdStatusProgramme(forcattle)

WBNp WoodBuffaloNationalPark,AlbertaandNorthwestTerritories

WBp WainwrightBuffaloPark,Alberta

WCNp WindCaveNationalPark,SouthDakota

WCS WildlifeConservationSociety

WhO WorldHealthOrganization

WMNWr WichitaMountainsNationalWildlifeRefuge,Oklahoma

WWF WorldWildlifeFund

YDOE YukonDepartmentoftheEnvironment

YNp YellowstoneNationalPark,Idaho,MontanaandWyoming

Yt YukonTerritory,Canada

AmericanBison:StatusSurveyandConservationGuidelines2010xv

ExecutiveSummaryCurtis h. Freese and C. Cormack Gates

ThepublicationofthisIUCNAmericanBisonStatusSurveyand

ConservationGuidelinesistimelyowingtoarecentconvergence

offactors:newresearchfindingsonbisongeneticsandecology,

assessmentandawarenessoftheprecariousstatusofmany

bisonconservationherds,newinitiativesbygovernmentand

non-profitinstitutionstoimprovemanagementofexistingherds

andtoestablishconservationherds,growinginterestamong

NativeAmericansinrestoringbisonaspartoftheircultural

heritage,andanincreasingawarenessbythecommercialbison

industrythatconservationofwild-typebisonisinthelong-

terminterestoftheindustry.Thereisalsoagrowingbodyof

evidencethatthebiodiversityofecosystemswithintheoriginal

rangeofbisoncanbenefitfrombisonrestoration,fromthe

desertgrasslandsofnorthernMexico,throughtheGreatPlains,

tothelowlandmeadowsystemsofinteriorAlaska.Theten

chaptersofthisbookexaminetheseandotheraspectsofthe

biologyandconservationofthespecies,andofferguidelines

forwhatweanticipatewillbeaneweraofbisonconservation

inNorthAmerica.UndertheauspicesoftheIUCNAmerican

BisonSpecialistGroup,twenty-ninechaptercoordinators

andcontributorssharetheirknowledgeandideasinthis

comprehensivereviewofthediversetopicsthatneedtobe

consideredbyresearchers,managers,policymakersandothers

interestedinrestoringandconservingthismagnificentanimal.

In the introductory chapter, C.GatesandP.Goganexplain

theoverallpurposeoftheIUCNAmericanBisonSpecialist

Groupandthisdocument.TheSpecialistGroupiscomposedof

morethan60registeredmembersandnumerouscollaborators

fromthethreenationscomprisingNorthAmericaandranging

fromChihuahuaStateinMexicototheStateofAlaska.The

GroupoperatesundertheaegisoftheIUCNSpeciesSurvival

Commission.Theauthorsnotethatthepurposeofthisvolume

istocontributetothedevelopmentofstrategiesandactions

that,wherefeasible,willconserveandecologicallyrestore

bisonaswildlifethroughouttheiroriginalrange.Gatesand

Goganacknowledgethatlarge-scalerestorationofbisonis

anambitiousandcomplexundertaking,perhapsunparalleled

inspeciesconservationeffortsinNorthAmerica.Their

introductionbrieflyreviewsthemajorissuesfacingbison

conservationandthestronginfluencethatbisonhistorically

exertedonecosystemsacrossmuchofthecontinent.Apart

fromtheecologicalimportanceofbison,thesocialandcultural

significanceofbisonrestorationisrecognisedwhenthey

state,“nootherwildlifespecieshasexercisedsuchaprofound

influenceonthehumanhistoryofacontinent.”

In Chapter 2, B.Potterandco-authorstracetheevolutionary

andrecenthistoryofbison,beginningwiththeearliestfossil

recordsshowingbisoninAsiaatleasttwomillionyearsago,and

continuingwiththeirexpansion,muchlater,intoNorthAmerica

acrosstheBeringLandBridgeduringthemiddlePleistocene.

Theevolutionanddistributionofvariousbisonspeciesand

subspeciesinNorthAmericapresentacomplexstoryshaped,

inlargepart,bybisonhabitatandrangesthatshiftedwidelywith

advancingandretreatingcontinentalicesheets.Theresultofthis

evolutionaryhistorytodayistwospecies,theEuropeanbison

andAmericanbison,andtwosubspeciesofAmericanbison,

woodbisonandplainsbison.Fivehundredyearsago,tensof

millionsofplainsbisonprobablyinhabitedNorthAmerica,from

southernCanadatonorthernMexico,andfromnearlythewest

coasttotheeastcoast,withtheGreatPlainsastheircentreof

abundance.Woodbison,becauseofamorerestrictedboreal

foresthabitat,weremuchlessnumerous.Formanynative

peoplesofNorthAmerica,thousandsofyearsofcoexistence

hadledtobisonbeingcentraltotheirsurvivalandcultures,

ahistorythatPotteret al.exploreinsomedetail.European

colonisationofNorthAmericabroughtrapidchangetoboth

bisonandNativeAmericans.Commercialhunting,competition

withlivestock,killingofbisonasgovernmentpolicytosubjugate

Indiantribes,andothercausesledtotheprecipitousdecline

ofbothplainsandwoodbison.Bytheendofthe19thCentury

afewhundredbisonsurvivedinvarioussmallcaptiveandwild

herdsacrossNorthAmerica.Fortunately,conservationefforts

quicklyemergedinbothCanadaandtheUnitedStates(U.S.)

and,onceprotected,bisonnumbersbegantorecover.Their

iconicstatusnowseemstoberecoveringalso.Potteret al.echo

whatotherauthorsofthisvolumehaveexpressedwhenthey

notethatnootherNorthAmericanspeciesholdssuchgreat

culturalandpoliticalsignificance.

In Chapter 3,D.Boydandco-authorsreviewtheconfusing

anddisputedevidencefor,anddiverseopinionsabout,bison

taxonomy.Agreementseemstoendwiththeconsensusthat

bisonbelongtothefamilyBovidae.Muchofthedebatecentres

onwhetherbisonbelongtothegenusBos,thegenusofcattle,

guar,yak,andoxen,ortotheirowngenus,Bison.Bothnames

arecurrentlyusedinthescientificliterature.Differencesof

opinionarelargelybasedontheimportanceofmorphological

(phenetic)versusmolecular(phylogenetic)linesofevidence,and

onhistoricalprecedenceandusage.WithinBison,therearealso

somepeoplewhoquestionthedesignationofEuropeanbison

andAmericanbisonasseparatespecies.Boydet al.conclude

xvi AmericanBison:StatusSurveyandConservationGuidelines2010

that“Furtherresearchanddebatebytaxonomists,andthebison

conservationcommunity,isrequiredtoreconcilemolecular,

behaviouralandmorphologicalevidencebeforeachangein

nomenclaturecouldbesupported,andthus,forthisdocument,

theAmericanBisonSpecialistGroupadherestothegenusBison

withtwospecies,B. bonasusandB. bison.Notsurprisingly,

disagreementalsoexistsregardingthesubspeciesstatusof

woodandplainsbison.However,Boydet al.emphasisethatthis

debatedoesnotnegatetheimportanceofconservingthetwo

formsasseparateentities.Fromaconservationperspective,the

goalistoconserve“evolutionarilysignificantunits”or“distinct

populationsegments,”amongothertermsusedtodefine

geographicvariationamongpopulations,aconceptrecognised

byboththeU.S.EndangeredSpeciesActandtheCommittee

ontheStatusofEndangeredWildlifeinCanada.Keepingwood

bisonandplainsbisonasseparatenon-interbreedingunitsisthe

recommendedprecaution.

Geneticsplayaparticularlycomplexandimportantrolein

bisonconservation,asexplainedbyD.Boydandco-authors

inChapter 4.Therapidlyadvancingscienceofgeneticshas

recentlybroughtnewinformationandinsightsintonotjust

theevolutionaryrelationshipsamongbisontaxa,butalsoto

managingforviablebisonpopulationsandconservingthe

wildbisongenome.Boydet al.reviewthecurrentstateof

bisongeneticsandwhatneedstobedonetoaddressthe

majorthreatstogeneticdiversityandintegrity—demographic

bottlenecks,foundereffects,geneticdrift,andinbreeding—allof

whichbisonhaveexperienced.Althoughpopulationbottlenecks

canleadtosignificantlossofgeneticdiversity,bisonappear

tohavelargelyavoidedthisproblemduringtheirpopulation

bottleneckinthelate1800s.Giventhegooddiversitywithin

thebisongenepool,andrecentevidencethatshowsseveral

conservationherdsaregeneticallydistinguishable,oneofthe

mostimportantmanagementquestionsishowtomanagethe

populationgeneticsoftheseoftenrelativelysmallherds.Should

thisbeaccomplishedasonelargemetapopulationorasclosed

herdstomaintainlocaliseddiversity?Thebestconservation

strategyistodoboth,and,wherepossible,toincreasethe

sizeofsmallherdstoattainalargeeffectivepopulationsize.

Hybridisationalsoposeschallengesforbisonconservation.

Althoughtheintroductionofplainsbisonintowoodbisonrange

hasresultedinsomehybridisation,thetwoformsremaindistinct

andavoidingfurtherhybridisationisapriority.Muchmore

widespread,andofgreaterconcern,istheintrogressionofcattle

genesintothebisongenome,alegacyofattemptstocross-

breedcattleandbisonthatbeganwhenbisonnumberswere

stilllowintheearly1900s.GenetictestingreviewedbyBoydet

al.indicatesthatmostconservationherdshavesomelevelof

cattle-geneintrogressioninthenuclearand(or)mitochondrial

DNA.Byinferencethisstronglysuggeststhatavastmajorityof

commercialherdshavecattle-geneintrogression.Theeffects

ofintrogressiononbisonbiologyarelargelyunknown.No

introgressionhasbeendetectedinseveralconservationherds,

whichconsequentlydeservepriorityattentionformaintaining

inreproductiveisolation,andassourcestockforestablishing

newconservationherds.Finally,Boydet al.notethatthe

approximately400,000bisonincommercialherdsinNorth

America,some93%ofthetotalcontinentalpopulation,are

undergoingartificialselectionfordomestictraits,suchasease

ofhandling,bodyconformation,carcasscomposition,andso

on.Domestication,whetherintentionalornot,posesaspecial

challengetoconservingthewildbisongenome.

In Chapter 5,K.Auneandco-authorsprovideacomprehensive

reviewofhowdiseases,particularlythosethatare“reportable”

accordingtofederalorstate/provincialregulations,haveamajor

influenceonbisonrestorationandmanagement.Theydescribe

thecharacteristicsandimplicationsofninediseasesforbison

conservation,rangingfromanthraxandbluetonguetobovine

brucellosisandbovinespongiformencephalopathy.Federaland

state/provincialregulationsfor,andmanagementresponsesto,a

particulardiseasedependonseveralfactors,includingpotential

effectsonbison,threattolivestockandhumans,andwhetherit

isindigenousorexotictobisonandtheecosystem.Theauthors

describethecomplexanddifficultmanagementchallengesthat

diseasespresentinthreeofNorthAmerica’smostimportant

conservationherds:theplainsbisonherdsofYellowstone

NationalPark(YNP)andGrandTetonNationalPark/NationalElk

Refugethatharbourbrucellosis,andthewoodbisonherdsin

andaroundWoodBuffaloNationalParkthatareinfectedwith

bothbovinetuberculosis(BTB)andbrucellosis.Diseasessuch

asbrucellosisalsoseverelylimitthetranslocationofbisonfrom

infected,importantconservationherds,suchastheYellowstone

herd,toestablishnewherdsinnewareasbecauseofconcerns

aboutpotentialtransmissiontocattle.Whilethepoliciesand

legalframeworkforcontrollingdiseaseindomesticlivestockare

wellestablished,theydonotworkwellwhenappliedtowildlife,

includingbison,becausetheyoftenconflictwithconservation

goalsandourabilitytomanageandmaintainwildpopulations.

Therecentdevelopmentofnationalwildlifehealthstrategiesin

bothCanadaandtheU.S.couldhelpaddressthisproblem.

Chapter 6,byP.Goganandco-authors,addressesgeneral

biology,ecology,anddemographicsofbison.Bisonare

remarkablyadaptabletoawiderangeofecosystemsand

climaticregimes.Physiologically,bisonaremuchbetteradapted

toclimateextremesthancattle.Behaviourally,bisonexhibita

relativelysimplesocialstructurewithcow-calfpairsatthecore

and,morelooselyandsomewhatseasonally,largegroupsof

cows,calvesandimmaturemales,andseparate,smallergroups

ofmaturebulls.Bisonexhibitindividualandgroupdefence

againstlargepredatorssuchaswolves.Historically,plainsbison

madeseasonalmigrationsbetweensummerandwinterranges,

insomecasesnorth-southandinothersbetweentheprairies

AmericanBison:StatusSurveyandConservationGuidelines2010xvii

andfoothills.Bisonhaveaprofoundinfluenceonecosystems

andcreatehabitatheterogeneitythroughvariousmeans.As

primarilygraminoid(grassesandsedges)eaters,variablegrazing

pressurebyfree-rangingbisonandtheirinteractionwithfire

createhabitatpatchinessonwhichgrasslandbirddiversity

depends.Wallowingbehaviourfurtherpromotesheterogeneity

byformingtemporarypoolsandchangingsurfacehydrologyand

runoffandcreatinglocalpatchesofdisturbedsoilinwhichsome

floweringplantspeciesprosper.Bisonaredispersersofseeds,

andaresourcesandredistributorsofnutrientsforpredators,

scavengers,plants,andecosystemprocesses.Goganet al.

describeforagingpatternsandhabitatusebywoodandplains

bisoninvariousecoregions,fromthearidsouthwesttohumid

coldborealregions.Theauthorsalsoreviewbisonpopulation

structureandreproductionanddemonstratethatundernatural

conditionsnewlyestablishedbisonpopulationscandouble

everyfourtosixyears.Populationnumbersareaffectedbyboth

density-independentevents,suchasseverewintersandwild

fires,anddensity-dependentfactorssuchasdiseaseandwolf

predation.Whilehumanswereabisonpredatorforthousandsof

years,theadventoffirearmsgreatlyincreasedhumanpredation,

sothatbythemid-1800s,anestimated500,000plainsbison

werekilledannuallyforsubsistenceand100,000forhides.The

human-firearm-commercecombination,itwouldseem,largely

voidedthedensity-dependentrelationshipbetweenbisonand

humanpredationuntilitwasalmosttoolatefortheAmerican

bison.

In Chapter 7,C.Gatesandco-authorsassessthestatusof

conservationherdsusingsevencriteria:numericalstatus,

geographicstatus,populationsizeandclassdistribution,

opportunityformatecompetitionamongmaturemales,presence

ofwolves,presenceofdiseasesthatcouldaffectconservation

status,andoccurrenceorlikelyoccurrenceofcattle-gene

introgression.Thedesignation“conservationherd”isassigned

toherdsmanagedbyfederalorstate/provincialgovernments

ornon-governmentalorganisations(NGOs)whosemission

isnatureconservation.Remarkably,littleprogresshasbeen

madeinrecentdecadesinincreasingthenumberofanimals

inconservationherds.Fromthefewhundredthatremainedin

thelate1800s,thenumberofanimalsinconservationherds

increasedinthefirsthalfofthe1900s,butthenlevelledoff,or

inthecaseofthewoodbison,evendeclined,whilethenumber

ofconservationherdshascontinuedtogrowtothepresent

day.Asof2008,therewere62plainsbisonconservationherds

containingabout20,500animals,and11conservationherds

ofwoodbisoncontainingnearly11,000animals.Meanwhile,

startinginthe1980s,thecommercialbisonindustryprospered

withthetotalpopulationgrowingtoaround400,000animals

in2007,roughlyevenlydividedbetweentheU.S.andCanada.

Althoughafewconservationherdsexceed1,000animals,most

conservationherdsofbothwoodandplainsbisonhavefewer

than400animalsand,inthecaseoftheplainsbison,many

arefencedinareasofonlyafewthousandshectaresandnot

subjecttonaturalpredation.Untilrecently,therewasawild

bisonherdinhabitingatrans-boundaryareabetweenMexico

andtheU.S.,theonlyherdmeritingconservationstatusin

Mexico.Butnow,ithasbeenrestrictedtoaprivateranchon

theU.S.side.TheAmericanbisonnearlyqualifiesforlistingas

VulnerableCa2(1)underIUCNcriteriaandiscurrentlylistedas

NearThreatenedontheIUCNRedList.

AsK.Auneandco-authorsdescribeinChapter 8,bison

conservationmustdealwithacomplexmazeoflegaland

policyissues.Muchofthiscomplexityisduetoahistoryof

bisonbeingtreatedlikelivestock.Astheauthorsnote,“During

thegreatrestorationperiodofwildlifemanagement,bison

wereroutinelyclassifiedandmanagedbystate/provincialand

federalagenciesacrossNorthAmericaasaformoflivestock,

whileotherwildlifewereclassedandmanagedasfree-roaming

wildanimals.”Theysubsequentlyprovideadetailedreviewof

thelegalstatusof,andconservationinitiativesunderwayfor,

bisoninMexico,theU.S.,andCanada.Thelegalrecognition

ofbisonaswildlifeorlivestock,orboth,variesacrossvarious

federal,state,andprovincialjurisdictionsinNorthAmerica.For

example,onlytenU.S.states,fourCanadianprovincesand

twoterritories,andoneMexicanstateclassifybisonaswildlife;

allotherstatesandprovinceswithinthebison’shistoricrange

designatethemasdomesticlivestock.Overlayingthislegal

mapforbisonareseveralstakeholdergroupsthatmanage

bison:publicwildlifeandlandmanagementagencies,Native

Americangroups,non-profitconservationorganisations,and

privateproducers.Reportablediseasespresentanotherset

oflegalissuesthataffectinternationalandinterstatetransport

ofbison.Auneet al.suggestthataparadigmshiftisrequired

wherebythepublicrecognisesbisonaswildlife,andthatthere

ismuchgreatersocialtolerance,especiallyintheagricultural

community,ifmajorprogressistobemadeinre-establishing

free-rangingbisonontheirnativerange.Moreover,large-scale

restorationoverbiglandscapeswilltypicallyrequirepartnerships

andco-managementamongmultiplelandownersandresource

managers,andmoreenlightenedandcoordinatedgovernment

regulationsandpolicies.

In Chapter 9,J.E.Grossandco-authorsprovideguidelines

forpopulation,genetic,anddiseasemanagementforboth

existingconservationherdsandforthefullrecoveryofbison

overboththeshortandlongterm.Astheauthorsexplain,

conservationfocusesonretainingexistingecological,cultural,

andgeneticcharacteristicsofbison,whereasfullrecovery

entailsabroadervisionofbisoninhabitinglandscapesthat

permitthefullexpressionofnaturalbehavioursandecosystem

interactionsthatonceexisted.Theguidelinesfirstaddressbison

behaviour,particularlytheimportanceofensuringnaturalmating

systemsthatinvolveavoidingaskewedsexratioandallowing

xviii AmericanBison:StatusSurveyandConservationGuidelines2010

competitionamongbulls,aswellasotherfactors,suchas

naturalmovementsandmortalityrates.Giventhesmallsizeof

manyexistingherdsandnewlyestablishedherds,guidelinesfor

populationandgeneticmanagementareparticularlyimportant.

Herdsof1,000ormoreanimalsareimportantforconserving

geneticdiversity,andfactorssuchasnon-randommating,

skewedsexratios,andlargeswingsinpopulationsizeneedto

beavoidedinrelativelysmallherds.Managingbisonforrestoring

andmaintainingbiodiversityinvolvesallowinganimalsto

naturallymoveandforageacrossthelandscape,andtointeract

withothernaturalprocessessuchasfire,drought,andsnow

cover.Guidelinesareprovidedforactivemanagement,including

handlingandherdingandthetypeofinfrastructurerequired,

withthecaveatthatactivemanagementandhandlingshouldbe

minimised.Diseaseguidelinesaddressprevention,surveillance

and,whenpathogensaredetected,management.Gross

et al.stresstheimportanceofwell-designedreintroduction

programsforestablishingnewherdsandoffersuggestions

rangingfromstakeholderinvolvementtosourcinganimalsand

ensuringproperherdstructure.Givenconcernsaboutthe

geneticuniquenessofsomeherdsandcattle-geneintrogression,

similarcareneedstobegivenintransferringanimalsbetween

herdswiththegoalofmaintaininggeneticdiversityand(or)

aidingintherecoveryofsmallorthreatenedherds.Thechapter

concludeswithrecommendationsforusingmodellingand

computersimulationstoassessbisonpopulationsandhabitat.

the concluding chapter(10) onguidelinesforecological

restorationbyC.Gatesandco-authorsisdirectedat

establishingnew,largepopulationsofbisononlargelandscapes.

Becausebisonwereanecologicallydominantspeciesover

muchoftheirrange,restoringhistoricecologicalprocessesand

biodiversityinareastheyonceinhabiteddependsonrestoring

large,free-roamingherds.Fullecologicalrestorationisdefined

as“there-establishmentofapopulationofseveralthousand

individualsoftheappropriatesubspeciesinanareaoforiginal

rangeinwhichbisoninteractinecologicallysignificantwayswith

thefullestpossiblesetofothernativespeciesandbiophysical

elementsofthelandscape,withminimalnecessarymanagement

interventions.”Althoughthefocusofthischapterisonrestoring

largeherdsoverlargeareas,whereprocessessuchasmigration

andnaturalselectionaremostlikelyfulfilled,Gateset al.point

outthatsmallherdscanalsocontributetorestoringmany

ecologicalprocessesthatoccuratsmallerscales.Thechapter

providesguidelinesforplanningandexecutinglarge-scale

re-introductions,includingafeasibilityanalysisthataddresses

bothbiologicalquestionsandathoroughassessmentof

socioeconomicvariablesandlegalrequirements,sourcingand

thenreintroducingsuitablestock,andfollow-upmonitoring,

evaluationandadaptationasexperienceisgainedandlessons

learned.Asnotedaswellinchapter8,oneofthebiggest

challengesfacinglarge-scalerestorationisthatassemblinga

landscapeofhundredsofthousandsormillionsofhectareswill

usuallyrequirepartnershipsandco-managementofmultiple

landowners,bothpublicandprivate,andthesupportofmany

stakeholders.

AmericanBison:StatusSurveyandConservationGuidelines20101

TheABSGisagroupofvolunteersrepresentingavarietyof

disciplinarybackgrounds,expertise,andprofessional

experience.Theyaregeographicallydistributedacrossthe

breadthoftheoriginalcontinentalrangeofthespecies,from

MexicotoAlaska,andfromtheTallgrassPrairieintheeasttothe

intermountainwest.Theyworkforavarietyofinstitutions

includinggovernments,conservationorganisations,and

academicinstitutions(seeAcknowledgements).

1.2 Context

PriortoEuropeansettlement,theAmericanbisonhadthe

largestoriginaldistributionofanyindigenouslargeherbivorein

NorthAmerica,rangingfromthedesertgrasslandsofnorthern

MexicotothefloodplainmeadowsofinteriorAlaska(Listet al.

2006;Stephensonet al.2001)andalmostfromcoasttocoast.

Theecologicalscopeofthespecieswaslimitedonlybyits

habitatrequirementsandspecialiseddiet.Anobligategrazer,

grassesandsedgespresentingrasslandsandmeadowsarethe

mainstayoftheAmericanbison’sdietandhabitat.Bisonhave

beencontinuouslypresentinNorthAmericaforatleast300,000

years,persistinginvariousformsduringthelatePleistocene

throughsequentialglacialandinterglacialperiods,theninto

theHoloceneandpresenttimes(MacDonald1981;Shapiroet

al.2004;Wilsonet al.2008).Theyhavebeenassociatedwith

successiveculturessincehumansfirstoccupiedthecontinent

about12,000yearsago.

Overhundredsofthousandsofyears,bisonhavecontributed

totheco-evolutionofotherbiota,includinggrazingadaptations

inplants,mutualistic,commensalandtrophicinterrelationships,

andbisonhavefunctionedasakeycomponentofthenative

biodiversityinvastareasofthecontinent.Keyspecies,suchas

bison,haveamarkedinfluenceonthepatternsofoccurrence,

distribution,anddensityofotherspecies(MeffeandCarroll1994;

Paine1969).Wherepresent,bisonplayimportantecological

rolesbyinfluencingthestructure,compositionandstabilityof

bothplant(Campbellet al.1994;Knappet al.1999)andanimal

communities(Bogan1997;Roe1970;Truettet al.2001).

the primary goal of the American Bison

Specialist Group (ABSG) is to contribute

to the development of comprehensive and

viable strategies and management actions

to enhance conservation and achieve

ecological restoration of bison as wildlife

where feasible throughout the original

the primary goal of the American Bison

Specialist Group (ABSG) is to contribute

to the development of comprehensive and

viable strategies and management actions

to enhance conservation and achieve

ecological restoration of bison as wildlife

where feasible throughout the original

1.1 the Species Survival Commission and the American Bison Specialist Group

TheInternationalUnionforConservationofNature(IUCN)

SpeciesSurvivalCommission(SSC)isascience-basednetwork

ofapproximately8,000volunteerexpertsfromalmostevery

countryoftheworld,workingtogethertowards“A world that

values and conserves present levels of biodiversity.”Withinthe

SSC,over100specialistgroupsandmorethan15independent

RedListAuthoritiesaresetuptotrack

species’status,monitorbiodiversity,analyse

issues,developsolutions,andimplement

actions(SSCStrategicPlan2001-2010).

Amongthem,theBisonSpecialistGroupis

distinguishedbytwoorganisationalunits,

onefortheEuropeanbison(Bison bonasus),

andtheother,fortheAmericanbison(Bison

bison).

TheprimarygoalsoftheAmericanBisonSpecialistGroup

(ABSG),andtheintentofthisdocument,aretocontributeto

thedevelopmentofcomprehensiveandviablestrategiesand

managementactionstopromoteconservationandecological

restorationofbisonaswildlifewherefeasiblethroughoutthe

originalrangeofthespecies.Conservationandecological

restorationofbison,aswildlife,atthescaleofitsoriginal

continentalrangeareambitiousandcomplexendeavours,

perhapsmoresothanforanyotherNorthAmericanspecies.

Enhancingthelong-termsecurityofbison,aswildlife,willrequire

thecommitmentandparticipationofkeysectors,including

publicwildlifeandlandmanagementagencies,non-government

environmentalorganisations,aboriginalgovernmentsand

communities,localcommunities,andconservation-oriented

commercialproducers.Towardthisgoal,theABSGwas

establishedtoincludeabroadnetworkofpeopleinterested

inbisonconservationandrecovery.Therearemorethan60

registeredmembersandnumerousothercollaborators.Aswith

otherspecialistgroups,thisnetworkofvolunteersrepresents

thefunctionalcapacityoftheIUCNtomonitorthestatusand

managementofAmericanbisoninrelationtoglobalandlocal

biodiversity.SpecialistGroupparticipantscontributedthe

scientificandpracticalknowledgeassembledinthisreport,and

canofferexpertadviceand,inmanyinstances,themeansto

makethingshappenonthegroundbyimplementingactionsor

encouragingandfacilitatingotherstoadvancetheconservation

andecologicalrestorationofbisonaswildlife.

Chapter1 Introduction:TheContextLead authors: C. Cormack Gates and Peter J. P. Gogan

the primary goal of the American Bison Specialist

Group (ABSG) is to contribute to the development of

comprehensive and viable strategies and management

actions to enhance conservation and achieve ecological

restoration of bison as wildlife where feasible throughout

2 AmericanBison:StatusSurveyandConservationGuidelines2010

Nootherwildlifespecieshasexercisedsuchaprofound

influenceonthehumanhistoryofacontinent.Asthegreat

icesheetsreceded,andgrassesandsedgescolonisedthe

emerginglandscape,beginning14,000yearsago,bison,then

humanculturesfollowed.Widespreadandabundant(Shaw

1995),bisonwereastapleresourceformorethan12,000years

inthesubsistenceeconomiesofsuccessiveculturesofNative

NorthAmericans.Duringbriefrecenthistory,overthelast

500yearsorso,Europeanscolonisedtheeasternseaboard,

exploredwestwardintotheNative-occupiedprairiesandthe

North,foughtforresources,dominatedindigenouspeoples,

andprosperedasnewsettlersandindustrialsocieties.Trading

postsrecruitedindigenouspeopletoharvestbisonformeatand

pemmicanforthefortsandtofuelthetradeinfurs(Gateset

al.1992).Armiesclashedundertheprairieskies(Greene1996)

andrailwayswerebuilttoconnecttheWesttoeasternmarkets.

Millionsofplainsbisonwerekilledfortheirmeat,hidesfor

machinebeltsandrobes,forsport,andtosubjugatetheFirst

Nations,makingwayforsettlersocietyanddomesticEuropean

livestock(Hornaday1889;Isenberg2000).Inlessthanacentury,

fromChihuahuaStateinMexicototheStateofAlaska,the

mostabundantindigenouslargeherbivoreinNorthAmerica

wasdrivenclosetoextinction.Haditnotbeenfortheinterest

ofprivatecitizensinrearingafewsurvivorsincaptivity(Coder

1975),andtheremotenessofalonewildpopulationinwhatis

nowYellowstoneNationalPark(YNP)(Meagher1973),plains

bisonwouldhavedisappearedfromthecontinent.Similarly,by

theendofthe“GreatContraction”ofplainsbisonlateinthe19th

Century(Flores1996),woodbisonwerealsoreducedtoasingle

survivingpopulationoffewerthan300animalsinaremotearea

intheforestedborderlandsofAlberta(AB)andtheNorthwest

Territories(NWT)(Gateset al.1992;2001).

Duringthe20thandintothe21stCentury,federal

andstate/provincialagenciesandconservation

organisationsplayedanimportantroleinthe

conservationandrecoveryofbisonaswildlife.Sixty-

twoplainsbisonand11woodbisonherdshavebeen

establishedforconservation,representingabout

7%ofthecontinentalpopulation.Inparallel,since

about1980,thenumberofbisonraisedundercaptive

commercialpropagationhasincreasedmarkedly,and

nowrepresentabout93%ofthecontinentalpopulation

(Chapter7).

1.3 Current Challenges for Conservation and Ecological restoration of Bison as Wildlife

Conservationofanywildlifespeciesrequiresensuring

bothlong-termpersistenceofasufficientnumber

ofpopulationsandmaintainingthepotentialfor plate 1.1 Free ranging bison in Yellowstone National Park. Photo: John Gross.

ecologicaladaptationresultingfromnaturalselectionoperating

onindividualsinviablepopulationsinthewild(IUCN2003;

SecretariatoftheConventiononBiologicalDiversity1992;Soulé

1987).Inwildmammalpopulations,limitingfactors,suchas

predation,seasonalresourcelimitation,andmatecompetition,

contributetomaintainingthewildcharacter,geneticdiversity,

andheritabletraitsthatenableaspeciestoadaptto,and

persist,inanaturalsetting(Knowleset al.1998).Thelong-

termconservationofAmericanbisonaswildlifeisfacedwith

severalimportantchallengesthatneedtobeacknowledged

andaddressedbypublicagencies,non-profitorganisations

andproducerorganisations.Theyincludetherarityoflargewild

populationsinextensivenativelandscapes,conservingthewild

characterandgenomeofbison,andthepresenceofregulated

diseases.

1.4 Large Wild populations

Bisoncanbestachievetheirfullpotentialasanevolving,

ecologicallyinteractivespeciesinlargepopulationsoccupying

extensivenativelandscapeswherehumaninfluenceisminimal

andafullsuiteofnaturallimitingfactorsispresent.Whilesuch

conditionsremainavailableinthenorthofthecontinent,it

ischallengingtofindextensivelandscapesforrestoringand

sustaininglargefree-roamingwildbisonpopulationsinsouthern,

agriculture-dominatedregions.Ecologicalrestorationisthe

intentionalprocessofassistingrecoveryofanecosystemthat

hasbeenmodified,degraded,damagedordestroyedrelative

toareferencestateortrajectorythroughtime(SERIandIUCN

CommissiononEcosystemManagement2004).Asdescribed

bytheIUCNCommissiononEcosystemManagement,

ecologicalrestorationhas,asitsgoal,anecosystemthatis

resilientandself-sustainingwithrespecttostructure,species


compositionandfunction,aswellasbeingintegratedinto

thelargerlandscape,andsupportingsustainablehuman

livelihoods.Ecologicalrestorationinvolvingbisonasanintegral

componentofecosystemsfacestwomajorchallenges:1)

howtoundertakerestorationacrosslargeareaswithdiverse

land-useandownershippatterns;and2)howtoundertake

restorationinawaythatimprovesbothbiodiversityandhuman

wellbeing.Large-scaleecologicalrestorationinvolvesbiological

andsocialcomplexity.Attitudes,economicsandpolitics,from

localtoregionalandinternationalscales,willshapethefuture

ofbisonconservationonoccupiedlands.Thesechallengesare

addressedinChapter10.

1.5 Conserving the Wild Character and Genome of Bison

Bisonincaptiveherdsmaybemanagedtoachievevarious

objectives,includingtheecologicalservicesthatbisonprovide

(e.g.,grazing,nutrientcycling,andterraindisturbance),

educationanddisplay,commercialproduction,and

conservationofbisonaswildlife.Conservingbisonaswildlife

isnotnecessarilyservedbymanagingapopulationforother

purposes.Forexample,theecologicaleffectsofherbivorymay

beachievedbygrazingavarietyoflivestockspecies.Although

somerangelandsformerlyusedforcattleproductionhave

beenconvertedtobisonproduction,thesubstitutionofbison

forcattleproductiondoesnot,byitself,necessarilycontribute

tobisonconservation,ortoecologicalrestorationofbison

aswildlife.Similarly,displayherdsmayserveconservation

educationobjectiveswithoutotherwisecontributingtospecies

conservation.

Intheabsenceofintentionalpoliciesandactionstoconserve

thewildcharacterandgenomeofbison,captivityand

commercialisationcanleadinadvertentlyorintentionallyto

avarietyofeffectsthatmaybedeleterioustobisonasa

wildlifespeciesintheintermediatetolongterm(Chapter4).

Theseincludeeffectsonthegenome:foundereffect;reduced

geneticdiversity;persistenceandphenotypicpenetration

ofdeleteriousgenes;orinadvertentselectionforheritable

morphology,tamenessoradaptationtocaptivity.Small

populationsareparticularlysusceptibletosucheffects.Thesex

andagestructureofcaptiveconservationpopulationsmaybe

manipulatedtoreducetheriskofescape,removeaggressive

animals,ortoalterfecundityortherateofpopulationincrease.

Theagecompositionofmalesincaptiveherdsistypically

substantiallydifferentfromwildpopulations.

Thecommonpracticeincaptivecommercialherdsofeliminating

males,beforetheybecomemorphologicallyandbehaviourally

mature,posesachallengingquestionabouttherolesof

matecompetitionandnaturalselectionforfitnessinsuch

populations.Ingeneral,selectionpressuresoncaptivewildlife

aresubstantiallydifferentfromthoseinthewild.O’Regan

andKitchener(2005)positedthatdomesticationmayoccur

inadvertentlyincaptivewildmammalsthroughpassiveselection

forindividualsbehaviourallysuitedtocaptivity,withconcomitant

morphologicalchangesoverseveralgenerations.Mostchanges

arethoughttoresultfromincreasingpaedomorphosis,whereby

juvenilecharacteristicsareretainedintheadultformofan

organism(O’ReganandKitchner2005).Clutton-Brock(1999)

describedchangesinlargemammalsundercaptiveconditions

includingreducedbodyandbrainsize,alteredexternal

appearance,thegainingofafatlayerbeneaththeskinanda

reductionofthefacialregion.Inadvertentselectionfortameness

andadaptationtoacaptiveenvironmentistypicalinmammals

(Frankhamet al.1986),andinadditiontoaltering“wildness”,

canreducethechancesforsuccessfulreintroductionofcaptives

intothewild.Alossofresponsetopredatorsandalteration

ofdefensiveandsexualbehaviourshavealsobeenreported

incaptivewildlife(Price1999;2002).Manycommercialbison

producersdirectlyselectformarketabletraitssuchasearly

maturity,coatcolour,bodysizeandconformation.Thelatter

“showringtraits”arepromotedinbisonindustryadvertisements,

publicationsandatauctions.

Thelargesizeofthecommercialcaptivepopulationisthebasis

forapopularmisconceptionthatthespeciesis“secure”,leading

wildlifemanagementagenciestoignoreactionsnecessaryfor

conservationofwildtypebison.Today,amongNorthAmerican

jurisdictions,thereisaconfusingarrayofclassificationsofbison

aswildlife,domesticlivestock,orboth(Chapter8).

Hybridisationwithcattleisanotherseriouschallengeforbison

conservation.IntheU.S.,Canada,andEurope,agricultural

interestsattemptedtodevelopanimprovedrangeanimalby

hybridisingbisonandcattle.Forced-matingofbisonandcattle

canbereadilyachievedinacontrolledenvironment.However,

theypreferentiallymatewiththeirownspeciesunderopen

rangeconditions(Boyd1908;Goodnight1914;Jones1907).

InEurope,theEuropeanbison(Bison bonasus),arelativeof

theAmericanbison,andtheaurochs(Bos taurus primigeneus),

progenitorofmoderncattle,weresympatric,yetevolutionarily

divergent,units.Typicalofsympatricspeciesoccupyingsimilar

trophicniches,behaviouralandecologicalspecialisation

providesnicheseparation,leadingtoreproductiveisolation

andprogressivelytospeciation(Bush1975;RiceandHostert

1993).Speciesdivergenceandreproductiveincompatibility

areevidentfromthelowfertilityoffirstgeneration(F1)bisonx

cattleoffspring(Boyd1908;SteklenevandYasinetskaya1982)

andthedifficultyproducingviablemaleoffspring(Boyd1914;

Goodnight1914;SteklenevandYasinetskaya1982;Steklenevet

al.1986).Unfortunately,forcedhybridisationsbetweenB. bison

andBos taurusinNorthAmericahaveleftalegacyofcattle

mitochondrial(Polziehnet al.1995;Wardet al.1999)andnuclear

DNA(Halbert2003;Halbertet al.2005).Thisintrogressionis


widespreadamongcontemporarybisonpopulations,inboth

publicandprivatesectorherds(Chapter4).Theimplications

forbisonconservationarejustbeginningtobeunderstoodand

appropriateinterventionsconsidered.

1.6 reportable Diseases

Bisonhostnumerousparasitesandpathogens(Reynolds

et al.2003;Tessaro1989),someofwhichareimportantto

conservation.Livestockdiseasesthatrestricttradeorposearisk

tohumanhealthandare‘reportable’underfederal,provincial,

andstatelegislationareparticularlyimportantbecause

theymayinducemanagementactionsthatnegativelyaffect

bisonconservationandrestoration(Chapter5).Management

interventionsmayincludedepopulation,limitingdispersal

andrangeexpansiontoprotectadjacentbisonorlivestock

populations,andrestrainingtranslocations.Thepresenceor

perceivedriskofreportablediseasesinbisondevaluesthem

aswildlifeandconstrainsconservationandrecoverypotential.

Largefree-rangingbisonpopulationsareinfectedwithexotic

(non-native)reportablediseasesintwoareasofthecontinent,

theGreaterYellowstoneArea(GYA)mainlyinMontanaand

Wyoming(bovinebrucellosis),andtheGreaterWoodBuffalo

EcosysteminAlbertaandtheNorthwestTerritories(bovine

brucellosisandtuberculosis).Balancingconservationwith

intensiveinterventionsisaperpetualchallengefortheagencies

responsibleformanagingthesepopulations.

1.7 purpose of this Document

Thisdocumentprovidesanauthoritativesummaryofthebiology

andstatusofAmericanbison,including:prehistorictorecent

historyandculturalcontext(Chapter2);taxonomyandrelated

issues(Chapter3);geneticvariationandeffectsofhuman

interventionsonthegenome(Chapter4);diseasesthatdirectly

orindirectlyaffectbisonconservation(Chapter5);biologyand

ecologyofthespecies(Chapter6);thenumericandgeographic

statusofAmericanbison,emphasizingherdsmanagedprimarily

forconservation(Chapter7);legislationandpoliciespertaining

tobisoninallrangestates(Chapter8).Guidelinesforbison

conservationareprovidedinthefinaltwochaptersofthis

document(Chapter9PopulationandGeneticsGuidelines;

Chapter10EcologicalRestorationGuidelines).Throughoutthe

documentreferenceismadetochallengesrequiringactions

rangingfromurgenttolongterm.

Non-prescriptiveguidanceisofferedonhowconservationand

ecologicalrestorationofbisonaswildlifemaybeachieved,while

respectingtheprinciplesofdemocraticgovernanceinthethree

nationsformingNorthAmerica,thesustainabilityofeconomic

useofecologicalresources,culturalheritagevalues,and

ecologicalvaluesofintactecosystems.


2.1 palaeobiology and phylogeny

Bisonhaveexistedinvariousformsformorethan2,000,000

years(Danz1997;McDonald1981).Earlyformsoriginatedin

AsiaandappearinVillafranchiandeposits,andintheearly

fossilrecordinIndia,China,andEurope(Guthrie1990;Shapiro

et al.2004).BisonoccupiedEurasiaabout700,000yearsago

thenmovedacrosstheBeringLandBridgeintoAlaskaduring

themiddlePleistocene300,000–130,000yearsago(Illinoin

Glaciation;MarineOxygenIsotopeStages(MIS)8to6(Shapiro

et al.2004).AllSiberianandAmericanbisonsharedacommon

maternalancestorabout160,000yearsago(Shapiroet al.2004).

Fossilevidenceindicatestherewasasinglespecies,oratleast

asimilarlarge-hornedformwithvariablespecies/sub-species

designations,thesteppebison,Bison priscus,throughout

Beringia(Guthrie1990).

Chapter2 HistoryofBisoninNorthAmericaLead Authors: Ben A. Potter, S. Craig Gerlach, and C. Cormack Gates,

Contributors: Delaney P. Boyd, Gerald A. Oetelaar, and James H. Shaw

Villafranchian: a major division of early pleistocene time, named for a sequence of terrestrial sediments studied in the region of Villafranca d’Asti, an Italian town near turin. this was a time when new mammals suddenly appeared.

holarctic: a term used by zoologists to delineate much of Eurasia and North America, which have been connected by the Bering land bridge when sea levels are low during glacial periods.

plesitocene: Ice Age. A division of geological time; epoch of the Quaternary period following the pliocene. During the pleistocene, large areas of the northern hemisphere were covered with ice and there were successive glacial advances and retreats.

Beringia: a 1,000 mile wide ice-free grassland steppe, in Asia and North America linked together by the “Bering Land Bridge” when sea levels were low. Animals traveled in both directions across this vast steppe, and humans entered the Americas from what is now Siberia.

Glacial periods: there have been at least four major ice ages. the present ice age began 40 million years ago with the growth of an ice sheet in Antarctica. Since then, the world has seen cycles of glaciation with ice sheets advancing and retreating on 40,000- and 100,000-year time scales. the most recent glacial period ended about ten thousand years ago.

Marine isotopic stages (MIS): alternating warm and cool periods in the Earth’s ancient climate, deduced from oxygen isotope data reflecting temperature curves derived from data from deep sea core samples.

Ural Mountains: a mountain range that runs

plate 2.1 Skull of Bisonpriscus, Yukon Canada. Photo: Cormack Gates.

Steppebisonprobablyreachedtheirmaximumdistributionand

abundanceduringthelastglacialperiod(Wisconsinan,100,000–

12,000yearsB.P.;MIS2-4and5a-d).Thesearethetypicalbison

fossilsfoundintheYukonandAlaskaduringthatperiod.Steppe

bisonhadrelativelylonghindlegs,similartotheEuropean

bison(B. bonasus),andlargehornswithtipscurvedback,anda

secondhump(Guthrie1990).Analysisofancientmitochondrial

DNA(mtDNA)(Shapiroet al.2004)suggeststhatLate

Pleistocenebison,foundfromtheUralMountainstonorthern

China,weredescendantsofoneormorereversedispersalsfrom

NorthAmerica.Themostrecentcommonancestorofbison

specimensanalysedbyShapiroet al.(2004)existedtowardsthe

endoftheIllinoianGlacialPeriod(MIS6).


BisonmovedsouthintothegrasslandsofcentralNorthAmerica

whentheicesheetsretreatedatthebeginningoftheSangamon

Interglacial(MIS5e)130,000-75,000yearsB.P.(MacDonald

1981),evolvingthereintoalargeform,B. latifrons.Thisgiant

bisonpossessedahornspanofmorethantwometresand

wasabundantinthecentralcontinentduringtheSangamon

Interglacial.Itunderwentagradualreductioninbodysizeand

hornspan(Guthrie1980;vanZylldeJong1993).Duringthe

subsequentWisconsinGlaciation(110,000-12,000yearsB.P.;

MIS2-4and5a-d),Beringianandsouthernpopulationsbecame

separatedastheLaurentidecontinentalicesheetextended

intowesternCanadafrom20,000-13,000yearsB.P.(Burns

1996;Wilson1996).Geographicseparationhadprofound

biological,taxonomic,andevolutionaryeffects.Southernbison

evolvedintodistinctivephenotypes(vanZylldeJong1993)and

separatemtDNAclades.AllmodernAmericanbisonnowbelong

toasinglecladethatisdistinctfromBeringeanbison,witha

mostrecentcommonancestorbetween22,000and15,000

yearsB.P.(Shapiroet al.2004).Thisinterpretationisconsistent

phenotype: Observable physical or biochemical characteristics of an organism. phenotype is determined by both genetic makeup and environmental influences.

Clade: A biological group (taxa) that share features inherited from a common ancestor.

holocene: A geological period, which began approximately 11,550 calendar years B.p. (about 9600 BC) and continues to the present. It has been identified with MIS 1 and can be considered an interglacial in the current ice age.

phylogenetics: the study of evolutionary relatedness among groups of organisms.

Glacial maximum: the time of maximum extent of the ice sheets during the last glaciation (the Würm or Wisconsin glaciation), approximately 20,000 years ago.

taphonomic processes: the transition of the remains, parts, or products of organisms in soil, e.g. the creation of fossil assemblages through burial.

taxonomy: the science of classification of organisms. Nomenclature is the system of naming

withcompleteseparationbetweennorthernandsouthern

populationsatthetimeofthelastglacialmaximum(20,000-

18,000yearsB.P.).

DatapresentedbyShapiroet al.(2004)andWilsonet al.(2008)

supportthehypothesisthatmodernbisonaredescended

frompopulationsthatoccurredsouthoftheicesheetbefore

theLastGlacialMaximum.Southernbisonunderwentrapid

in situevolutionduringtheearlyHolocenefromB. antiquusto

anintermediateformB. occidentalis,thentothemodernform

B. bison(Wilsonet al.2008).Whenthecontinentalicesheets

begantomelt,bisoninvadedtheemergingice-freecorridor

fromthesouthwherethawingandmeltingoccurredfirst.

ColonisationfromBeringiawaslimited(Shapiroet al.2004).

Overlapbetweennorthernandsouthernbisonoccurredinthe

vicinityofthePeaceRiverinnorth-easternBritishColumbia

wherenorthernbisonwerepresentby11,200-10,200yearsB.P.

(Shapiroet al.2004),andsouthernformsofbisonwerepresent

10,500yearsB.P.MolecularresearchbyShapiroet al.(2004)

indicatesthatallmodernbisonaredescendedfrompopulations

livingsouthoftheicesheetbeforetheLastGlacialMaximum.

ThetwomodernNorthAmericansubspecies(plainsbisonand

woodbison)divergedbyabout5,000yearsago(Gateset al.

2001;vanZylldeJong1986).Thewoodbison(B.b. athabascae)

wasthemostrecentvarianttooccurinAlaska,theYukonand

NorthwestTerritoriesandtheplainsbison(B.b. bison)isthe

mostrecentsouthernvariantoftheNorthAmericanspecies

(vanZylldeJong1993Stephensonet al.2001).Small-horned

bisonsimilartowoodbisonalsooccurredinnorthernEurasia

duringtheHolocene(Flerov1979;Lazarevet al.1998;vanZyll

deJong1986,1993).AlthoughtheEuropeanbison(B. bonasus)

ismorphologicallysimilartoandreadilyinterbreedswiththe

Americanbison,theyformdistinctlydifferentcladesbasedon

mtDNAsequencesofthe273bp-longfragmentofcytochrome

bgene(Prusaket al.2004).Thisisconsistentwithgeographic

separationbetweenthesetwospeciesstartingduringthemid-

Pleistoceneandbeforereverse-dispersaloccurredfromNorth

AmericatoSiberia.

2.2 Original range

PrevioustypologiesdividetheHolocenerangeofbisoninto

“prehistoric”and“historic”periods(vanZylldeJong1986).

Thedistinctionbetweenthemisnotbasedonobjectiveor

biologicallymeaningfulcriteria,andprovidesanartificialand

confusingtemporaldichotomythatpersistsdespitewell-

informedargumentstothecontrary(Stephensonet al.2001).

Apreferredandmoreaccuratealternativeistorefertothe

previousrangeofbisonas“original”range,therebyavoiding

thenecessitytodistinguishbetweenwrittenrecordsand

othersourcesincludingzooarchaeologicalevidenceandorally

transmittedknowledge(Gateset al.2001).


ModernbisonoriginallyrangedacrossmostofNorthAmerica

(Figure2.1).PlainsbisonweremostabundantontheGreat

Plains,butalsoradiatedeastwardintotheGreatLakesregion,

overtheAlleghenyMountainstowardtheeasternseaboard,

northwardasfarasnorthernNewEngland,andthensouthinto

Florida;westward,theywerefoundinNevadaandpartsofthe

GreatBasin,theCascadeandRockyMountainsnorthwardto

mid-AlbertaandSaskatchewanprairielands,andfurthersouth

alongtheGulfofMexicointoMexico(Danz1997;Reynoldset

al.1982).Therearerecordsofbisonoccurringatsurprisingly

highelevationsinmountainousregions,particularlyalong

theFrontRangeoftheRockyMountains(Fryxell1928;Kay

andWhite2001;Meagher1986).Evidencealsoindicatesthat

bisoninhabitedareasoftheGreaterSouthwest,including

Arizona,NewMexico,andnorthernMexico,areasnotgenerally

recognisedaswithintheoriginalrangeofplainsbison(Truett

1996).Whetherapparentorreal,bisonscarcityintheAmerican

Southwestisusuallyattributedtoacombinationofinsufficient

waterandgrassandhumanhunting(Truett1996).Theoriginal

rangeofwoodbisonincludesnorthernAlberta,north-eastern

BritishColumbia,asmallareaofnorth-westernSaskatchewan,

thewesternNorthwestTerritories,Yukon,andmuchofAlaska

(Stephensonet al.2001).Morerecentresearchincorporating

Figure 2.1 Original ranges of plains bison and wood bison. Recreated by Boyd (2003) based on van Zyll de Jong (1986) and Stephenson et al. (2001).

oralnarrativesofaboriginalpeopleinAlaska,Yukon,and

NorthwestTerritories,incombinationwitharchaeologicaland

palaeontologicalrecords,demonstratesthatwoodbisonwere

presentintheYukonandAlaskawithinthelasttwocenturies,

andthattheseareasarewithintheoriginalrangeofthe

subspecies(Lotenberg1996;Stephensonet al.2001).

2.3 Abundance

Historicalandarchaeologicalrecordsdemonstratethatplains

bisonthrivedonthegrasslandsoftheGreatPlains(Malainey

andSherriff1996;ShawandLee1997).Explorers,settlers,

andEuroamericanhuntersdescribedenormousherdsof

plainsbison,withpopulationestimatesrangingfrom15to100

million(Dary1989;Shaw1995).Inthe1890s,naturalistErnest

ThompsonSetonpositedthewidelyacceptedestimatefor

Americanbisonat60million(Dary1989;McHugh1972;Roe

1970;Shaw1995).

Severalquantitativeandqualitativemethodshavebeenused

toestimatepre-settlementbisonabundance,includingdirect

observation,carryingcapacitycalculations,andcountsof

bisonkilledformarketinthelate1800s.Evenwhenusedin

combination,allmethodsarefraughtwithuncertainty,untested,


fromdomesticlivestock(horses,cattle,sheep)andwild

horsesalsoplayedaroleinreducingbisonnumbers(Flores

1991;Isenberg2000).Furthermore,becausebisonprovided

sustenanceforNorthAmericanaboriginalsandcommodities

fortheirbartereconomy,theeliminationofbisonwasviewedby

Euroamericansasthemostexpedientmethodtosubjugatethe

NativeAmericansandforcethemontoreserves,makingwayfor

agrariansettlementandcontinuedwesterndevelopment(Danz

1997;Geist1996;Isenberg2000;MayerandRoth1958).Tothis

end,theU.S.governmentunofficiallysupportedtheslaughter

ofbisonbyprovidingammunitionandsuppliestocommercial

buffalohunters(MayerandRoth1958).Althoughanovert

politicalpolicytodecimatebisonwasneverformallyestablished,

theCanadianandU.S.governmentscapitalisedonwidespread

hungeramongaboriginalcommunitiescausedbythenear

extirpationofbisonasameanstosubjugateandcontrolthe

aboriginalpopulation(Geist1996;StonechildandWaiser1997).

Bythelate19thCenturyitwasestimatedthattherewerefewer

than1,000remainingbisoninNorthAmerica(Hornaday1889;

Seton1927).WoodbisonwereconcentratedinnorthernAlberta

andtheNorthwestTerritories,andplainsbisonwerescatteredin

isolatedgroupsacrosstheCentralGreatPlainsand,notably,in

whatisnowYellowstoneNationalPark(YNP).

2.5 Early recovery

Asthegreatherdsdiminished,therewassomepublicoutcry,

butfewlawswereenactedtoprotectthebison(Danz1997).

Mostearlyplainsbisonconservationeffortshappenedthrough

theindependentactionsofprivatecitizens.Prominentfigures

intheconservationmovementincludedJamesMcKayand

evenunwarrantedassumptions,andarbitrarypopulation

attributions(Shaw1995).Regardless,thereislittledoubtthat

priortoEuroamericansettlement,plainsbisonnumberedinthe

millions,andprobablyeveninthetensofmillions(Shaw1995).

Woodbisonwerenotasnumerousasplainsbisonowingto

limitedhabitat,althoughtheydidinhabitavastregionofthe

borealforestinnorth-westernNorthAmerica(Gateset al.

2001c).Soper(1941)estimatedthetotalwoodbisonpopulation

in1800tobe168,000,anestimatethatwashighlyspeculative.

TheSoperestimateisbasedonthenumberanddistribution

ofwoodbisonexistinginWoodBuffaloNationalPark(WBNP)

duringthe1930s,withsomefuzzyextrapolationfromthe

WBNPdensitytothepresumedareaoftheoriginalwoodbison

range.Theestimatedidnotaccountforregionalvariability

inhabitatavailability.Furthermore,Stephensonet al.(2001)

documentedaconsiderablylargeroriginalrangethanSoper

(1941).Therefore,woodbisonmayhavebeenmorenumerous

thanestimatedbySoper.

2.4 Extirpation

Continentalbisonnumbersdeclineddramaticallyandrapidly

followingEuropeansettlement.Specificregionalimpactson

numbers,distribution,andabundancearerecordedinmany

historicalaccountsandreferences(e.g.,Dary1974).Large-

scaleseasonalmigrationsofboththenorthernandsouthern

plainsbisonherdsmayhavetemporarilymaskedtheirdecline,

althoughbythelate1800sitwasobviousthattheAmerican

bisonpopulationhadbeendecimatedandwasinseriousdecline

(Krech1999).CommercialhuntingbyEuroamericansandsome

NativeNorthAmericansformeatandhideswasaprimary

cause(Hornaday1889;Isenberg2000).The

Americanmilitaryquietlyapprovedillicitmarket

huntingonfederallyprotectedtriballandsin

thenorthernandsouthernplains.Otherfactors

includedindiscriminateslaughterforsportand

recreation.Sporthuntingwasexacerbatedby

thewestwardpushofcolonizationfromthe

eastandacrosstheprairieswiththeimplicit

andexplicitapprovalofpoliticiansandmilitary

leadersanxioustoresolvethefoodsupply

sideoftheso-called“Indianproblem.”(Danz

1997;Dary1989;Hewitt1919;Isenberg2000;

McHugh1972).

Environmentalfactors,suchasregionaldrought,

introducedbovinediseases,andcompetition

plate 2.2 An enormous pile of bison skulls waiting

to be ground for fertilizer (c. mid-1870s). Copyright

expired - Courtesy of the Burton Historical Collection,

Detroit Public Library - downloaded from English

Wikipedia 20 Aug 2009.


CharlesAlloway(Manitoba),CharlesGoodnight(Texas),Walking

Coyote(Montana),FrederickDupree(SouthDakota),CharlesJ.

Jones(Kansas),andMichelPabloandCharlesAllard(Montana)

(Coder1975;Danz1997;Dary1989;Geist1996).Theirefforts

toestablishherdsfromthefewremainingbisonsecuredthe

foundationstockformostcontemporarypublicandprivate

plainsbisonherds.Formedin1905,theAmericanBison

Society(ABS)pressedCongresstoestablishseveralpublic

bisonherdsatWichitaMountainsNationalWildlifeRefuge,the

NationalBisonRange(NBR),Sully’sHillNationalGamePreserve

(SHNGP),andFortNiobraraNationalWildlifeRefuge(Coder

1975;Danz1997).NationalparksinboththeU.S.andCanada

alsofiguredprominentlyinbisonrecoveryefforts(Danz1997;

Ogilvie1979).

Onceplainsbisonwereprotectedfromhunting(beginninginthe

1870s),theirnumbersincreasedconsiderably,doublingbetween

1888and1902.By1909,thesubspecieswasconsideredsafe

fromextinction(Coder1975).Initiallysparkedbynostalgiaand

reverencefortheanimal,motivationsforbisonrecoverybecame

increasinglydrivenbytheircommercialvalue(YorksandCapels

1998).By1970,therewere30,000plainsbisoninNorthAmerica,

withapproximatelyhalfinpublicherdslocatedinnationalparks,

wildliferefuges,andstatewildlifeareas,andhalfinprivateherds

(ShawandMeagher2000).Asreviewedinchapter7,thenumber

ofplainsbisoncurrentlyismorethan20,500in62conservation

herds,whilethenumberundercommercialpropagationisabout

400,000.

Thewoodbisonpopulationfelltoalowof250animalsatthe

closeofthe19thCentury,thenslowlygrewto1,500-2,000by

1922owingtotheenforcementofCanadianlawsenactedto

protecttheanimal(Gateset al.2001c;Soper1941).In2008,

therewereabout10,870woodbisonin11conservationherds

(Chapter7).

2.6 Cultural Significance

Fewspeciesenjoyahistoryasrichinarchaeology,

palaeontology,storyandlegend,oralanddocumentaryhistory

astheAmericanbison.NoristhereanotherNorthAmerican

speciesforwhichtheculturalandpoliticalsignificanceofan

animalissogreat.Forthousandsofyearsvariousformsand

populationsofbisonhavecoexistedwithhumansinNorth

America,providingsustenanceandshapinghumansocial

andeconomicpatterns,andinfluencingnationalhistoryand

internationalpoliticalrelationships.Althoughacomprehensive

reviewofhuman-bisoninteractionsfromthecolonisationof

NorthAmericatorecenttimesisencyclopaedicinscope,abrief

summaryanddiscussionisprovidedhere.

Bisonwereimportantinthesubsistenceeconomiesofthe

firstBeringiancolonisersofthewesternhemisphere,andlater

figuredprominently,butdifferentially,inPalaeo-Indian,Archaic,

andsubsequentNorthAmericanculturalhorizonsandtraditions.

Bisonwereeconomicallyandculturallyimportantthroughout

mostofNorthAmerica,includinginteriorAlaska,Yukonand

NorthwestTerritories,buttheywereparticularlysignificantfor

groupslivingintheGreatPlains,fromnorth-centralTexasto

southernAlberta.Variousformsofbisonhavebeenidentifiedas

keysubsistenceresourcesinthePalaeolithicofnorth-eastern

Asia,formingpartofamegafaunalcomplexadaptedtothe

steppe-tundraofLatePleistocenenorthernEurasiaandBeringia,

alongwithmammothsandhorses(Guthrie1990).Whilebison

remainsarecommonlyfoundinSiberianarchaeologicalsites,

standardzooarchaeologicalmethods(Ermolova1978)indicate

theydonotappeartohavecontributedgreatlytosubsistence.

Bycomparison,reindeer,mammoths,andhorsesarerelatively

abundantinSiberianarchaeologicalsites.Bisonseemtohave

playedamoreimportantroleinNorthAmericanarchaeological

complexes.InAlaska,thereisempiricalevidencefromnumerous

archaeologicalcomplexesspanning12,000to1,000years

B.P.thatlinksbisonwithculturaltraditionsusingconservative,

palaeo-Indian: (12,000-6,000 B.p.) A group

of Late pleistocene–Early holocene cultures

associated with the colonisation of central North

America. While their subsistence economies are

debated, many archaeologists consider them

to be big game hunting specialists (including

mammoth).

Folsom: (11,000-10,200 B.p.) A palaeoindian

culture, characterised by very high mobility and

specialised bison hunting.

Archaic: (6,000-2,300 B.p.) A group of Middle

holocene cultures characterised by broad

spectrum foraging (i.e., subsisting on a wide

variety of big and small game, fish, shellfish,

and plant foods). they do not have permanent

villages or agriculture.

plains Woodland: (2,300-1,000 B.p.) A group of

Late holocene cultures characterised by semi-

permanent villages, horticulture (maize and

beans) in addition to hunting and gathering.

Altithermal: also the holocene Climate Optimum.

A warm period during the interval 9,000 to

5,000 years B.p. this event is also known by

other names, including: hypsithermal, Climatic


burialordestructionthrougherosion)(Artz1996;Walker1992).

Someevidenceindicatesthatduringthisperiodbisonand

peopleconcentratedtheiractivitiesinlocalisedrefugia,suchas

rivervalleys(Buchner1982).ThroughoutNorthAmerica,there

wasageneralshifttomixedforagingeconomiesbasedonmore

locallyabundantresources,withbisonplayingamuchsmaller

roleexceptinspecificareasoftheGreatPlains.

After2,000yearsB.P.,archaeologicalrecordsfortheNorth

Americangrasslandsshowevidenceofwidespreadhuman

occupationandregionalspecialisationinhabitatuse(Manning

1995;Speth1983).Theso-calledPlainsWoodlandcomplexes

showedlocalpatternsofadaptationrepresentedaswidespread

networksofculturalinteractionsthatlinkedtheeastern

woodlands,andperhapseventheGreaterSouthwest,tothe

grasslandsthroughtradeandreligiousorceremonialinteractions

(Frison1991).Technologiesshiftedagaintoincludebowsand

arrows,potteryanddistinctiveregionalceramictraditions.

Muchlater,theuseofhorsesformedthebasisforthemounted,

efficientmicrobladetechnology(HolmesandBacon

1982;Potter2005;2008).Microbladesaresmallelongate

sharpstonebladesinsertedintopiecesofboneorwood

tomakecompositetools(Guthrie1983).

BisonplayedakeyroleinPalaeo-Indian,Archaic,

andlatereconomiesinNorthAmerica,particularlyin

theGreatPlains.Whilesomehavequestionedearly

Palaeo-Indiandependenceonbisonandotherlarge-

bodiedungulates(GraysonandMeltzer2002),other

studiesshowaclearpatternofspecialisedlarge

mammalhuntingduringtheLatePleistoceneandEarly

HoloceneinNorthAmerica(HofmanandTodd2001;

WaguespackandSurovell2003).Althoughthereare

disagreementsastowhetherEarlyPalaeo-Indians

shouldbeclassifiedasspecialisedbig-gamehunters

orbroad-spectrumforagers,bisonevidentlyplayedan

importantroleintheirsubsistenceeconomies.Arecent

surveybyWaguespackandSurovell(2003)reported

that52%of35EarlyPalaeo-Indiancomponents(Clovis,

11,300-10,900yearsB.P.)includedbisonremains.With

theextinctionofthemammothandotherPleistocene

megafauna,bisonbecameagreatereconomicfocus

forlatePalaeo-Indiancomplexes(Folsomandothers

presentduringtheEarlyHolocene).Changesinprojectile

pointformshavebeenlinkedtospecialisationsforbison

hunting(Stanford1999).Inparticular,Folsomcomplex

adaptationshavebeenlinkedtointensivebisonhunting

(Amick1996).Communalbisonhuntingprobablyplayed

animportantroleinseasonalaggregationsofPalaeo-

Indianpopulations,withhumangroupscombiningtohuntand

thendispersingintosmallergroupsinrelationtoseasonalbison

migrations(KellyandTodd1988).

OntheGreatPlains,theHoloceneClimaticOptimumor

Altithermal(about7,500yearsB.P.inmid-latitudeNorth

America)resultedinwarmeranddrierconditionsandincreased

seasonality.Climatechangeapparentlylimitedbisonabundance

andgeographicdistribution,andinducedhumanadaptations

tonewclimaticandecologicalconditions(Sheehan2002;but

seeLovvornet al.2001).Humanpopulationsadjustedprimarily

bydevelopingneweconomicstrategies,termed“Archaic”

byNorthAmericanarchaeologists.Adaptationsinvolvednew

technologiessuchasgroundstoneforprocessingavarietyof

plantfoods,andincorporatingamorediversearrayofsmaller

gameandplantsintothesubsistenceeconomy.Duringthis

period,someportionsoftheGreatPlainsappeartohavebeen

abandonedentirelybypeople(Meltzer1999).However,the

dearthofsitescouldalsobeexplainedbytaphonomy(deep

plate 2.3 Arvo Looking Horse performing a ceremony

honouring slaughtered bison after a harvest near Yellowstone

National Park. Photo: Jim Peaco, National Park Service.


nomadic“PlainsIndianCulture”observedbyEuropeanexplorers

andmissionariesatfirstcontact(Duke1991;Wedel1959).

NativeNorthAmericans,during,andevenafterthePlains

Woodlandtradition,livedinlargermorepermanentvillages.They

dependedonmaize,bean,andgourdhorticulturetonamesome

ofthemostimportantdomesticates,withwinterdependence

ondeerandseasonalmovementsinthefallandspringtotake

advantageofmigratingbisonherds(Wilson1987).Thispatternis

wellrepresentedethnographicallyintheMiddleMissouriRegion.

GroupsliketheSiouxan-speakingMandanandHidatsa,andthe

Caddoan-speakingPawneeandArikara,withtheWichitaand

others,werescatteredalongmajorPrairieriversandtributaries

liketheLoup,LowerLoup,Canadian,andWashita,asfarsouth

asNebraska,Kansas,andOklahoma(Weltfish1965).Largekill

events,suchasthoserepresentedattheHead-Smashed-Insite

inAlberta,generallydidnotoccuruntilverylateinthehistory

ofbisonhuntingonthePlains,andarerepresentedfromthe

LateArchaicandlaterperiods(Byerlyet al.2005).Theshiftin

huntingstrategiesmayhavebeenaresponsetoincreasingherd

sizes,introductionofbowandarrow,and/orchangesinsocial

organisation(Driver1990;Reeves1990;Walde2006).

Withincreasedresolutionandclarityaffordedbyethnohistoric

andethnographicinvestigations,human-bisoninteractions

amonghistoricnativepeoplesarebetterdescribedand

documentedthanforthelatePleistoceneandHolocene.

Bisoncontinuedtobethepreferredgameformanynative

NorthAmericancultures,especiallyontheGreatPlainsand

Prairies,providingfood,clothing,shelter,andtools(Geist1996;

Roe1970).Sustainedbybisonandplantresources,many

nativegroupslikelyaffecteddensitiesofotherlargeherbivore

species(Kayet al.2000;MartinandSzuter1999).Inaddition

tosignificantecologicalrelationships,thebisonwasacentral

elementinoraltradition,rituals,dances,andceremoniesof

nativepeoplesofthePlains(Wissler1927),anditremains

symbolicallyimportantintheculturaltraditionsofmanynative

Tribestothisday.

ThearrivalofEuropeansinNorthAmerica,after1492,

resultedinsignificantchangesinhuman-bisoninteractions,

andchangedthefabricofNativeAmericanlifeforever.

Introduceddiseasessuchassmallpoxdecimatedindigenous

humanpopulations(Crosby1986),andalteredsubsistence,

settlement,demography,andsocialorganisationformany

differentgroups.Bisonhuntingbynativepeoplewasseasonal

innature.Bisonwereincorporatedintoabroadspectrumof

plantandanimalprocurementactivities(Holder1970;Isenberg

2000).Bisonprovidedtheeconomicbasisforstable,resilient

landuseregimesandsocialsystems.However,effectsof

NativeAmericanwarfareandraidingduringthehistoricperiod

disruptedanddestabilisedtheselanduseandsocialsystems.

ThespreadofhorsesintoGreatPlainsaboriginaleconomiesby

the1750s,andincreasingcommoditisationofbisonproducts

causedbytheemergenceofaEuropeancommercialmarket

forwildlifeproductsbythe1820s,contributedtothenear

extinctionofthebison(Flores1994;Isenberg2000:27).Native

peoplestradedbisonhidesforEuro-americancommodities,with

themarketinbisonrobesreachingapeakinthe1840s.Hide

huntersbegantosignificantlyparticipateinthemarkethuntingof

plainsbisoninthe1850s,andbythe1890shaddecimatedthe

herds.Evenboneswerecleanedforsaletotheeasternfertilizer

market,anactivitythatcontinuedto1906(Dary1974).

NumerousnativeNorthAmericantribesmanagebisononnative

andtriballands,butcultural,socialandspiritualrelationships

withthisanimalarechanging.FormanyNativeAmericans

thereisstillastrongspiritualandsymbolicconnection,but

forothersitisthepotentialcommercialvalueofbisonthat

ismostimportant.Forstillothers,itisthepragmaticuseof

bisonforfood,andtherelationshipbetweenlocalcontrolover

foodproductionandland,foodsecurity,tribalsovereignty,

anddecreasingrelianceonoutsidesourcesforfoodand

commoditiesthatisemergingasatopicofconcern,anda

themeunderlyingtribaldecision-making.

ItisnotjusttherelationshipbetweenNativeAmericansand

bisonthatischanging,buttheroleofbisonintheoverall

NorthAmericanfoodsystemischangingaswell.TheNorth

Americanperspectiveisshiftingfromtheviewthatbisonare

anartifactfromthepasttobeviewedassuchinparksand

preserves,toonethatseesbisonasadynamiccomponent

oftheAmericandiet.Alongwithanewvisionforahealthy

ecologicalandgeneticfuturefortheAmericanbison,food

systemresearchers,foodsystementhusiasts,andthe

biomedicalresearchcommunityenvisionanewroleforbisonin

theAmericandiet.Thisroleelevatestheanimaltopriorityover

industriallyraisedbeefandpork,andsecuresforitaplaceas

thehealthyalternativetoafatty,sugar-baseddietthatalready

hassignificanthealthimpactsintermsofincreasedratesof

cardiovasculardisease,colorectalandotherformsofcancer,

anddiabetes.Free-rangebisonmeatishigherinomega-3fatty

acidsthanaregrain-fedanimals,perhapsevenashighaswild

salmonandothercoldwaterfishspecies,anditisalsohigh

inconjugatedlinoleicacid,afat-blockerandanti-carcinogen

withthepotentialtoreducetheriskofcancer,diabetes,and

obesity.Theextenttowhichbisoncanbeproducedefficiently

andinhealthywaysthatdonotfurtherdegradeecosystems

andecosystemservices,andmarketedasahealthyfoodat

anaffordableprice,willperhapsbethetippingpointsforhow

importantbisonbecomeinafutureAmericanfoodsystem.

WhetherNativeAmericanornot,culturalvalues,attitudes,and

perspectivesarereflectedinhowwethinkabout,manage,and

handleanimalsinthewild,incommercialproductionsystems,

andafterbutcheringandprocessingthroughmarketing.Bison

areperhapsuniqueinthatwemanagethembothaswildlifeand


aslivestock,withwoodbisoninAlaskaandCanadaanexample

oftheformer,andplainsbisonintheCanadianandAmerican

Plainsanexampleofthelatter.Thejuryisprobablystillout

onwhetherwewillmanagebisonaswildlife,aslivestock,or

asbothinthefuture,butitisclearthatthereisabrightrolefor

thisanimalinanemergingNorthAmericanfoodsystemand

tradition.NativeAmericansarebothrecoveringandrestoring

theirlong-establishedculturalrelationshipwiththeAmerican

bison,andNativeAmericansandothernon-nativeNorth

Americansarefindingnewwaystorelatetothisanimalinways

thatwillenhancetheconservationofthespecies.


Chapter3 TaxonomyandNomenclatureLead Authors: Delaney P. Boyd, Gregory A. Wilson, and C. Cormack Gates

Thepurposeofnamingorganismsistofacilitaterecognition

andcommunicationandtoidentifypatternsandapplypractical

structuretothenaturalworld.Taxonomycansupportthe

conservationandsustainableuseofbiologicaldiversityby

contributingtoidentification,assessment,andmonitoring

programmes(EnvironmentAustralia1998).Taxonomyisalso

vitalforthecreationandinterpretationoflaws,treaties,and

conservationprogrammesbecauseitcreateslegalidentities

fororganisms(Geist1991).Whileitisimportanttostrivefor

accuracyintaxonomicclassification,semanticissuesand

uncertaintycancreatesubstantialmanagementchallenges

bydistractingconservationdecisionmakersfromtheissues

threateningataxonorbiologicalunitworthyofconservation.

Despitetheextensivehistory,andtheeconomicandsymbolic

importanceofbisontoNorthAmericansocieties,there

remainssignificantconfusionanddisagreementaboutbison

taxonomy.Theissuesrangefromanhistoricaldiscrepancy

overthecommonname,toongoingscientificdebateoverthe

systematicsofthegenus,species,andsubspeciesdesignations.

3.1 An historical Misnomer: Bison vs. Buffalo

Thebisonisnotabuffalo.True‘buffalo’arenativeonlyto

Africa(capebuffalo,Syncerus caffer)andAsia(fourspecies

ofwaterbuffalo, Bubalus spp.).Theuseofthetermbuffalo

forAmericanbisonderivedperhapsfromotherlanguages

usedbyexplorerstodescribetheunfamiliarbeast,e.g.,

bisonte, buffes, buffelo, buffles,andbuffilo(Danz1997;Dary

1989).Thesetermsaresimilartobufleandbuffe,whichwere

commonlyusedtorefertoanyanimalthatprovidedgoodhide

forbuffleather(Danz1997).Despitethemisnomer,theterm

‘buffalo’hasbeenusedinterchangeablywith“bison”since

earlyexplorersfirstdiscoveredtheNorthAmericanspecies

(Reynoldset al.1982).Thetermhasbecomeentrenchedasa

colloquialisminNorthAmericancultureandlanguage.Although

scientificconventiondictatesuseof‘bison’,theterm‘buffalo’

persistsasanaccepted,non-scientificconventionforhabitual

andnostalgicreasons.

3.2 Genus: Bos vs. Bison

WhenLinnaeusfirstclassifiedthebisonin1758forhis10th

EditionoftheSystema Naturae,heassignedtheanimalto

Bos,thesamegenusasdomesticcattle(WilsonandReeder

2005).Duringthe19thCentury,taxonomistsdeterminedthat

therewasadequateanatomicaldistinctivenesstowarrant

assigningthebisontoitsowngenus(ShawandMeagher

2000).Therefore,in1827,C.HamiltonSmithassignedthesub-

genericnameBisontotheAmericanbisonandtheEuropean

bison(SkinnerandKaisen1947).In1849,Knightelevated

thesubgenusBisontothelevelofgenus(SkinnerandKaisen

1947).Sincethen,taxonomistshavedebatedthevalidityof

thegenus,somearguingthatbisonarenotsufficientlydistinct

fromcattle,guar,yak,andoxentowarrantadistinctgenus

(Gardner2002,personalcommunication).Duringthelasttwo

decades,asmoleculargeneticandevolutionaryevidence

hasemerged,scientistshaveusedBoswithincreasing

frequency.Discrepanciesinthegenusarereflectedinmajor

cataloguingcentresandbooks.Forexample,theCanadian

MuseumofNature(Balkwill2002,personalcommunication)

andtheSmithsonianNationalMuseumofNaturalHistoryinits

publicationMammal Species of the World(WilsonandReeder

2005)useBison,whiletheRoyalOntarioMuseum(Eger2002,

personalcommunication)andtheMuseumofTexasTech

University,initsRevised Checklist of North American Mammals

North of Mexico(Jones,Jr.et al.1992;Joneset al.1997;Baker

et al.2003),haverevertedtoBos.

Thedebateovertheappropriategenusarisesfromthe

conflictbetweenthetraditionalpracticeofassigningnames

basedonsimilarfeaturesdistinguishablebymorphology(the

pheneticapproach)versususingevolutionaryrelationships(the

phylogeneticapproach)(FreemanandHerron2001;Winston

1999).Systematistsdevelopevolutionarytreesbyanalysing

sharedderivedcharacteristics(FreemanandHerron2001;

Winston1999).Inthisscheme,onlymonophyleticgroups,or

clades,whichrepresentalldescendantsofacommonancestor,

arenamed.Apheneticschememightassignnamestopartial

clades,orparaphyleticgroups,whichexcludeoneormore

descendants(FreemanandHerron2001).Sometaxonomists

andsystematistssuggestthatthetraditionalnamingsystem

bereplacedwithaphylogeneticscheme(FreemanandHerron

2001).Whilenotallbiologistsagreethisisprudent,giventhat

astrictlyphylogeneticschemecouldignorefunctionallyand

ecologicallyimportantdifferencesamongspecies(Freeman

andHerron2001),thephylogeneticapproachprovidessome

usefulinsightsaboutevolutionaryrelationshipswithinthefamily

Bovidae.

BisonresideinthefamilyBovidae,subfamilyBovinae,tribe

Bovini,whichcurrentlycontainsfourgenera:Bubalus(Asian

waterbuffalo);Syncerus(Africanbuffalo);Bos(domesticcattle


andtheirwildrelatives),andBison(bison)(Wallet al.1992;

WilsonandReeder2005).Studiesofnuclear-ribosomalDNA

(Wallet al.1992),mitochondrialDNA(Miyamotoet al.1989;

Miyamotoet al.1993),andrepetitiveDNAsequences(Modiet

al.1996)withinthistribehaverevealedthatthegenusBosis

paraphyleticwithrespecttothegenusBison.MitochondrialDNA

studiesdonotsupportthetraditionalorganisationofthetribe

Bovinibecausetheyak(Bosgrunniens)ismorecloselyrelated

tobisonthantoitscongenercattle(Bos taurus)(Miyamotoet al.

1989;Miyamotoet al.1993).RibosomalDNAstudieshavenot

fullyclarifiedthisrelationship(Wallet al.1992).However,skeletal

analysisbyGroves(1981)notedthatbisonandyakhave14

thoracicvertebraewhileothermembersoftheTribeBovinihave

only13,underscoringtheimportanceofconsideringheritable

morphologicaldifferencesthatmaynotberevealedusing

molecularmethods.

AcomparisonofvariousphylogenetictreesforthetribeBovini

furtherillustratesthenamingconflict.Figure3.1(a)depictsa

Figure 3.1 Comparison of phylogenetic hypotheses for the tribe Bovini based on: (a) conventional morphological analysis (Bohlken 1958); (b) cladistic

analysis of cranial characteristics (Groves 1981); (c) mtDNA sequences (Miyamoto et al. 1989); and (d) ribosomal DNA analysis (Wall et al. 1992).

conventionalschemebasedonmorphologicalcharacteristics

(Bohlken1958),whileFigures3.1(b-d)showdifferent

interpretationsbasedoncranialorgeneticevidence.Although

non-conventionalschemesdonotshareidenticalbranching

patternsforeveryspecies,thepositionofBisonwithinthepattern

ofdevelopmentforeachalternativeisequallyincongruous.Inthe

conventionalscheme,BosbranchedoffthetreelaterthanBison;

however,thearrangementsbasedonmorerecentevidence

suggestthataBosbranchwasfollowedbyBison,thenbyBos.

EachalternativedemonstratesthatBosisparaphyleticbecause

itislackingoneofitsdescendantbranches(denotedasBison).

Underaphylogeneticscheme,bisonwouldbeincludedinthe

Boscladetocorrectthisincongruity.

Forfourdecades,therehavebeensuggestionstocombine

BisonandBosintoonegenus(Baccuset al.1983;Gentry1978;

Groves1981;Miyamotoet al.1989;Modiet al.1996;Stormont

et al.1961;VanGelder1977).StudiesofDNA,bloodtypes,

andchromosomal,immunological,andproteinsequences


demonstratethatBisonandBosweregeneticallysimilar,given

molecularmethodsexistingatthetime(Beintemaet al.1986;

BhambhaniandKuspira1969;Dayhoff1972;Kleinschmidt

andSgouros1987;Stormontet al.1961;Wilsonet al.1985).

Additionally,thepercentdivergencesamongmitochondrial

DNA(MtDNA)sequencesofBison bison,Bosgrunniens,and

Bos tauruswerecomparabletothosecalculatedamongother

setsofcongenericspeciesassesseduntil1989(Miyamotoet

al.1989).Reproductiveinformationalsosupportstheinference

ofaclosephylogeneticrelationshipbetweenBosandBison;

BisonandsomemembersofBoscanhybridiseunderforced

matingtoproducepartiallyfertilefemaleoffspring(Miyamotoet

al.1989;VanGelder1977;Wallet al.1992;Ward2000).Species

divergenceandreproductiveincompatibilityareevidentwith

thelowfertilityoffirstgeneration(F1)bisonxcattleoffspring

(Boyd1908;SteklenevandYasinetskaya1982)andthedifficulty

producingviablemaleoffspring(Boyd1908;Goodnight1914;

SteklenevandYasinetskaya1982;Steklenevet al.1986).

Behaviouralincompatibilityisalsoevident.Althoughmating

ofbisonandcattlecanreadilybeachievedinacontrolled

environment,theypreferentiallyassociateandmatewith

individualsoftheirownspeciesunderopenrangeconditions

(Boyd1908;1914;Goodnight1914;Jones1907).Differences

indigestivephysiologyanddietselectionbetweencattleand

Americanbison(reviewedbyReynoldset al.2003)andEuropean

bison (Gębczyńska and Krasińska 1972) provide further evidence

oftheantiquityofdivergencebetweencattleandbison.Based

onpalaeontologicalevidence,Loftuset al.(1994)concluded

thatthegeneraBosandBisonsharedacommonancestor

1,000,000–1,400,000yearsago.

InNorthAmerica,sympatrybetweenbisonandcattleisan

artefactoftherecenthistoryofcolonisationbyEuropeansand

theirlivestock.However,inprehistoricEurope,thewisent(Bison

bonasus)andaurochs(Bos taurusprimigeneus),theprogenitor

ofmoderncattle,weresympatricyetevolutionarilydivergent

units.Thedivergenceinbehaviour,morphology,physiology,and

ecologyobservedbetweenbisonandcattleisconsistentwith

thetheorythatecologicalspecialisationinsympatricspecies

occupyingsimilartrophicnichesprovidesamechanismfor

reducingcompetitionintheabsenceofgeographicisolation

(Bush1975;RiceandHostert1993).

Theassignmentofananimaltoagenusintraditionalnaming

schemescanbesubjective,andchanginggenericnamescan

createconfusionandcontravenethegoaloftaxonomy,whichis

tostabilisenomenclature(Winston1999).However,wecaution

thatmaintainingastablenomenclatureshouldnotoccurat

theexpenseofmisrepresentingrelationships.Achangeof

BisontoBosmayreflectinferredevolutionaryrelationshipsand

geneticsimilaritiesbetweenBisonandBosspecies.Itcould

alsopotentiallyprovidecontinuityandstabilitytothescientific

referenceforbison,whichcurrentlyhastwospeciesnamesinuse

(B. bonasusandB. bison).However,andincontrast,basedon

divergenceonacytochromebgenesequenceanalysis,Prusaket

al.(2004)concludedthatalthoughAmericanandEuropeanbison

arecloselyrelated,theyshouldbetreatedasseparatespeciesof

thegenusBison,ratherthansubspeciesofabisonspecies.There

isalsothepotentialthatchangingthegenusfromBisontoBos

wouldcomplicatemanagementofEuropean(threesubspecies)

andAmericanbison(twosubspecies)atthesubspecieslevel

anddisruptanestablishedhistoryofpublicpolicyandscientific

communityidentificationwiththegenusBison.

Furtherresearchanddebatebytaxonomistsandthebison

conservationcommunityisrequiredtoreconcilemolecular,

behaviouralandmorphologicalevidencebeforeachange

innomenclaturecouldbesupportedbytheAmericanBison

SpecialistGroup(ABSG).Inconsiderationoftheuncertainties

explainedabove,andinkeepingwiththenamingconventions

formammalsusedforthe1996RedListandthe2008RedList

(WilsonandReeder1993;WilsonandReeder2005),theABSG

adherestothegenusBisonwithtwospecies,Europeanbison

(B. bonasus)andAmericanbison(B. bison),inthisdocument.

3.3 Subspecies

AcontroversialaspectofAmericanbisontaxonomyisthe

legitimacyofthesubspeciesdesignationsforplainsbison(B.

Bison bison)andwoodbison(B. bisonathabascae).Thetwo

subspecieswerefirstdistinguishedin1897,whenRhoads

formallyrecognisedthewoodbisonsubspeciesasB. bison

athabascaebasedondescriptionsoftheanimal(Rhoads

1897).Althoughthetwovariantsdifferinskeletalandexternal

morphologyandpelagecharacteristics(Table3.1),some

scientistshavearguedthatthesedifferencesalonedonot

adequatelysubstantiatesubspeciesdesignation(Geist1991).

Theissueiscomplicatedbythehuman-inducedhybridisation

betweenplainsbisonandwoodbisonthatwasencouraged

inWoodBuffaloNationalPark(WBNP)duringthe1920s.

Furthermore,theconceptofwhatconstitutesasubspecies

continuestoevolve.

Theassignmentofsubspecificstatusvarieswiththeorganism,

thetaxonomist,andwhichofthevariousdefinitionsof

subspeciesisapplied.MayrandAshlock(1991:430)definea

subspeciesas“an aggregate of local populations of a species

inhabiting a geographic subdivision of the range of the species

and differing taxonomically from other populations of the

species.” AviseandBall(1990:59-60)adaptedtheirdefinition

fromtheBiologicalSpeciesConcept,whichdefinesspeciesas

groupsoforganismsthatarereproductivelyisolatedfromother

groups(MayrandAshlock1991):“Subspecies are groups of

actually or potentially interbreeding populations phylogenetically

distinguishable from, but reproductively compatible with, other

such groups.”


Crucialtothisdefinitionistheargumentthatevidencefor

phylogeneticdistinctionmustderivefrommultipleconcordant,

independent,genetically-based(heritable)traits(Aviseand

Ball1990).Essentially,subspeciesshoulddemonstrateseveral

conspicuousmorphologicaldifferences,geographicallopatric

populationpatterns,andnormallypossessgeneticdivergences

atseveralgenes(Winston1999).Hybridisationbetween

subspeciesispossiblealongcontactinterfaces(Winston1999).

Thefossilrecordandobservationsofvariabilityamong

livingbisonsuggestthatthespeciesexhibitedconsiderable

geographicvariation.Thisvariationledtoclaimsidentifying

variousformsofthespecies,mostnotablyanorthernanda

southernplainsbison,whichdifferedinpelageandconformation

(vanZylldeJong1993).Analysisofcranial,horn,andlimb

measurementsforplainsbisonsuggestsclinalvariationalong

anorth-southaxis(McDonald1981;vanZylldeJong1993).

Itispossiblethatexternalcharacteristics,suchaspelage

plains bison Bison bison bison

Wood bison Bison bison athabascae

pelage characteristics

Densewoollybonnetofhairbetweenhorns Forelockdark,hanginginstrandsoverforehead

Thickbeardandfullthroatmane,extendingbelowribcage

Thinbeardandrudimentarythroatmane

Well-developedchaps Reducedchaps

Well-demarcatedcape,lighterincolourthanwoodbison

Noclearcapedemarcation,hairusuallydarkerthanplainsbison

Structural Characteristics

Highestpointofthehumpoverfrontlegs Highestpointofthehumpforwardoffrontlegs

Hornsrarelyextendabovebonnet Hornsusuallyextendaboveforelock

Smallerandlighterthanthewoodbison(withinsimilarageandsexclasses)

Largerandheavierthanplainsbison(withinsimilarageandsexclasses)

table 3.1 Comparison of structural and pelage characteristics for the two bison subspecies. colouration,alsovariedalong

thisaxis(vanZylldeJonget al.

1995).Therefore,thecontinuous

gradationofintermediate

bisonformspreventsdefinitive

recognitionofnorthernand

southernformsofplainsbisonat

thetrinomiallevel.

Unliketheclinalvariation

reportedforplainsbison,a

phenotypicdiscontinuityexists

betweenplainsbisonand

woodbison(vanZylldeJong

1993),reflectedinsizeand

inmorphologicaldifferences

independentofsize(vanZyll

deJong1986;Gateset al.

2001).Discontinuousvariation

occurswhenabarrierimpedes

geneflowbetweenpopulations

ofaspecies,causinggenetic

differencestoaccumulateon

eithersideofthebarrier(vanZyll

deJong1992).Reproductive

isolationcausedbydiffering

habitatpreferencesandseasonal

movements,andthenatural

barrierformedbytheboreal

forest,contributedtomaintaining

thephenotypicdifferences

betweenplainsbisonandwood

bison(vanZylldeJong1986;van

ZylldeJong1993;Gateset al.

2001).TheSocietyforEcological

RestorationInternational(SERI)andIUCNCommissionon

EcosystemManagement(2004)explicitlyrecognisethe

continuousnatureofbiologicalprocesses,suchasspeciation,

initsguidelinesforrestorationofecosystemsthathavebeen“…

degraded, damaged, or destroyed relative to a reference state or

a trajectory through time” (Chapter9).AnalysisofancientmtDNA

indicatesthatmodernAmericanbisonarederivedfromamost

recentcommonancestorexisting22,000to15,000yearsB.P.

(Shapiroet al.2004;Chapter2).

Theallopatricdistributionandquantifiedphenotypicdifferences

betweenthebisonsubspeciesareconsistentwiththe

subspeciesconcept.Nevertheless,therehasbeenasuggestion

thatthetwosubspeciesareactuallyecotypes,thatis,forms

exhibitingmorphologicaldifferencesthataresimplyareflection

oflocalenvironmentalinfluencesratherthanheritabletraits

(Geist1991).Thishypothesisisnotsupportedbyobservations

oftransplantedplainsandwoodbison.Woodbisontransplanted


fromtheiroriginalhabitatneartheNyarlingRiverinWBNPto

verydifferentenvironmentsintheMackenzieBisonSanctuary

(MBS)(in1963)andElkIslandNationalPark(EINP)(in1965)do

notdifferfromeachother,orfromlaterspecimenstakenfrom

theoriginalhabitat(vanZylldeJong1986;vanZylldeJonget

al.1995).Furthermore,despitethepassingofover40years,

theEINPwoodbison,whichliveunderthesameconditions

asplainsbisonresidingseparatelywithinthepark,showno

evidenceofmorphologicalconvergencewiththeplainsbison

form(vanZylldeJong1986;vanZylldeJonget al.1995).

Similarly,plainsbisonintroducedtoDeltaJunction,Alaska

(in1928)fromtheNationalBisonRange(NBR)haveclearly

maintainedthephenotypictraitsofplainsbison(vanZylldeJong

1992;vanZylldeJonget al.1995).Suchempiricalevidence

suggeststhatthemorphologicalcharacteristicsthatdistinguish

plainsandwoodbisonaregeneticallycontrolled(vanZyllde

Jonget al.1995).

HybridisationbetweenthesubspeciesinWBNPafteran

introductionofplainsbisonduringthe1920shascomplicated

theconsiderationofsubspeciesdesignations.Thecontroversial

decisiontomoveplainsbisonfromWainwrightBuffaloPark

(WBP)insouthernAlbertatoWBNP(from1925to1928)resulted

intheintroductionofdomesticbovinediseasestowoodbison

(Chapter5),andthreatenedthedistinctivenessandgenetic

purityofthesubspecies.In1957,CanadianWildlifeService

researchersdiscoveredapresumablyisolatedpopulationof

200woodbisonnearNyarlingRiverandBuffaloLake.The

researchersbelievedthatthisherdhadremainedisolatedfrom

thehybridherds,andtherefore,representedthelastreservoir

oforiginalwoodbison(BanfieldandNovakowski1960;Ogilvie

1979;VanCamp1989).Inanefforttosalvagethewoodbison

subspecies,bisonfromtheNyarlingherdwererelocatedto

establishtheMBSandEINPwoodbisonherdsinthe1960s.

LateranalysishasindicatedthattheNyarlingherd,andbison

elsewhereinWBNPandadjacentareas,didhavecontactwith

theintroducedplainsbison(vanZylldeJong1986;Aniskowicz

1990),butitwasminimalenoughthattheanimalscontinuedto

exhibitpredominatelywoodbisontraits(vanZylldeJonget al.

1995).Studiesontheimpactoftheplainsbisonintroduction

havedeterminedthatthehybridisationdidnotresultina

phenotypicallyhomogeneouspopulation,aswasfeared(vanZyll

deJonget al.1995).Sub-populationswithinWBNPdemonstrate

varyingdegreesofplainsbisontraitsdependingontheir

proximityto,oreaseofaccessfrom,theoriginalplainsbison

introductionsite(vanZylldeJonget al.1995).

Althoughdescriptivemorphologyandquantitativemorphometry

providesubstantialevidencesupportingthesubspecific

designations(vanZylldeJonget al.1995),earlyanalysisof

bloodcharacteristicsandchromosomalhomologydidnotdetect

adifference(PedenandKraay1979;Stormontet al.1961;Ying

andPeden1977).Preliminaryanalysisofgrowthregulating

geneswithinthetwosubspeciessuggeststhatthebison

subspecieshavereachedastageofevolutionarydivergence

duetogeographicisolation(Borket al.1991);however,under

theBiologicalSpeciesConcept,subspeciesmaybedefined

atthenextstageofspeciation,thatiswhenhybridoffspring

exhibitreducedfitness,whichdoesnotappeartobethecasein

WBNP(Borket al.1991).Furthermore,analysisofmtDNAfrom

NyarlingRiverwoodbisonandplainsbisondidnotproduce

monophyleticgroups(Strobeck1991;1992).This,however,does

notmeanthatthereisnodifference.Inisolatedpopulations,

mtDNAdivergesatarateof1to2%permillionyears(Wilsonet

al.1985).Itisestimatedthatthetwobisonsubspeciesdiverged

approximately5,000yearsago(vanZylldeJong1993;Wilson

1969),andhuman-inducedsubspecieshybridisationfurther

complicatedthephylogeny.Therefore,currentgeneticanalysis

techniquesmaynotbeabletodetectexistingdifferencesin

themitochondrialgenome.Inaddition,becausemtDNAis

maternallyinherited,mtDNAwithintheNyarlingRiverherd,as

wellasotherherdsinWBNP,reflectsthecontributionsfrom

maternalpopulations,whichhadabiasedrepresentationof

plainsbisoncows(Gateset al.2001).Therefore,theinability

todetectadifferencewithamoleculartestcomparinglimited

sequencesofgenomicmaterialdoesnotnecessarilymean

thereisnogeneticdifference;itmayjustbebeyondthecurrent

resolutionoftechnology.

RecentstudiesofDNAmicrosatellitesindicatethatthegenetic

distancesbetweenplainsbisonandwoodbisonaregreater

thanthosewithineitherofthetwosubspecies(Wilson2001;

WilsonandStrobeck1999).Thewoodbisonpopulationsstudied

formedadistinctivegrouponaNei’sminimumunrootedtree;a

stronggroupingdespitethepervasivehybridisationwithplains

bison(Wilson2001;WilsonandStrobeck1999).Wilsonand

Strobeck(1999)andWilson(2001)concludedsuchastrong

clusteringindicateswoodbisonandplainsbisonarefunctioning

asdistinctgeneticentities,andshouldcontinuetobemanaged

separately.Basedontheavailableevidence,Canada’sNational

WoodBisonRecoveryTeamconcluded:(1)historically,multiple

morphologicalandgeneticcharacteristicsdistinguishedwood

bisonfromtheplainsbison;(2)woodbisonandplainsbison

continuetobemorphologicallyandgeneticallydistinct,despite

hybridisation;and(3)woodbisonconstituteasubspeciesof

bison,andtherefore,shouldbemanagedseparatelyfromplains

bison(Gateset al.2001).

Theissueofsubspeciesdesignationsisrelevanttoconservation

inthatadecisiontocombineformsatthespecieslevelwould

invitehybridisationandeffectivelyeliminateanyevolutionary

divergencethathadoccurred.Establishingdefinitiverecognition

ofbisonsubspeciesiscomplicatedbyongoingchangeofgenus,

speciesandsubspeciesconcepts(Winston1999).However,

otherclassificationsandconcepts,suchastheevolutionarily

significantunit(ESU;Ryder1986),andgeneticandecological


exchangeability,movebeyondtraditionaltrinomialtaxonomy

toincorporateevolutionaryconsiderations.Conservation

biologistsarereconsideringdefinitionsofconservationunitsthat

incorporateboththehistoryofpopulationsreflectedinmolecular

analysis,andadaptivedifferencesrevealedbylifehistoryand

otherecologicalinformation(Crandallet al.2000;DeWeerdt

2002).Forexample,thegeminateevolutionaryunitidentifies

conservationunitsthataregeneticallysimilarbutecologically

orbehaviourallydistinct(Bowen1998).Crandallet al.(2000)

argueforabroadcategorisationofpopulationdistinctiveness

basedonnon-exchangeabilityofecologicalandgenetictraits.

Eachoftheseconceptspresentschallenges,asdoesany

conceptthatattemptstodividethebiologicalcontinuumfor

theconvenienceofhumaninterests.Essentially,differentiation

onanylevelwithinaspecieswarrantsaformaldecision

andrecognition.Ofnote,TheU.S.EndangeredSpeciesAct

recognisesthisconservationissueandprovidesforprotection

of“distinctpopulationsegments”.Similarly,theCommittee

ontheStatusofEndangeredWildlifeinCanada(COSEWIC),

whichisresponsibleforassessingthestatusofwildlife,includes

anyindigenousspecies,subspecies,varietyorgeographically

definedpopulationofwildfaunaorfloraasa“species”.

Whilethereappeartobesufficientgroundsforformal

recognitionofAmericanbisonsubspecies,thedebatemay

continue.This,however,shouldnotprecludeconservationof

thetwoformsasseparateentities(vanZylldeJonget al.1995;

WilsonandStrobeck1999).Regardlessofcurrentgenetic,

biochemicalorotherevidenceaboutthesubspeciesquestion,

therearenotablephenotypicdifferences,andpotentiallyother

typesofvariationthatmaynotbedetectablewithtechnologies

availableatthistime.Geneticistspredictthatgeneticanalysisin

thefuturewillbeabletobetteridentifygroupingswithinspecies

(Wilson2001).

Althoughgeneticandmorphologicalevidenceoftencorrespond,

thisisnotalwaysthecase(Winston1999).Thiscanleadto

debateoverrecognisingvariationthatcannotbemeasuredusing

alternativemorphologicalormolecularmethods.Nevertheless,

allformsofgeographicandecologicalvariationwithinaspecies

contributetobiodiversity(SecretariatoftheConventionon

BiologicalDiversity2000).Allvariantsofaspeciesmaycarry

evolutionarilyimportantecologicaladaptations(Chapter4),and

possessthepotentialtodevelopgeneticisolatingmechanisms

leadinginevolutionarytimetonewspecies(O’BrienandMayr

1991).Predictionofwhichvariantswillevolvetobecome

speciesisnotpossible;thisisanoutcomeofnaturalselection

andchance.Therefore,tomaintainbiodiversityandevolutionary

potential,itisimportanttonotdismissanyformofdifferentiation

withinaspecies,andtomaintaintheopportunityforevolutionary

processestofunction(Crandallet al.2000).Debatingwhethera

nameiswarrantedwithinarelativelyarbitrarytaxonomicsystem

doesnotabsolvehumansoftheresponsibilitytorecogniseand

maintainintraspecificdiversityastherawmaterialofevolution.

Theriskoflosingevolutionarypotentialsuggestsitwouldnotbe

prudenttoprematurelydismissexistinggroupingssuchasthe

plainsandwoodbison.


Chapter4 GeneticsLead Authors: Delaney P. Boyd, Gregory A. Wilson, James N. Derr, and Natalie D. Halbert

Asascience,populationgeneticsisconcernedwiththeorigin,

nature,amount,distributionandfateofgeneticvariationpresent

inpopulationsthroughtimeandspace.Geneticvariation

constitutesthefundamentalbasisofevolutionarychangeand

providesthefoundationforspeciestoadaptandsurvivein

responsetochangingintrinsicandextrinsicstressors.Therefore,

lossofgeneticdiversityisgenerallyconsidereddetrimental

tolong-termspeciessurvival.Intheshort-term,populations

withlowlevelsofgeneticdiversitymaysufferfrominbreeding

depression,whichcanincreasetheirprobabilityofextirpation

andreducefitness.Plainsandwoodbisonexperiencedsevere

andwell-documentedpopulationdeclinesinthe19thCentury

thatreducedthecensussizeofthisspeciesbyover99.99%.

Thespectacularrecoverytoaround430,000animalstoday

(Chapter7)isatestamenttotheirgeneticconstitution,and

representsoneofthemostsignificantaccomplishmentsin

modernconservationbiology.Americanbisonhave,however,

undergoneartificialhybridisationwithdomesticcattle,been

subjectedtodomesticationandartificialselection,andbeen

separatedintomanyrelativelysmallisolatedpopulations

occupyingtinyfractionsoftheiroriginalrange.Aswell,allwood

bisonpopulationscontainsomelevelofplainsbisongenetic

materialduetoartificialhybridisationbetweenthesubspecies.

Allofthesefactorshavehadaneffectonthecurrentlevelsof

geneticdiversityandontheintegrityofthebisongenome.As

aresult,preservationofbisongeneticdiversityisakeylong-

termconservationconsideration.Thefollowingsectionsdiscuss

someofthemajorissuesthatareimportantforthegenetic

managementofthisspeciesintothefuture.

4.1 reduction of Genetic Diversity

Withinspecies,geneticdiversityprovidesthemechanismfor

evolutionarychangeandadaptation(AllendorfandLeary1986;

Chambers1998;MeffeandCarroll1994;MittonandGrant

1984).Reductioningeneticdiversitycanresultinreduced

fitness,diminishedgrowth,increasedmortalityofindividuals,

andreducedevolutionaryflexibility(AllendorfandLeary1986;

BallouandRalls1982;Franklin1980;Frankhamet al.1999;

MittonandGrant1984;).Therearefourinterrelatedmechanisms

thatcanreducegeneticdiversity(heterozygosityandnumberof

alleles):demographicbottlenecks,foundereffects,geneticdrift,

andinbreeding(MeffeandCarroll1994).Unfortunately,overthe

lasttwocenturies,bisoninNorthAmericahave,tosomedegree,

experiencedallofthesemechanisms.

AsAmericanbisonapproachedextinctioninthelate1800s,

theyexperiencedaseveredemographicbottleneck,leadingto

aconcernthatextantbisonpopulationsmayhavelowergenetic

diversitythanpre-declinepopulations.Theconsequencesof

ageneticbottleneckdependonthepre-bottleneckgenetic

diversitywithinaspecies,theseverityofthedecline,andhow

quicklythepopulationreboundsafterthebottleneck(Meffeand

Carroll1994;Neiet al.1975).Thedeclineofbisonwassevere,

withareductionfrommillionstofewerthan1,000individuals.

Recoveryefforts,however,enabledbisonpopulationstogrow

quickly,morethandoublingbetween1888and1902(Coder

1975).Althoughtheeffectsofthebottleneckonthegenetic

diversityofthespeciesarenotclear(Wilson2001),thereare

severalpossiblerepercussions.First,afteraseverereductionin

populationsize,averageheterozygosityisexpectedtodecline

(Allendorf1986;Neiet al.1975).Heterozygosityisameasureof

geneticvariationthatisadirectreflectionofthepastbreeding

historyofapopulation.Heterozygosityvaluesareexpressedas

thefrequencyofheterozygotes(i.e.,geneswithdissimilaralleles)

expectedatagivenlocus(Griffithset al.1993).Areductionin

thelevelofheterozygositycanresultininbreedingeffects.At

thesametime,alossofallelesmaylimitapopulation’sability

torespondtonaturalselectionforcesandreducetheadaptive

potentialofapopulation(Allendorf1986;MeffeandCarroll1994;

Neiet al.1975;Robertson1960).

Afterthedemographiccrash,severalsmallbisonherds

remainedinNorthAmerica,manyofwhichwerederivedfrom

veryfewanimals.Overalllevelsofgeneticvariationincurrent

populationscan,intheory,varydirectlywiththenumberof

originalfounders(MeffeandCarroll1994;WilsonandStrobeck

1999).Remnantpopulationsmaynothavebeenrepresentative

oftheoriginalgenepooland,consequently,sufferedreduced

geneticvariability.Throughtime,thedetrimentaleffectsof

geneticdriftmayhavecompoundedtheeffectsoftheearlier

bottleneck.Geneticdriftinvolvestherandomchangeingene

frequenciesandleadstothelossofallelesovertime.Therate

ofthisloss,orfixationofalleles,isroughlyinverselyproportional

tothepopulationsize(Allendorf1986;MeffeandCarroll

1994).However,theactualcountofbreedingindividualsina

populationisnotappropriatefordeterminingtherateofgenetic

driftbecausefactorssuchasunequalsexratios,differential

reproductivesuccess,overlappinggenerations,andnon-random

matingresultinthe“effective”populationsizealwaysbeingless

thanthecensussize.Forbison,theratioofeffectivepopulation

size(Ne)tothecensuspopulationsize(N)hasmostcommonly


beenestimatedtobebetween0.16and0.42(Bergerand

Cunningham1994;ShullandTipton,1987;WilsonandZittlau,

2004),althoughShullandTipton(1987)suggestedthattheratio

couldbeaslowas0.09insomemanagedpopulations.

ItispossiblethatAmericanbisonexperiencedreductionsin

overallgeneticdiversityduetothepopulationbottleneckofthe

late1800s;however,thiseffectmaynothavebeenasgreatas

onceexpected.McClenaghan,Jr.et al.(1990)foundthatplains

bisonhavegreatergeneticvariabilitythanseveralothermammals

thatexperiencedseveredemographicbottlenecks.Furthermore,

WilsonandStrobeck(1999),Halbert(2003)andHalbertand

Derr(2008)foundlevelsofDNAmicrosatellitevariabilityinbison

populationstobesimilartootherNorthAmericanungulates.

Someauthorsspeculatethatpriortothebottleneck,the

Americanbison,withthepossibleexceptionofthewoodbison,

expressedsurprisinghomogeneitydespiteitsextensiverange

(Roe1970;Seton1910).Plainsbisonrangedoverlargeareas.

Thissuggeststhatextensiveanimalmovements,andthereby

geneflow,mayhaveexistedamongpopulations(Bergerand

Cunningham1994;WilsonandStrobeck1999).Similartoother

largemammals,bisonareexpectedtobelessgeneticallydiverse

thansmallmammals(SageandWolff1986).Despitefounder

effectsandlowgeneflow,whichincreasegeneticdistance

values,recentstudiesdemonstratethatthegeneticdistances

betweenexistingbisonherdsarelowerthanexpected,indicating

thatexistingisolatedpopulationsarelikelyderivedfromonelarge

genepool(WilsonandStrobeck1999).Furthermore,foundation

herdsforcontemporarybisonoriginatedfromacrossthespecies’

range,suggestingthatmuchofthepre-existingdiversitywas

likelyretained(Halbert2003).AnalysisofancientDNAmay

provideanopportunityforassessingpre-bottleneckgenetic

diversityforcomparativepurposes(Amos1999;Cannon2001;

Chambers1998).Unfortunately,itisnotpossibletorecoverthe

geneticmateriallostasaresultofthebottleneckunderscoring

theimportanceofmaintainingexistinggeneticdiversitywhile

minimisinganyfuturegeneticerosion.

Inbreeding,orthematingofrelatedindividuals,canleadtothe

expressionofdeleteriousalleles,decreasedheterozygosity,

lowerfecundity,anddevelopmentaldefects(Allendorfand

Leary1986;BergerandCunningham1994;Lande1999;Meffe

andCarroll1994).Inbreedingisdifficulttoassessanddoesnot

alwayshavemeasurabledeleteriousconsequences(Bergerand

Cunningham1994;MeffeandCarroll1994);however,itremains

apotentialcauseofreduceddiversityinbison.Todecrease

theeffectsofinbreeding,somebisonherdswerefoundedor

augmentedwithanimalsfromdifferentregions(Wilson2001).

Overtime,thetranslocationofanimalsamongherdsmayhave

reducedtheimpactsofinbreedingandfoundereffects,which

aremostsevereinisolated,smallpopulationswithlowlevels

ofgeneticdiversity.Whilefewbisonherdshavetrulyexhibited

signsthoughttobetheresultofinbreedingdepression,such

ashighratesofphysicalabnormalities,reducedgrowthrates,

andreducedfertility,inbreedingdepressionhasbeenlinkedto

lowlevelsofcalfrecruitmentandhighlevelsofcalfmortality

inaplainsbisonherd(Halbertet al.2004;2005),andhasbeen

suggestedtoaffectmalereproductivesuccessinanother

population(BergerandCunningham1994).

Althoughexistingbisonpopulationsmaybederivedfroma

largelyhomogeneousgenepool,recentstudiesusingDNA

microsatellitesrevealthatseveralplainsbisonherdsare

geneticallydistinguishable(HalbertandDerr2008;Wilsonand

Strobeck1999).Thisraisestheissueofwhetherconservation

herdsshouldbemanagedasalargemetapopulation,with

translocationofbisonamongherdstomaintainlocaldiversity,or

asclosedherdstopreserveemerginglocalizeddifferentiation.

Somepopulationsmaybeadaptingtonon-nativehabitats

orchangingconditionsinthenaturalenvironment,and

would,therefore,benefitfromlocalizeddifferentiation.Other

populationsmaybeadaptingto,orinadvertentlyselected

for,unnaturalconditions,andwouldbenefitfromperiodic

augmentation(Wilsonet al.2002b).Aprecautionaryapproach

maybetodiversifyconservationeffortsbytransferringrandomly

selectedanimalsamongsomeherds,tomaximiseintra-

populationgeneticdiversity,whilemaintainingotherherdsas

closedpopulationswiththepossibilityoftheestablishmentof

satellitepopulationstoincreaseoveralleffectivepopulationsizes

(HalbertandDerr2008).Managersshouldcarefullyconsider

theimplementationofmetapopulationmanagementplansasa

tooltopreservegeneticdiversityduetohistoricaldifferencesin

morphology,behaviour,physiology,anddiseasestatus(Lande

1999;RyderandFleischer1996;Wilsonet al.2002b)andtolimit

thespreadofdomesticcattlegenesbetweenbisonpopulations

(Halbertet al.2005a;2006).

Geneticanalysiscouldbeusedtomonitorgeneticdiversity

bybuildinganinventoryofdiversityheldwithinconservation

herds.Thereareseveralmeasuresofgeneticdiversityincluding

heterozygosity,allelesperlocus,andproportionofpolymorphic

loci(Amos1999;Templeton1994;Wilsonet al.2002b).While

earlyworkonbisongeneticsinvolvedbloodgroups(Stormont

1982;Stormontet al.1961),someauthorssuggestthatsuch

studiesareinappropriateforassessinggeneticdiversity

becauseselectionforbloodgrouptypemaybehigh,violating

theassumptionofselectiveneutrality(BergerandCunningham

1994;KnudsenandAllendorf1987;YamazakiandMaruyama

1974).Morerecentstudieshaveusedallozymes(Knudsenand

Allendorf1987;McClenaghanet al.1990),mitochondrialDNA

(MtDNA)(Polziehnet al.1996),nuclearDNArestrictionfragment

lengthpolymorphisms(Borket al.1991),andDNAmicrosatellites

(WilsonandStrobeck1999)toassessdiversity.Investigationof

individualgenomicregionscanreflectoveralldiversity,allowing

fordatafromvarioustechniquestobecombinedtoprovidean

accuraterepresentationofgeneticdiversity(Chambers1998).


Selectionfordiversityinonesystem,suchasbloodgroup

proteins,orbiasedselectionformaintainingspecificraregenetic

characteristicscouldleadtoreduceddiversityinotherparts

ofthegenome(Chambers1998;Hedricket al.1986).Biased

selectionformaintainingrareallelesisespeciallyquestionable

ifitisnotknownwhattherarealleledoes,orifitisdetrimental

(i.e.,itmayberarebecauseitisbeingexpungedfromthe

bisongenomethroughnaturalselection).Variationthroughout

thegenome,ratherthanthemaintenanceofonespecificrare

allele,conveysevolutionaryflexibilitytoaspecies(Chambers

1998;VrijenhoekandLeberg1991).Therefore,itiscrucialfora

geneticmanagementplantoconsiderallavailablemeasuresfor

managinggeneticdiversityinthepoliciesandproceduresfor

breedingandcullingdecisions.

Anassessmentofoverallgeneticdiversityshouldexamine

atleast25-30locidistributedacrossthenucleargenome

(Chambers1998;Nei1987).Whilegeneticdiversityforsome

herdshasbeenassessed(Baccuset al.1983;Bergerand

Cunningham1994;KnudsenandAllendorf1987;Wilsonand

Strobeck1999),thesestudiesdidnotincludeasufficient

numberoflociandcomparisonsbetweenstudiesarenot

possibleduetodifferencesinmarkersystems(allozymesvs.

microsatellites).Otherstudieshaveincludedlargernumbersof

lociandpopulations;however,severalconservationherdshave

notbeenfullyexamined(e.g.,someU.S.,CanadianandMexican

state,federalandprivatebisonherds;Halbert2003;Halbertand

Derr2008).Clearlyitisimportanttocreateamorecomplete

assessmentofbisongeneticdiversitytoallowformoreinformed

managementdecisions.

Ingeneral,maintaininggeneticdiversityofAmericanbison

requiresanunderstandingofherdpopulationdynamicsto

assesstheprobabilityoflong-termpersistenceofthatdiversity.

Mostbisonpopulationsarecomposedoffewerthan1,000

individuals,anditispossibleforarelativelysmallnumberof

dominantmalestoberesponsibleforahighpercentofthe

matinginagivenyear(BergerandCunningham1994;Wilson

et al.2002;Wilsonet al.2005;Halbertet al.2004).This,in

turn,canreducegeneticdiversityovertime,especiallyin

theabsenceofnaturalmigrationandexchangeofgenetic

diversityamongpopulations(BergerandCunningham1994).

Thepotentialfordisproportionatereproductivecontributions

emphasisestheimportanceofmaintaininglargeherdswith

largeeffectivepopulationsizes,thatgivenpropermanagement,

willpreventlossofgeneticdiversity(Frankham1995;Franklin

1980).Assessmentofgeneticuncertainty,basedonNe,founder

effects,geneticdrift,andinbreeding,isarequiredcomponent

ofapopulationviabilityanalysis(PVA)(GilpinandSoulé1986;

Shaffer1987).

4.2 hybridisation

Hybridisationinvolvestheinterbreedingofindividualsfrom

geneticallydistinctgroups,whichcanrepresentdifferent

species,subspecies,orgeographicvariants(Rhymerand

Simberloff1996).Someauthorsarguethathybridisationisa

potentiallycreativeevolutionaryforce,whichgeneratesnovel

combinationsofgenesthatcanhelpspeciesadapttohabitat

change,althoughsuchhybridsoftenexperiencereducedfitness

(AndersonandStebbins1954;LewontinandBirch1966;Hewitt

1989).Hybridisationthroughartificialmanipulationorrelocation

ofanimals,however,cancompromisegeneticintegritythrough

geneticswampingofonegenomeoveranotheranddisruption

oflocallyadaptedgenecomplexes(Avise1994).Itcanalso

produceoffspringthataredevaluedbytheconservationand

legalcommunities(O’BrienandMayr1991;Chapter7).The

geneticlegacyofintroducingplainsbisonintoawoodbison

populationinnorthernCanada,andcrossbreedingbisonand

cattle,havemadehybridisationacontroversialtopicinbison

conservation.

4.2.1 plains bison x wood bison

Basedontheirgeographicdistributionandmorphology,plains

bisonandwoodbisonwerehistoricallydistinctentities(Chapter

3).Itcanbearguedthattheintroductionofplainsbisoninto

rangeoccupiedbywoodbisonwasa“negligibletragedy”(Geist

1996),becausesomeconsiderthetwogroupstobeecotypes

(Geist1991).Othersmaintainthattheinterbreedingofthese

twotypesshouldhavebeenavoidedtopreservegeographic

andenvironmentalvariation(vanZylldeJonget al.1995).The

introductionofeithersubspeciesintotheoriginalrangeofthe

othercould,intheory,erodethegeneticbasisofadaptation

tolocalenvironmentalconditions(Lande1999).Therefore,

hybridisationbetweenplainsandwoodbisonshouldbe

considereddetrimentaltomaintainingthegeneticintegrityand

distinctivenessofthesetwogeographicandmorphologically

distinctforms.

Whilehistoricallytheremayhavebeennaturalhybridisation

eventsbetweenthesubspeciesinareasofrangeoverlap,the

currenthybridisationissueistheconsequenceofanill-advised

andirreversibledecisionmadenearly85yearsago.In1925,

theCanadiangovernmentimplementedaplantomovemore

than6,000plainsbisonfromtheovercrowdedWainwright

NationalParktoWoodBuffaloNationalPark(WBNP).Biological

societiesfromU.S.andCanadastrenuouslychallengedthis

action,asinterbreedingwouldeliminatethewoodbisonform,

resultinghybridsmightnotbeasfitfortheenvironment,and

diseasessuchasbovinetuberculosis(BTB)wouldspreadto

formerlyhealthyanimals(Howell1925;Harper1925;Lothian

1981;Saunders1925).Proponentsoftheplancounteredthe

criticismbyquestioningthesubspeciesdesignations,arguing


4.2.2 Domestic cattle x bison

Theconceptofcrossingbisonwithdomesticcattledatesback

toSpanishcolonisersofthe16thCentury(Dary1989).There

aremanyaccountsofhistoricalattemptstohybridisebison

andcattle(Coder1975;Dary1989;Ogilvie1979;McHugh

1972;Ward2000).Privateranchersinvolvedwithsalvaging

bisonhadaspirationstocombine,throughhybridisation,the

hardinessandwinterforagingabilityofbisonwiththemeat

productiontraitsofcattle(Dary1989;Ogilvie1979;Ward2000).

TheCanadiangovernmentactivelypursuedtheexperimental

productionofcrossbredanimalsfrom1916-1964(Ogilvie1979;

Polziehnet al.1995).

Historicalcrossbreedingattemptshavecreatedalegacy

ofgeneticissuesrelatedtotheintrogressionofcattleDNA

intobisonherds.Introgressionreferstogeneflowbetween

populationscausedbyhybridisationfollowedbybreedingof

thehybridoffspringtoatleastoneoftheirrespectiveparental

populations(RhymerandSimberloff1996).Theintrogressed

DNAreplacessectionsoftheoriginalgenome,thereby

affectingthegeneticintegrityofaspecies,andhamperingthe

maintenanceofnaturalgeneticdiversity.Manycontemporary

bisonherdsarefoundedon,andsupplementedwith,animals

fromherdswithahistoryofhybridisation(Halbert2003;Halbert

et al.2005a;2006;Wardet al.1999;2000).Thisextensivehistory

ofhybridisationbetweenthesetwospeciesraisesquestions

abouttheintegrityofthebisongenomeandthebiological

effectsofcattleDNAintrogression.

Fertilityproblemsthwartedmanyoftheoriginalcrossbreeding

attemptsbecausecrossesresultinhighmortalityforoffspring

andmother(Ward2000).Experimentationhasrevealedthat

crossesofbisonfemaleswithdomesticcattlemalesproduce

lessmortalityintheoffspringthanthemoredeadlyreverse

thattheintroductionsitewasisolatedfrom,

andunusedby,thewoodbisonpopulation,

andsuggestingthattheintroducedanimals

weretooyoungtocarryBTB(Fuller2002;

Graham1924).Theseargumentsdidnot

considerthefuturehabitatneedsofthe

growingwoodorplainsbisonpopulations,

northelikelihoodthatthetwosubspecies

wouldnotremainisolated.Aswell,a

recommendationthatonlyyearlingsthat

passedatuberculintestbeshippedto

WBNPwasrejected(Fuller2002).

Itwasnotuntil1957thatthediscoveryof

aseeminglyisolatedherdof200animals

neartheNyarlingRiverandBuffaloLake

alleviatedfearsthatwoodbisonwaslost

tohybridisation(vanCamp1989).CanadianWildlifeService

researchersdeterminedthattheseanimalsweremorphologically

representativeofwoodbison(BanfieldandNovakowski1960).

Tosalvagethewoodbisonsubspecies,bisonfromtheNyarling

herdwerecapturedandrelocatedtoestablishtwonewherds.

SixteenanimalsweremovedtotheMBSnorthofGreatSlave

Lakein1963(Fuller2002;Gateset al.2001c),and22animals

weresuccessfullytransferredtoElkIslandNationalPark(EINP)

eastofEdmonton,Albertain1965(BlythandHudson1987).

TwoadditionalcalvesweretransferredtoEINPbetween1966

and1968(BlythandHudson1987;Gateset al.2001c).Ofthose

bisontransferred,11neonatesformedthefoundingherd.

Subsequentstudiesrevealedthattherewascontactbetween

theNyarlingherdandtheintroducedplainsbison(vanZyllde

Jong1986).AlthoughhybridisationwithinWBNPdidnotresult

inaphenotypicallyhomogenouspopulation(vanZylldeJong

et al.1995),geneticdistancesamongsubpopulationsinthe

parkaresmall,indicatingthatthereisgeneflowandinfluence

oftheplainsbisongenomethroughoutallregionsofthepark

(Wilson2001;WilsonandStrobeck1999).Despitehybridization,

geneticdistancesbetweenplainsandwoodbisonaregenerally

greaterthanthoseobservedwithinsubspecies.Moreover,wood

bisonformageneticgroupingonaNei’sminimumunrooted

tree,suggestinggeneticuniqueness(Wilson2001;Wilsonand

Strobeck1999).

Morphologicalandgeneticevidencesuggestthatcareshould

nowbetakentomaintainseparationbetweenthesehistorically

differentiatedsubspecies.Effortsareinplacetoensure

representativewoodbisonandplainsbisonherdsareisolated

fromeachothertopreventfuturehybridisationbetweenthese

importantconservationherds(Harperet al.2000).

plate 4.1 Hereford x bison hybrid; cattle gene

introgression is morphologically evident. Photo:

Bob Heinonen.


cross,however,thelatterismorecommonbecauseitisvery

difficulttocompeldomesticcattlebullstomatewithbison

females.AllF1generationhybridsexperiencereducedfertility

andviabilityrelativetoeitherparent:F1malesaretypically

sterile,butthefertilityofF1femalesmakesintrogressive

hybridisationpossible(Ward2000).Geneticstudieshavefound

noevidenceofcattleY-chromosomeintrogressioninbison,

whichissupportedbythesterilityofF1hybridmalesfromthe

crossofcattlemaleswithbisonfemales,andbythebehavioural

constraintpreventingdomesticbullsfrommatingwithfemale

bison(Ward2000).

However,anumberofstudiesusingmodernmoleculargenetic

technologieshavereportedbothmtDNAandnuclearDNA

introgressioninplainsbisonfromdomesticcattle.Thefirst

ofthesestudies(Polziehnet al.1995)foundcattlemtDNA

amongCusterStateParkplainsbison.Subsequently,more

comprehensiveexaminationsofpublicbisonherdsrevealed

cattlemtDNAinsevenof21bisonconservationherds

(Ward2000;Wardet al.1999),suggestingthathybridisation

issuesbetweenthesetwospecieswerewidespreadanda

significantconcerntolong-termbisonconservationefforts.

Furtherinvestigationsbasedonhigh-resolutionnuclearDNA

microsatellitesdetecteddomesticcattlenuclearDNAmarkersin

14ofthese21U.S.federalconservationherds(Ward2000).

AllmajorpublicbisonpopulationsintheU.S.andCanadahave

nowbeenexaminedusingmtDNA,microsatellitemarkers,or

acombinationofthesetwotechnologies.Combiningevidence

frombothmtDNAandnuclearmicrosatellitemarkerswith

informationregardingpopulationhistoriesprovidesamore

completeviewofhybridisationbetweenthetwospecies.To

date,nogeneticevidenceofdomesticcattleintrogressionhas

plate 4.2 Custer State Park plains bison

bull; a high level of cattle gene introgression

is not morphologically evident. Photo:

Cormack Gates.

beenreportedinnineoftheseconservationpopulations(plains

bisonunlessotherwisenoted;n=samplesizeexamined):EINP

(woodbison,n=25);MBS(woodbison,n=36);WBNP(wood

bison,n=23);EINPplainsbison(n=25);GTNP(n=39);HMSP

(n=21);SHNGP(n=31);WindCaveNationalPark(WCNP)(n=

352);andYNP(n=520)(Halbertet al.2005a;2006;Wardet al.

1999).

However,theabilitytodetectnuclearmicrosatelliteDNA

introgressionishighlydependentonthenumberofbisonineach

population,thenumberofbisonsampledfromeachpopulation

andtheactualamountofdomesticcattlegeneticmaterial

presentinthepopulation(Halbertet al.2005a).Considering

statisticalconfidence(greaterthan95%)allowedbydetection

limitsofthetechnology(Halbertet al.2006),adequatenumbers

ofbisonhavebeenevaluatedfromonlytwooftheseherds

thatdisplayednoevidenceofhybridisation(WCNPandYNP).

Thesetwoherdsrepresentlessthan1.0%ofthe420,000plains

bisoninNorthAmericatoday(Freeseet al.2007;Chapter7)

andbothoftheseherdsarecurrentlyprovidinganimalsforthe

establishmentofnewsatelliteherdsforconservationsefforts

(Chapter7).Furtherevaluationisurgentlyneededtomore

accuratelyassesslevelsofdomesticcattlegeneticsinother

publicbisonherds.

Hybridisationissueswithdomesticcattlemustbeconsidered

alongwithothergeneticandnon-geneticfactorsindetermining

whichpopulationsaredesignatedas‘conservationherds’.

Forexample,althoughsomepublicherdsareknowntohave

lowlevelsofdomesticcattlegenetics,theseherdsmayalso

representdistinctlineagesthatreflecthistoricalandgeographic

differencesingeneticdiversity(Halbert2003;HalbertandDerr

2006;HalbertandDerrsubmitted).Cautionisneededinlong-

termconservationplanningtoensure

thatgeneticdiversitythatrepresents

historicalbisongeographicdifferences

isidentifiedandconservedforall

importantpopulationsandnotjust

thosethoughttobefreeofdomestic

cattleintrogression.Nevertheless,

defininggenetichistoriesthat

includehybridisationisafirst

stepindevelopingaspecies-wide

conservationmanagementplan.Given

thatthereareseveralsubstantialbison

herdsthatappeartobefreeofcattle

geneintrogression,itisofparamount

importancetomaintaintheseherdsin


reproductiveisolationfromherdscontaininghybrids.

4.3 Domestication

Thenumberofbisonincommercialherdshasgrownrapidlyover

thepastfivedecadesasmanyranchersenterthebisonindustry

tocapitaliseontheeconomicopportunitiesofferedbythis

species(Dey1997).Theincreaseincommercialbisonproduction

mayreflecttherecognitionoftheadvantagesaffordedbythe

adaptationsandecologicalefficiencyofbisonasanindigenous

rangeanimal.Bisonpossessseveraltraitsthatmakethem

preferabletocattleasarangeanimal,includingagreaterability

todigestlowqualityforage(Hawleyet al.1981;Plumband

Dodd1993),theabilitytodefendagainstpredators(Carbynet

al.1993),theabilitytosurviveharshwinterconditions,andalow

incidenceofcalvingdifficulties(Haighet al.2001).Accordingto

federalgovernmentsurveys,thecommercialbisonpopulation

inNorthAmericaisabout400,000,dividedalmostequally

betweentheU.S.andCanada(Chapter7).Despitethecurrent

plateauinbeefandbisonmeatprices,boththeCanadianBison

AssociationandtheU.S.-basedNationalBisonAssociation

predictveryfavourablelong-termgrowthofthebisonindustry.

Thenumberofbisoninconservationherdsiscurrentlyestimated

atonly20,504plainsbisonand10,871woodbison.Therefore,

approximately93%ofAmericanbisonareundercommercial

productionandexperiencingsomedegreeofdomestication.

Domesticationisaprocessinvolvingthegenotypicadaptationof

animalstothecaptiveenvironment(Price1984;PriceandKing

1968).Purposefulselectionoverseveralgenerationsfortraits

favourableforhumanneeds,resultsindetectabledifferences

inmorphology,physiology,andbehaviourbetweendomestic

speciesandtheirwildprogenitors(Darwin1859;Clutton-Brock

1981;Price1984).Humanshavepracticeddomesticationof

livestockspeciesforatleast9,000years(Clutton-Brock1981).

Asagricultureprecipitatedthesettlementofnomadichuman

cultures,thedomesticationofseveralwildmammalspecies

madelivestockfarmingpossible(Clutton-Brock1981).Intensive

managementpracticesandcompetitionbetweendomesticated

animalsandtheirwildancestorsoftenpushedwildvarieties

andpotentialpredatorstotheperipheryoftheirrangesorto

extinction(BaerselmanandVera1995;Hartnettet al.1997;

Price1984).Examplesofextinctancestorsofdomesticated

animalsincludethetarpan(Equus przewalski gmelini),the

wilddromedary(Camelus dromedarius),andtheaurochs(Bos

primigenius)(BaerselmanandVera1995).

Thedomesticationofcattleprovidesarelevanthistoryfrom

whichtoconsidertheissuesofbisondomestication.Before

cattle(Bos taurus)wereintroducedtoNorthAmericatheyhad

experiencedthousandsofyearsofcoevolutionwithhuman

culturesinEurope(Clutton-Brock1981;Hartnettet al.1997).

Duringthedomesticationprocesscattlewereselectedfor

docilityandvaluedmorphologicalandphysiologicaltraits,

butnotwithoutadverseconsequences.Geneticselectionhas

producedananimalthatisdependentonhumans,isunableto

defenditselfagainstpredators,andhasanatomicalanomalies,

suchasasmallerpelvicgirdle,whichcausecalvingandwalking

difficulties(Kampf1998;Knowleset al.1998;Pauls1995).

Domesticationhasalteredthewildcharacterofcattle,producing

animalsmaladaptedtothenaturalenvironment.Furthermore,

becausetheaurochs,thewildancestorofEuropeandomestic

cattle,becameextinctin1627(Silverberg1967),domesticcattle

havenowildcounterparttoprovideasourceofgeneticdiversity

forgeneticenhancementandmaintenance.

Whileithasbeensuggestedthatdomesticatedanimalscan

bereintroducedintothewildandreverttoaferalstate(Kampf

1998;Lott1998;Turnbull2001),suchattemptsdonotrestore

theoriginalgeneticdiversityofaspecies(Price1984;vanZyll

deJonget al.1995).Experiencehasshownthatrecoveryof

originalgeneticdiversityisdifficultorimpossibleoncedomestic

breedsarehighlyselectedforspecifictraitsandwildstocksare

extinct(Price1984;Turnbull2001;vanZylldeJonget al.1995).

Forexample,inthe1920s,twoGermanbrothers,Heinzand

LutzHeck,setoutto“re-create”theaurochsbyback-breeding

domesticcattlewithothercattledemonstratingaurochs-like

qualities(Fox2001;Silverberg1967;Turnbull2001).They

producedonesuccessfulline,theHellabrunnbreed,alsoknown

asHeckcattle.Thisisananimalthatlooksverymuchlikean

aurochs,butisdevoidofthewildtraitsandhardinessofthe

originalwildform(Fox2001;Silverberg1967).Thisillustrates

thattheoriginalwildgenotypeisnolongeravailabletothe

cattleindustryforimprovingdomesticbreeds.Thehistoryof

theaurochsoffersalessonforbison:domesticationcanlead

toalteredgeneticallybasedbehaviour,morphology,physiology,

andfunction,andthelossofthewildtypeandthegenetic

diversityitcontains.

Theprimarygoalofmanycommercialbisonranchersisto

increaseprofitsbymaximisingcalfproduction,feed-to-meat

conversionefficiency,andmeatquality(Schneider1998).

Thisrequiresnon-randomselectionfortraitsthatservethis

purpose,includingconformation,docility,reducedagility,growth

performance,andcarcasscomposition.Selectionforthese

traitsreducesgeneticvariationandchangesthecharacterof

theanimalovertime(Schneider1998).Althoughagrowing

numberofconsumersprefernaturallyproducedmeatproducts

withouthormones,antibiotics,orintensivemanagement(Morris

2001),thedemandforbisoncannotcurrentlycompetewiththe

muchlargerscaleofthebeefindustry.Therefore,manybison

producersapplycattlehusbandrypracticesandstandardsto

bison.Artificialselectionbasedonhusbandryandeconomics

maymakegoodbusinesssenseintheshortterm,butitwillnot

conservenativebisongermplasm.


Thelongtermobjectivesandgoalsthatdrivecommercialbison

productiongenerallydifferfromthemajorissuesassociatedwith

theconservationofthewildspecies.Furthermore,commercial

bisonoperationscouldposeathreattoconservation

populationsthroughaformofgeneticpollutionifgenetically

selectedcommercialanimalsaremixedintoconservationherds

orescapeandjoinwildherds.Themostprudentactionisto

identifyandmaintainexistingconservationherds,andavoid

mixingcommerciallypropagatedstockintothoseherds.Bison

producersandthebisonindustrycouldbenefitinthelongterm

bysupportingeffortstorestoreandmaintainconservationherds,

particularlythosesubjecttoafullrangeofnaturalselection

pressures(Chapter7).Conservationherdssecurethebison

genomeforthefutureuseofproducers—anoptionnotavailable

formostotherdomesticanimals.


Chapter5 ReportableorNotifiableDiseases

Contributors: Brett T. Elkin, Martin Hugh-Jones, Damien O. Joly, and John Nishi.

Throughouttheirrange,bisonhostnumerouspathogensand

parasites,manyofwhichalsooccurindomesticcattle(see

reviews:Berezowski2002;Tessaro1989;Reynoldset al.2003).

Inthisreview,weconsideronlyinfectiveorganismsthatmay

negativelyaffectbisonpopulations,ortheirconservation,

eitherthroughdirectpathobiologicaleffects,orindirectlyasa

consequenceofmanagementinterventions.Livestockdiseases

thatrestricttradeorposearisktohumanhealthmaybe

“reportable”or“notifiable”underfederalandprovincial/state

legislation.

InCanada,reportableandimmediatelynotifiablediseasesare

listednationallyundertheauthorityoftheHealthofAnimalsAct

andRegulations(http://laws.justice.gc.ca/en/H-3.3/,accessed

15April2009)andunderprovincialstatutesandlegislation.The

CanadianHealthofAnimalsActrequiresownersandanyone

caringforanimals,orhavingcontroloveranimals,toimmediately

notifytheCanadianFoodInspectionAgency(CFIA)whenthey

suspectorconfirmthepresenceofadiseaseprescribedinthe

ReportableDiseasesRegulations.TheCFIAreactsbyeither

controllingoreradicatingthediseasebaseduponaprogramme

agreedtobystakeholders(CFIA2001).

IntheU.S.,theU.S.DepartmentofAgricultureAnimalandPlant

HealthInspectionService(APHIS)conductsfederaleradication

programmesforseveralreportablelivestockdiseasesand

isinvolvedinanegotiatedmulti-jurisdictionalbrucellosis

managementprogrammeforbisoninYellowstoneNationalPark

(YNP)(APHIS,USDA2007;NPS-USDOI2000).Inbothcountries,

Federallegislationsupersedesstateandprovincialdisease

controllegislation.IntheU.S.andCanadatherearespecific

stateandprovincialregulationsthatrequiretestingfor,and

reportingof,variousdiseases.Theseregulationsmaybemore

extensivethanfederalrequirements,buttypicallyincludethose

diseasesregulatedbythefederalanimalhealthauthorities.

MuchliketheU.SandCanada,Mexicohasfederalanimal

diseaseregulationsthatareadministeredbytheSecretaryof

Agriculture,LivestockProduction,RuralDevelopment,Fishery

andFood(SAGARPA).Diseasesurveillanceprogrammesand

zoosanitaryrequirements,includingdiseasereporting,are

establishedbyfederallawtoprotecttradeinMexicoandare

administeredbyadecentralisedbranchofSAGARPAtitledthe

NationalServiceofHealth,Safety,andAgriculturalFoodQuality

(SENASICA,seehttp://www.senasica.gob.mx).SAGARPA

alsonegotiatesbi-lateraldiseasemanagementagreementsfor

importantlivestockdiseasesalongtheU.S.border,including

bovinetuberculosis,brucellosis,andscrewworm.

Inadditiontofederal,stateandprovincialregulatoryagencies

thereisaninternationalorganisationthatinfluencesanimal

diseasereportinginNorthAmerica.TheWorldOrganization

forAnimalHealth(OIE)isanintergovernmentalorganisation

createdbyinternationalagreementin1924.In2008theOIEhad

172membercountries.Everymembercountryiscommittedto

declaringtheanimaldiseasesitmaydetectinitsterritory.The

OIEdisseminatesthisinformationtohelpmembercountries

toprotectthemselvesfromthespreadofdiseaseacross

internationalboundaries.TheOIEproducessanitarycodeswith

rulesthatmustbeobservedbymembercountriestoprevent

thespreadofsignificantdiseasesaroundtheworld.OIEhas

establishedSanitaryCodesforTerrestrialAnimals,andthe

ManualforDiagnosticandVaccineTestsforTerrestrialAnimals,

whichmayinfluencetheinternationalmovementofbison(http://

www.oie.int/eng/normes/mcode/en_sommaire.htm).Allthree

countriesinNorthAmericaaremembersofOIE.

Dependingonthenatureofthedisease,managementof

reportablediseasesincaptiveorcommercialherdsinNorth

Americamayinvolvedevelopmentandapplicationofuniform

protocolstoreducediseaseprevalence,zoningofmanagement

areasbydiseasestatus,orimpositionofproceduresfordisease

eradication,includingtestandslaughter,ordepopulation.Where

reportablediseasesaredetected,federal,stateorprovincial

legislationaffectsmanagementofwildbisonpopulations.

Interventionsmayincludelimitingthegeographicdistributionof

aninfectedwildpopulation,(e.g.,removalsatparkboundaries

toreducetheriskofthediseasespreadingtoadjacentlivestock

population),quarantine,treatment,oreradicationofinfected

captiveconservationbreedingherds,orlimitinginter-population

orinter-jurisdictionaltransportofbison.Publicperception

ofbisonasspecific,ornon-specific,carriersofdiseases

isapotentialbarriertore-establishingconservationherds,

particularlyinregionswhereconventionallivestockgrazing

occurs.Nationalandstate/provincialgovernmentsmayrestrict

theimport/exportofbisonforconservationprojectsbasedon

realorperceivedrisksofinfectionandtransmissionofreportable

diseases.

Lead Authors: Keith Aune, C. Cormack Gates and Delaney Boyd.


5.1 Diseases of Conservation Concern

TheAmericanBisonSpecialistGroup(ABSG)recognisesnine

federallylisteddiseasesofconcernforbisonconservationin

NorthAmerica.Regulationsapplicabletoeachdiseasemayvary

amongjurisdictionsandintheirimpactonbisonconservation

andrestorationefforts.TheOIElistssevenofthesediseasesas

“notifiable”underinternationalstandards.

5.1.1 Anaplasmosis

TheetiologicagentofanaplasmosisisAnaplasma marginale,

arickettsiathatparasitisestheredbloodcellsofhostanimals.

Theorganismistransmittedbybloodsuckinginsects,such

asticks,whichserveasavectorbetweenhosts(Radostits

et al.2000).Theinterplayofsusceptiblewildruminants

andarthropodvectorsiscriticaltotheepizootiologyofthe

disease.Anaplasmosisisadiseaseofinternationalregulatory

concernand,therefore,significantlyimpactslivestocktrade

betweenCanadaandthenorth-centralandnorth-westernU.S.

Anaplasmosisisadiseaseofmajoreconomicimportanceto

thecattleindustryininfectedregions.Bisonareknownhosts

ofA. marginale(Zaug1986)andwildbisonhavedemonstrated

serologictitresforthedisease(Tayloret al.1997).Theyhave

alsobeenexperimentallyinfected(Kocanet al.2004;Zaugg

1986;ZauggandKuttler1985).Serodiagnosisinwildungulates

hasprovenlargelyunreliable,butmodernmoleculardiagnostic

procedureshaveprovidedanexcellentalternative(Davidsonand

Goff2001).Naturallyoccurringinfectionshavebeenreported

intheNationalBisonRange(NBR),Montana,where15.7%

ofbisontestedpositiveforanaplasmosis(ZauggandKuttler

1985).RecentstudiesdemonstratedA. marginaleinfectionin

twowidelyseparatedbisonherdsintheU.S.,oneinOklahoma

(NatureConservancyTallgrassPrairiePreserve)andonein

Saskatchewan(DeLaFuenteetal.2003).IntheCanadianherd,

serologyandpolymerasechainreactionsindicatedthat10

individualswereinfectedwithA. marginalewhereas42of50

bisonculledfromtheTallgrassPrairiePreserve(TGPP)tested

positiveserologicallyascarriersofA. marginale.TheU.S.bison

isolateofA. marginalewasfoundtobeinfectivewheninoculated

intosusceptiblesplenectomisedcalves.Clinicalsymptomsin

bisonaresimilartothosedescribedforcattle.Theyinclude

anaemia,jaundice,emaciation,anddebility(Radostitset al.

2000).Experimentallyinfectedbisoncalvesdemonstratedmild

clinicalsignssuggestingthatbisonmaybemoreresistantthan

cattle(ZauggandKuttler1985).Thediseaseoccurscommonly

inAfrica,theMiddleEast,Asia,Australia,theU.S.,Central

andSouthAmerica,andsouthernEurope.Ifanaplasmosis

isdiagnosedinCanadiancattleorbison,Canada’scurrent

foreignanimaldiseasestrategycallsforitseradicationthrough

thetestingofinfectedandexposedherdsandtheremovalof

infectedindividuals.EverybisonimportedintoCanadafrom

theU.S.mustbequarantinedfromthetimeofitsimportation

intoCanadauntilitprovesnegativetotestsforanaplasmosis

performedatleast60daysafteritwasimportedintoCanada

(CFIA2007).Programmesformanagingthisdiseaseindomestic

animalsincludevectorcontrol,vaccinationandantibiotictherapy

(DavidsonandGoff2001).Anaplasmosisisnotinfectiousto

humans.

5.1.2 Anthrax

Anthraxisaninfectiousbacterialdiseasecausedbythe

endospore-formingbacteriumBacillus anthracis(Dragonand

Rennie1995).Afterinhalationoringestionbyasusceptible

host,B. anthracisendosporesgerminateandthevegetative

formofthebacteriumreplicatesinthebloodstream,releasing

toxinsthatcausesepticaemiaanddeath(DragonandRennie

1995;Gateset al.2001b).Uponreleasefromacarcass,highly

resistantendosporescanremainviableinthesoilfordecades

beforeinfectinganewhost(DragonandRennie1995).Humans

haveplayedanimportantroleintheevolutionofanthraxby

increasingtheproliferationanddispersalofthisglobalpathogen.

Observationsoftheroleofclimaticfactors,suchasseasonof

year,ambienttemperature,anddroughtinpromotinganthrax

epizooticshavebeenmadeforseveraldecades(APHIS,USDA

2006).Thecommonalityofsummermonths,highambient

temperatures,drought,andanthraxepizooticsarenon-

contentious.Therolesofenvironmentalfactorssuchassoil

typesandsoildisturbancesviaexcavationarepoorlydefined

despiteattemptstoevaluatethesepotentialfactors.

Bacillus anthracisisdividedintothreegenotypebranches

withdistinctgeographicsub-lineagecompositionsthatvary

regionallyaroundtheglobe(VanErt2007).VanErt(2007)

analysed273isolatesofB. anthracisinNorthAmerica,reporting

acosmopolitanassortmentof44multiplelocus,variable

number,tandemrepeatanalysisgenotypes.Onehypothesis

holdsthatB. anthraciswasintroducedfromtheOldWorldto

theNewWorldinspore-infectedanimalproducts(wool,skins,

bonemeal,shavingbrushes)transportedtothesouth-eastern

seaboardduringtheEuropeancolonial-era(Hanson1959;Van

Ness1971).Consistentwiththishypothesis,VanErt(2007)

foundasingledominantsub-groupinNorthAmerican(A.Br.

WNA;70%ofgenotypes)thatiscloselyrelatedtothedominant

Europeansub-groupA.Br.008/009.Thediversityofsub-lineages

representedvariesgeographicallyinNorthAmerica.A.Br.WNA

predominatesinthenorth,whiletheindustrialisedsouth-eastern

regionofthecontinentcontainsacosmopolitanassortmentof

lesscommonB. anthracisgenotypesinadditiontothedominant

formA.Br.WNA.

Thegeographicpatternofsub-lineageoccurrenceinNorth

Americaisconsistentwiththehypothesisofanearlyinitial

introductionofalimitednumberofsub-lineages(perhaps


one)followedbyitswidespreaddispersalandecological

establishment.Wildbisonwereabundantandwidelydistributed

atthetimeofEuropeancolonisation.Onceinfectedwithanthrax

theymayhaveplayedanimportantearlyroleintheecological

establishmentandwidespreaddispersalofA.Br.WNA.Thebroad

diversityofanthraxlineagesrepresentedintheindustrialised

south-easternregionofthecontinent(VanErtet al.2007)is

suggestiveoftheaccumulationofadditionalsub-grouptypes

overtime.Alikelymechanismisimportationofcontaminated

animalproductsintomillsandtanneriesontheeasternseaboard

andNewEnglandwhichprocessimportedhair,wool,andhides.

TheWorldHealthOrganisation(WHO2008)commentedon

theroleoftanneriesasapointsourceofanthraxoutbreaks.

Contaminatedproductscomefromanimalsthatdiedofanthrax.

Wastewatereffluentfromplantscancontaminatedownstream

sedimentsandpastureswithanthraxspores,providinga

sourceoflocaloutbreaksinlivestockandfurtherproliferation

ofnovelintroducedvariantsofthepathogen.Marketingof

inadequatelysterilisedbonemealsandfertilisers,renderedfrom

contaminatedmaterials,canresultinlongdistanceredistribution

andintroducing“industrial”strainstolivestockremotefromthe

originalsource(Hugh-JonesandHussaini1975).

Undercertainenvironmentalconditions,concentrationsof

endosporeshavecausedperiodicoutbreaksamongwoodbison

intheSlaveRiverLowlands(SRL),MackenzieBisonSanctuary

(MBS),andWoodBuffaloNationalPark(WBNP)(Dragonand

Elkin2001;Gateset al.2001b;Pybus2000).Between1962and

1971,anthraxandtheassociateddepopulationandvaccination

programmesemployedtocontrolthedisease,accountedfor

over2,800woodbisondeaths(DragonandElkin2001).Further

outbreaksoccurredintheMBSin1993,intheSRLin1978,2000

and2006,andinWBNPin1978,1991,2000,and2001(Gates

et al.1995;Nishiet al.2002c).Fourfactorsthatareassociated

ratherconsistentlywiththeseepizooticsarehighambient

temperatures,intensematingactivity,highdensitiesofinsects,

andhighdensitiesofbisonastheycongregateandcompetefor

diminishingwaterandfoodsupplies(APHIS,USDA2006).Based

onthesefourfactors,twohypotheseshavebeenproposedto

explainoutbreaksofanthraxinbisoninnorthernCanada:(1)

“themodifiedhostresistancehypothesis”(GainerandSaunders

1989)and(2)“thewallowconcentratorhypothesis”(Dragonet

al.,1999).Thesetwohypothesesarenotmutuallyexclusive.

Arecentoutbreakwasreportedinacommercialherdin

south-westernMontanathatkilledover300bisonpasturing

onalargefoothillslandscapebeneaththeGallatinMountain

Range(Ronnow2008).Despitemassdeathsofbisonduring

anthraxoutbreaks,thesporadicnatureofoutbreaksand

predominanceofmaledeathssuggestthatthediseaseplaysa

minorroleinlong-termpopulationdynamicsunlessoperating

inconjunctionwithotherlimitingfactors(JolyandMessier

2001b;ShawandMeagher2000).Anthraxisnottreatablein

free-rangingwildlife,butcaptivebisoncanbevaccinatedor

treatedwithantibiotics(Gateset al.1995;Gateset al.2001b).

Carcassscavengingfacilitatesenvironmentalcontamination

withanthraxspores(Dragonet al.2005);thereforetimely

carcasstreatmentanddisposalduringanactiveoutbreakin

free-rangingbisonisconsideredanimportantpreventative

strategyforreducingthepotentialforfutureoutbreaks(Hugh-

JonesanddeVos2002;Nishiet al.2002a).Anthraxisapublic

healthconcernandhumansaresusceptible,however,exposure

fromnaturallyoccurringoutbreaksrequiresclosecontactwith

animalcarcassesorhides.Inaddition,humanshaverarelybeen

exposedtoanthraxthroughthepurchaseofcuriospurchasedby

tourists(Whitford1979).

5.1.3 Bluetongue

Bluetongue(BLU)isaninsect-borneviralhemorrhagicdisease

affectingmanyungulatesinthelowerlatitudesofNorthAmerica.

TheBLUvirusisamemberofthegenusOribivirusofthefamily

Reoviridae.Worldwidethereare24knownBLUserotypes,

butonlysixareactiveindomesticandwildruminantsfrom

NorthAmerica(Pearsonet al.1992).Bluetonguevirusesare

closelyrelatedtothevirusesintheepizootichemorrhagic

diseaseandBLUisknowntoinfectawidevarietyofwildand

domesticruminants(Howerthet al.2001).Bisonaresusceptible

toBLU,andthevirushasbeenisolatedunderfield,captive,

andexperimentalconditions(Dulacet al.1988).Thearthropod

vectorsofthebluetonguevirusarevariousspeciesofCulicoides

midges(GibbsandGreiner1989;Howerthet al.2001).Clinical

symptomsincludefever,stomatitis,oralulcerations,lameness,

andoccasionally,reproductivefailure(Howerthet al.2001).

Therearesubacute,acute,andevenchronicexpressionsofthe

diseaseinmanywildungulatesanddomesticlivestock.BLU

typicallyoccursinthelatesummerandearlyfalldepending

upontheseasonalpatternsofvectoractivity(Howerthet al.

2001).Factorsinfluencingthefrequencyandintensityofdisease

outbreaksareinnateherdimmunity,virulencefactorsassociated

withviruses,andvectorcompetencyandactivity.BLUoccursin

livestockovermuchoftheU.S.anditsdistributionparallelsthat

ofdomesticlivestock.ItsdistributionismorelimitedinCanada

whereitoncewasaregulateddiseaseuntilruleswererelaxed

inJuly2006(CFIAwebsite).Thereisconsiderabledifferencein

theepidemiologyofthediseasebetweennorthernandsouthern

portionsofNorthAmericadependingontheconsistencyof

vectoractivity.Inthesouthernareas,vectoractivityismore

commonandanimalpopulationsexhibitahigherprevalence

ofseroreactivityandantibodyprotection.BLUhasnotbeen

widelyreportedinbisonherdsinNorthAmerica.Serologic

surveysofseveralDepartmentofInteriorbisonherdsinthe

U.S.havenotfoundseroreactorsforbluetonguevirus(T.Roffe

personalcommunication;Tayloret al.1997).TheU.S.Fishand

WildlifeService(USFWS)hasopportunisticallyexaminedbison


neararecentoutbreakofBLUindeerandfoundnoevidence

ofexposure(T.Roffepersonalcommunication).Aswithmany

vector-borndiseases,climatechangeisapotentialfactor

affectingthedistributionofvectorsandthereforetheoccurrence

ofBLU(Gibb1992).Thereisnoeffectivetreatmentand,under

naturalconditions,thediseaseisnotconsideredasignificant

threattohumanhealth.Therehasbeenonehumaninfection

documentedinalaboratoryworker(WHOwebsite).

5.1.4 Bovine spongiform encephalopathy

Bovinespongiformencephalopathy(BSE),or“madcow

disease”asitiscommonlyknown,isoneofasuiteofdistinct

transmissiblespongiformencephalopathies(TSE)identified

duringthepast50years.TSEsareapparentlycausedbyrogue,

misfoldedproteinagentscalledprions(PrPSC)thataredevoid

ofnucleicacids(Prusiner1982).NootherTSEinmanoranimal

hasreceivedmoreworldwideattentionthanBSE(Hadlow1999).

Itwasfirstidentifiedin1986inEnglandandhassincehadfar

reachingeconomic,political,andpublichealthimplications.BSE

isaneurologicdiseasecharacterisedbyspongiformchange

ingraymatterneurophil,neuronaldegeneration,astrocytosis,

andaccumulationofmisfoldedPrPSC(Williamset al.2001).

Clinicallythediseaseisprogressive,displayinggradual

neurologicimpairmentovermonthsoryearsandisusuallyfatal.

Thediseasecausesprogressiveweightloss,low-leveltremors,

behaviouralchanges,ataxia,andposturalabnormalities.

Substantialevidenceexistsforgeneticvariationinsusceptibility

amongandwithinspecies(Williamset al.2001).Casesof

BSEwereidentifiedin10speciesofBovidaeandFelidae

atazoologicalcollectionintheBritishIsles(Kirkwoodand

Cunningham1994).Atleastoneofthesecasesincludedbison.

Worldwide,otherspeciessusceptibletoBSEincludecheetah,

macaquesandlemurs(Williamset al.2001).TherecentBSE

epidemicinEuropewaslinkedtooralingestionofcontaminated

feed(containingruminantderivedprotein),however,thereis

someevidenceforlow-levellateraltransmission.Thereareno

knowntreatmentsorpreventionsforBSE.Thehumanform

callednewvariantCreutzfeldt-Jakobdiseasehasbeenlinked

toconsumptionofBSEcontaminatedfoods.Duetotherisk

ofhumanexposuretoBSE,thisdiseaseishighlyregulated

worldwide.RecentcasesofBSEhavebeenreportedinCanada

andtheU.S.butareextremelyrareinthelivestockindustry.

Canadareportedacasein1993thatwasimportedfromEngland

andthefirstdomesticcasewasdetectedin2003.TheU.S.

reporteditsfirstcaseofBSEin2003.Sincethen,proteinby-

productswerebannedinlivestockfeed,nationalsurveillance

wasimplementedinbothcountries,andseveralregulations

werepromulgatedtorestrictimportsandexportsacrossthe

U.S.-Canadaboundary.Althoughbisonareconsideredtobe

susceptible,therehasnotbeenacaseofBSEreportedin

Americanbison.

5.1.5 Bovine brucellosis

Bovinebrucellosis,alsoknownasBang’sdisease,iscaused

byinfectionwiththebacteriumBrucella abortus(Tessaro1989;

Tessaro1992).Theprimaryhostsforbovinebrucellosisare

cattle,bison,andotherbovidspecies(Tessaro1992),however,

otherwildungulatessuchaselk(Cervus elaphus)arealso

susceptibleandseemtoplayaroleininterspeciestransmission

intheGreaterYellowstoneArea(GYA)(Davis1990;Rhyanet al.

1997;Thorneet al.1978).Evidencesuggeststhatbrucellosis

wasintroducedtoNorthAmericafromEuropeduringthe

1500s(MeagherandMayer1994;AguirreandStarkey1994).

Thediseaseisprimarilytransmittedthroughoralcontact

withabortedfoetuses,contaminatedplacentas,anduterine

discharges(Reynoldset al.1982;Tessaro1989).Theimpacts

ofbrucellosisonfemalebisonincludeabortion,inflammationof

theuterus,andretainedplacenta(Tessaro1989).Greaterthan

90%ofinfectedfemalebisonabortduringthefirstpregnancy;

however,naturallyacquiredimmunityreducesthisabortionrate

to20%afterthesecondpregnancy,andtonearlyzeroafterthe

thirdpregnancy(Daviset al.1990;Daviset al.1991).Malebison

experienceinflammationoftheseminalvessels,testicles,and

epididymis,and,inadvancedcases,sterility(Tessaro1992).

Bothsexesaresusceptibletobursitisandarthritiscausedby

concentrationsofthebacterialorganisminthejoints,resulting

inlameness,andpossiblyincreasedvulnerabilitytopredation

(Tessaro1989;Tessaro1992).

SerologyisusedtodetectexposuretoB. abortusbyidentifying

thepresenceofantibodiesintheblood.Sero-prevalenceisthe

percentageofanimalsinaherdthatcarryantibodies(Cheville

et al.1998).Asero-positiveresult,indicatingthepresence

ofantibodies,doesnotimplycurrentinfection,andmay

overestimatethetruelevelofbrucellosisinfection(Chevilleet

al.1998;DobsonandMeagher1996)becausetheorganism

mustbeculturedfromtissuesamplestodiagnoseananimal

asinfected.However,adisparitybetweenserologyresults

andlevelofinfectioncouldalsobeattributedtofalsenegative

cultureresultsrelatedtothedifficultiesinisolatingbacteriafrom

chronicallyinfectedanimals(Chevilleet al.1998).

Thereiscurrentlynohighlyeffectivevaccineforpreventing

bovinebrucellosis(Chevilleet al.1998;Davis1993).Strain19

(S19)wasacommonlyusedvaccineadministeredtocattle

fromthe1930suntil1996(Chevilleet al.1998).Itwasonly67%

effectiveinpreventinginfectionandabortionincattle(Cheville

et al.1998).S19wasfoundtoinduceahighfrequencyof

abortionsinpregnantbison(Daviset al.1991).Otherstudies

failedtodemonstrateefficacyofS19asabisoncalfhood

vaccine(Templetonet al.1998).Anewervaccine,strainRB51,is

nowpreferredoverS19becauseitdoesnotinduceantibodies

thatcaninterferewithbrucellosisserologytestsfordisease

exposure(Chevilleet al.1998;Roffeet al.1999a).RB51protects


cattleatsimilarlevelstoS19(Chevilleet al.1993).Dosesof

RB51consideredtobesafeincattlewerefoundtoinduce

endometritis,placentitis,andabortioninadultbison(Palmer

et al.1996).However,Roffeet al.(1999a)foundRB51hadno

significantadverseeffectsonbisoncalves.Thesafetyand

efficacyofRB51inbisonremainsunclearbut,nonetheless,

itwasprovisionallyapprovedforuseinbisonintheU.S.The

vaccineisnotrecognisedinCanadaandvaccinatedcattleare

notallowedintothecountry(CFIA2007).Everybisonimported

intoCanadafromtheU.S.mustbequarantinedfromthetimeof

itsimportationintoCanadauntilitprovesnegativetotestsfor

brucellosisperformednotlessthan60daysafteritwasimported

intoCanada(CFIA2007).

Quarantineprotocolshavebeendevelopedforbisonto

progressivelyeliminateallanimalsexposedtobrucellosisfrom

apopulation(APHIS,USDA2003;Nishiet al.2002b).These

protocolshavebeensuccessfulforeliminatingbrucellosisin

woodbisonthroughtheHookLakeprojectandarecurrently

beingattemptedintheGYA(AuneandLinfield2005;Nishiet al.

2002b).Resultsfromthesetwostudies,andothercasestudies

(HMSP,WCNPandEINP),haveshownthatbrucellosiscanbe

effectivelyeliminatedfromexposedpopulationswithahigh

degreeofcertaintyusingtestandslaughterprotocols.

5.1.6 Bovine tuberculosis

Bovinetuberculosis(BTB)isachronicinfectiousdiseasecaused

bythebacteriumMycobacterium bovis(Tessaroet al.1990).

TheprimaryhostsforBTBarecattleandotherbovidspecies,

suchasbison,waterbuffalo(Bubalus bubalis),Africanbuffalo

(Syncerus caffer),andyak(Bosgrunniens).Primaryhostsare

thosespeciesthataresusceptibletoinfectionandwillmaintain

andpropagateadiseaseindefinitelyundernaturalconditions

(Tessaro1992).Otheranimalsmaycontractadisease,but

notperpetuateitundernaturalconditions;thesespeciesare

secondaryhosts.Thebisonistheonlynativespeciesofwildlife

inNorthAmericathatcanactasatrueprimaryhostforM. bovis

(Tessaro1992).HistoricalevidenceindicatesthatBTBdidnot

occurinbisonpriortocontactwithinfecteddomesticcattle

(Tessaro1992).Currently,thediseaseisonlyendemicinbison

populationsinandnearWBNP,whereitwasintroducedwith

translocatedplainsbisonduringthe1920s.BTBisprimarily

transmittedbyinhalationandingestion(Tessaroet al.1990);

thebacteriummayalsopassfrommothertooffspringviathe

placentalconnection,orthroughcontaminatedmilk(FEARO

1990;Tessaro1992).Thediseasecanaffecttherespiratory,

digestive,urinary,nervous,skeletal,andreproductivesystems

(FEARO1990;Tessaroet al.1990).Onceinthebloodorlymph

systemsthebacteriummayspreadtoanypartofthehostand

establishchronicgranulomatouslesions,whichmaybecome

caseous,calcified,ornecrotic(Radostitset al.1994;Tessaro

1992).Thischronicdiseaseisprogressivelydebilitatingtothe

host,andmaycausereducedfertilityandweakness;advanced

casesarefatal(FEARO1990).Thediseasemanifestssimilarly

incattleandbison(Tessaro1989;Tessaroet al.1990).Boththe

U.S.andCanadaperformnationwidesurveillanceofabattoir

facilitiestomonitorBTBinfectionincattleanddomesticbison.

ThereisnosuitablevaccineavailableforBTB(FEARO1990;

CFIA2000;APHISUSDA2007).Everybisonimportedinto

CanadafromtheU.S.mustbequarantinedfromthetimeof

itsimportationintoCanadauntilitprovesnegativetotests

forBTBperformedatleast60daysafteritwasimportedinto

Canada(CFIA2007).Aquarantineprotocolhasbeendeveloped

andanexperimentalprojectwasattemptedtosalvagebison

fromaBTBexposedpopulation(Nishiet al.2002b).Although

atfirstitappearedtobeasuccessfultoolforsalvagingbison

fromanexposedherd,after10years,severalofthesalvaged

animalsexpressedBTB,andin2006allsalvagedanimalswere

slaughtered(Nishipersonalcommunication).Thereissome

evidencethatBTBcanbetreatedinindividualanimalsusing

longtermdosingwithantibiotics,butthedurationoftreatment,

costsoftherapy,andtheneedforcontainmentmakethis

optionimpracticalforwildlife.Theonlydefinitivemethodfor

completelyremovingBTBfromaherdisdepopulation(CFIA

2000;APHISUSDA2005).Theonlyalternativetodepopulation

iscontrollingthespatialdistributionandprevalenceofdisease

throughacooperativeriskmanagementapproachinvolvingall

stakeholders.Thebasicprerequisitesforeffectivelyaddressing

riskmanagementassociatedwithBBTBinbisonareteamwork,

collaborationacrossprofessionaldisciplines,andrespectfor

scientificandtraditionalecologicalknowledgeamongtechnical

andnontechnicalstakeholders(Nishiet al.2006).BTBcan

infecthumans,butitistreatablewithantimicrobialdrugs.

HumanTBduetoM. bovishasbecomeveryrareincountries

withpasteurisedmilkandBTBeradicationprogrammes.

5.1.7 Bovine viral diarrhoea

Bovineviraldiarrhoea(BVD)isapestivirusthatinfectsawide

varietyofungulates(Loken1995;Nettleston1990).Serologic

surveysinfree-rangingandcaptivepopulationsdemonstrate

priorexposureinmorethan40mammalspeciesinNorth

America(Nettleston1990;Tayloret al.1997).Thesuspected

sourceofBVDinwildanimalsisdirectcontactwithdomestic

livestock.Infectionsinwildruminants,likecattle,aredependent

uponthevirulenceoftheisolate,immunestatusoftheanimal

host,andtherouteoftransmission.Infectionsincattleare

usuallysubclinical,butsomeinfectionsmaycausedeath

orabortionsinpregnantanimals.Factorsinfluencingthe

persistenceofBVDincludepopulationsizeanddensity,herd

behaviour,timingofreproduction,andsurvivorshipofyoung

(Campenet al.2001).

Positiveserologicevidencewasreportedforbloodsamples

frombisonintheGYA(Tayloret al.1997;Williamset al.1993),


Alaska(Zarnke1993)andfrombisonatElkIslandNational

Park(EINP)inAlberta(Cool1999;Gateset al.2001b).InYNP,

positiveantibodytitresweredetectedin31%oftestedanimals

(Tayloret al.1997).Thereareunpublisheddataregardingsero-

reactivityfrombisontransportedtoMontanafromWCNPin

SouthDakota(K.Kunkel,personalcommunication).TheJackson

bisonherd,withaknownhistoryofcomminglingwithcattle,

hasdemonstratedlow-leveltitres,butnoevidenceofBVD

antigenorclinicaldiseasehasbeenfound(T.Roffe,personal

communication).ClinicalBVDwasreportedintheEINPplains

bisonherdin1996,promptingaserologicalsurveyofplains

bisonandwoodbisonherds(Cool1999;Gateset al.2001b).

Forty-sevenpercentof561plainsbisonfromEINPtestedsero-

positiveforBVD;onetestedpositiveforthevirusantigen.At

leastsixplainsbisondeathsinEINPwereattributedtotheBVD

virus(Cool1999).TissuesfromthesuspectedcasesofBVD

infectedplainsbisonweresubmittedtotheAnimalDisease

ResearchInstitute,Lethbridge,Alberta,Canada,andtype1

BVDviruswasisolated(TessaroandDeregt1999).Noneof352

woodbisonintheParktestedsero-positiveforBVDatthetime.

BothplainsandwoodbisonpopulationsatEINParevaccinated

forBVDduringannualroundups.However,calvesusedin

translocationsarenotvaccinatedtoallowfuturescreeningof

recipientpopulationsforBVD.InPoland,Sosnowski(1977)

reportedBVDinacaptiveEuropeanbison.BVDiscommonin

cattleinNorthAmericaandposesnoknownrisktohumans.

5.1.8 Johne’s disease

Johne’sdisease(JD)iscausedbytheetiologicagent

Mycobacterium avium subsp. paratuberculosis,ahardy

bacteriumrelatedtotheagentsofleprosyandtuberculosis.

Itoccursworldwideaffectingavarietyofdomesticandwild

ruminantsincludingbison,cattle,andsheep(Buergeltet

al.2000;Williams2001).Infectionsoftenleadtochronic

granulomatousenteritiswithclinicalsignsofdiarrhoea,weight

loss,decreasedmilkproduction,andmortality.JDiscommon

incattle.Recentstudieshaveshownthatmorethan20%of

dairyherdsintheU.S.haveJD(Chiet al.2002;Ottet al.1999)

causinganestimatedeconomiclossofmorethanUS$200

millionannually.JDtypicallyentersaherdwheninfected,

asymptomaticanimalsareintroduced.Unpasteurisedraw

milkorcolostrummaybeasourceofinfectionforartificially

raisedcalves.Animalsaremostsusceptibletoinfectionduring

theirfirstyearoflife.Neonatesmostoftenbecomeinfected

byswallowingsmallamountsofcontaminatedmanurefrom

thegroundorfromtheirmother’sudder.Animalsexposedtoa

verysmalldoseofbacteriaatayoungage,andolderanimals,

arenotlikelytodevelopclinicaldiseaseuntiltheyaremuch

older.Afterseveralyears,infectedanimalsmaybecomepatent

andshedmycobacteriaintheirfaeces.Typically,pre-patent

animalsdonotshowsymptomsofdisease;consequently,most

infectionsgounnoticedandundiagnosed.Thereisnotreatment

foranimalsinfectedwithJDandpreventionisthebestcontrol

measure.Humansarenotconsideredsusceptible,butM. a.

paratuberculosis hasbeenisolatedinpatientswithchronic

enteritis(Crohn’sdisease)(Chiodini1989).JDisnotconsidered

tobeadiseaseproblemwhenbisonareonopenrangelands

andmanagedatlowdensity.However,restrictionsmayapply

tointer-jurisdictionalmovementofanimalsfromknowninfected

herds.Hence,maintaininglowriskstatusforbisonherds

usedasasourceforconservationprojectsisanimportant

consideration.

In1998,theU.S.AnimalHealthAssociationapprovedthe

VoluntaryJohne’sDiseaseHerdStatusProgramforcattle

(VJDHSP).TheVJDHSPprovidestestingguidelinesforStates

tousetoidentifylivestockherdsaslowriskforJDinfection.

Withnumeroustestsoverseveralyears,herdsprogressto

higherstatuslevels.Thehigherthestatuslevel,themore

likelyitisthataherdisnotinfectedwithJD.InApril2002,

USDA-APHIS-VeterinaryServiceincorporatedportionsof

thisprogrammeintonationalprogrammestandards:Uniform

ProgramStandardsfortheVoluntaryBovineJohne’sDisease

ControlProgram(VBJDCP).VBJDCP-test-negativeherdsserve

asasourceoflowJDriskstock.TestingforJDinconservation

herdshasbeensporadicandopportunistic.Diagnostictools

arebeingdevelopedandimproved.TherearenoreportsofJD

inconservationbisonherdsintheliterature,however,some

commercialoperationshavediscoveredJD,andinmanycases

aremanagingtopreventitsspreadandreduceitsimpacton

theindustry.

5.1.9 Malignant catarrhal fever (sheep associated)

Malignantcatarrhalfever(MCF)isaserious,oftenfataldisease

affectingmanyspeciesoftheOrderArtiodactyla.Itiscaused

byvirusesofthegenusRhadinovirus.Atleast10MCFviruses

havebeenrecognisedworldwideandfiveviruseshavebeen

linkedtodisease.Thevirusesmostsignificanttolivestockare

thosecarriedbysheep,goatsorwildebeest(Connochaetes

spp.).Althoughovineherpesvirustype2(sheepassociated

MCF)doesnotcausediseaseinitsnaturalhost,domestic

sheep,itdoescauseMCFinbison.Serologicaltestingindicated

thatitiscommonindomesticgoats(61%)andsheep(53%)

intheU.S.(Liet al.1996).MCFisanimportantdiseaseinthe

commercialbisonindustryasitisoneofthemostinfectious

diseasesofbison,especiallyathighdensities(Heuscheleand

Reid2001).Itcauseshighlylethalinfectionsinbison,with

thereportedincidenceofmortalityinaherdofupto100%

(Schultheisset al.2001).Infectionsproceedrapidlytoclinical

disease.MCFisexpressedintwoforms,acuteandchronic,

butregardless,deathensuesinmostcases.Intheacuteform,

bisonusuallydiewithin7–10daysofinfectionorwithin48hr

ofbecomingsymptomatic.Alternatively,deathmayensueas


longas156dayspost-infection.Someanimalsrecoverand

remainpersistentlyinfected(Schultheisset al.1998).Clinical

signsinbisonincludehemorrhagiccystitis,colitis,conjunctivitis,

oculardischarge,nasaldischarge,excesssalivation,anorexia,

diarrhoea,melaena,haematuria,multifocalulcerationofthe

oralmucosa,fever,circling,ataxia,behaviourssuggestiveof

blindness,lameness,anddifficulturination(Liggittet al.1980;

Ruthet al.1977;Schultheisset al.1998).Lymphadenomegaly

andcornealopacityoccurinfewerthanhalfthecases

(Schultheisset al.2001).Directcontactbetweenbisonand

domesticsheepisconsideredthemostlikelysourceofinfection.

Hence,bisonshouldnotbegrazedinthesamepasturesor

adjacenttopastureswithsheep.Althoughmostinfectionsoccur

whenbisonareincloseassociationwithdomesticsheep,MCF

wasreportedinbisonherdsthatwerefivekilometres(three

miles)fromalambfeedlot(Schultheisset al.2001).Dr.T.Roffe

hasconductedserologicsurveysoftwoU.S.Departmentof

theInteriorbisonherdsnotassociatedwithdomesticsheep

andhasfoundnosero-reactorsforMCF(T.Roffe,personal

communication).Thereisnovaccineoreffectivetreatmentfor

MCFandthebestwaytocontrolthisdiseaseistominimise

contactwithreservoirhosts.Thereisnoevidencethatisolatesof

MCFareinfectioustohumans(HeuscheleandSeal1992).

5.2 Episodes of reportable Diseases in plains Bison

Basedonthissurvey,twoplainsbisonconservationherdsin

NorthAmericahavesignificantchronicdiseaseissues:YNP

herdandtheJacksonherdinGTNP/NER.Theseherds,which

accountfor4,700bison(asofwinter2008),or24%oftheentire

NorthAmericanplainsbisonconservationpopulation,harbour

brucellosis.

5.2.1 Yellowstone National park

BrucellosiswasfirstdetectedintheYNPbisonpopulation

in1917(Mohler1917).Theoriginofbrucellosisintheparkis

unclear,butwasprobablytheresultoftransmissionfromcattle

(MeagherandMayer1994).Opportunisticandsystematic

serologicalsurveysinthearearevealedsero-prevalencevarying

between20%and70%,whilebacterialculturesindicated

aninfectionprevalenceofapproximately10%(Dobsonand

Meagher1996;MeagherandMayer1994).Althoughthetrue

prevalenceofthediseaseisunknown,theYNPbisonpopulation

isconsideredtobechronicallyinfectedwithbrucellosis(Cheville

et al.1998).Morerecentresearchontheepidemiologyof

brucellosisinYellowstonebisonfoundthat46%ofthesero-

reactoranimalswereculturepositive(Roffeet al.1999b).Recent

demographicanalysisindicatesthatbrucellosishasasignificant

reproductiveeffect,thatthegrowthrateofthepopulationcould

increaseby29%intheabsenceofbrucellosis(Fulleret al.2007),

andthatbrucellosisisnotathreattothelong-termviabilityof

theYNPbison(MayerandMeagher1995;USDOIandUSDA

2000).Fulleret al.(2007)conductedadetailedanalysisofthe

demographicsoftheYellowstonepopulationfrom1900-2000

andfoundevidenceofdensitydependentchangesinpopulation

growthasnumbersapproached3,000animals.Thispopulation

appearsrobustandhasgrownattimestoexceed4,000,

althoughitwasreducedtofewerthan3,000severaltimesduring

thepastdecadeunderthecurrentherdmanagementregime(R.

Wallen,personalcommunication).

Herdmanagementisaffectedbythepresenceofbrucellosis

primarilybecauseofthepotentialriskthediseaseposestothe

livestockindustry(Keiter1997).Bisonleavingtheparkcould

potentiallytransmitthediseasetodomesticcattlegrazing

onadjacentNationalForestandprivatelandsinMontana,

WyomingorIdaho(USDOIandUSDA2000).Bisonleavethe

parkinthewinteronthenorthandwestboundarieswithin

Montana;movementtotheeastandsouthisrarebecauseof

topographicalbarriers(R.Wallen,personalcommunication).

Transmissionofbrucellosisfrombisontocattlehasbeen

demonstratedincaptivestudies;however,thereareno

confirmedcasesoftransmissioninthewild(Bienen2002;

Chevilleet al.1998;ShawandMeagher2000).Nevertheless,

thepotentialexists,andthishascreatedacontentiousbison

managementissueinthearea.

RelyingontheAnimalIndustryActof1884,theU.SDepartment

ofAgriculturebeganpreventingandcontrollingthespreadof

contagiouslivestockdiseasesintheU.S.In1947,federaland

stateofficialsbeganworkingcloselywiththelivestockindustry

toeradicatebrucellosis(Keiter1997;NPSUSDOI2000).Each

staterepresentedintheGYAisaco-operatorintheNational

BrucellosisProgramandhasauthoritytoimplementcontrol

programmesforbrucellosisinfectedorexposedanimals

withintheirrespectiveboundaries.Duetothetransmissionof

brucellosistocattle,presumablybyelk,Montana,Wyoming,and

Idahohaveeachperiodicallylosttheirbrucellosis-freestatus

ascertifiedbyAPHIS.Transmissionofbrucellosistocattlein

Montana,WyomingorIdahoindirectlyaffectsallproducersin

thesestates.IftheirAPHISstatusisdowngraded,otherstates

mayrefusetoacceptcattlefromproducersintheGYA(Cheville

et al.1998).

Resolutionofthisissuerequirestheinvolvementof,and

cooperationamong,agenciesinseveraljurisdictions:The

NationalParkService(NPS),theU.S.ForestService(USFS),

APHIS,andtheStateofMontanaDepartmentofLivestock

(MDOL)andMontanaDepartmentofFish,Wildlife,andParks

(MFWP).Aftermanyyearsofmediaandlegalcontroversyover

bisonmanagement,theagenciesacknowledgedtheneedto

cooperativelydevelopalong-termbisonmanagementplan

(PlumbandAune2002).In1990,theycommencedtheprocess


foraninteragencyenvironmentalimpactstatementtodevelop

alternativesfortheplan(USDOIandUSDA2000).Aseries

ofinteragencyinterimplansfollowed,whichprogressively

incorporatedgreatertoleranceforbisonoutsidetheparkin

certainareas,andenabledNPSandMFWPpersonneltolethally

removebisonmovingfromYNPintoMontana.

Legalandpolicydisagreementsbetweenthefederalagencies

andtheStateofMontanainhibitedthedevelopmentofalong-

terminteragencymanagementplanuntil2000whencourt-

orderedmediationresultedinafinaldecisionforalong-term

managementapproach.Thelong-termplanemploysanadaptive

managementapproachwiththreephasedstepsforeachofthe

northandwestboundaryareas(USDOIandUSDA2000).The

planincorporatesseveralriskmanagementstrategiesincluding

spatialandtemporalseparationofbisonandcattle,capture,

test,andslaughterofsero-positivebison,hazingofbisonback

intothepark,vaccination,andradio-telemetrymonitoringof

pregnantbisontolocatepossiblesourcesofinfectionifacow

givesbirthorabortsoutsidethepark(USDOIandUSDA2000).

Theultimatepurposeoftheplanistomaintainawild,free-

rangingpopulationofbisonwhile,atthesametime,protecting

theeconomicviabilityofthelivestockindustryinMontana

byaddressingtheriskofbrucellosistransmission;itisnota

brucellosiseradicationplan(PlumbandAune2002).Although

eradicationofbrucellosisfrombisonintheparkisapossible

futuregoal,suchaneffortiscomplicatedbyretransmission

potentialfromelkintheGYA,whichalsoharbourthedisease

(Chevilleet al.1998).Developmentofmoreeffectivevaccines

andvaccinationmethodsforbisonandelkarerequiredbefore

consideringeradicationalternatives(Chevilleet al.1998).Recent

researchongenesthatcontrolnaturalresistancetobrucellosis

mayalsoprovidefuturemethodsforeradicatingbrucellosis

(Templetonet al.1998).

Recenttransmissionofbrucellosisfromelktocattleand

thesubsequentlossofMontana’sbrucellosisstatushave

complicatedmanagement.Currentinitiativesareaimedat

managingtheproblemofbrucellosisinelkandbison.Changes

inthedistributionofbison,elk,andcattlewillgeneratefurther

publicdebateandperhapslegalaction.TheGYAsituation

illustratesthetremendousdifficultyinmanagingwildfreeranging

ungulatesaffectedbyasignificantdiseaseonalargelandscape

wherehumanlivelihoodsareatrisk.

5.2.2 Grand teton National park/National Elk refuge (Jackson herd)

TheJacksonherdofapproximately1,100animalsresidesinthe

southernendoftheGYA(USFWSandNPS2007),migrating

betweenGrandTetonNationalPark(GTNP)inthesummerand

theadjacentNationalElkRefuge(NER)inthewinter(Chevilleet

al.1998).AswiththeYNPherd,theJacksonherdischronically

infectedwithbrucellosis.Williamset al.(1993)reportedsero-

prevalenceof77%andinfectionprevalenceof36%fortheherd.

Serologytestsoverthepastfiveyearsindicateasero-prevalence

of80%(S.Cain,personalcommunication).Areductionof8%in

fecundityhasbeenestimated,however,thepopulationhasbeen

increasingsincethe1970sdespitethedisease(S.Cain,personal

communication,Chapter6;USFWS-NPS2007).

TheJacksonherdwasfoundedin1948withthereintroduction

of20bisonfromYNPtoa1,500-acredisplaypen.Thesebison

wereconfineduntil1963whenbrucellosiswasdiscoveredin

theherd(Chevilleet al.1998).Allbutfourvaccinatedyearlings

andfivevaccinatedcalvesweredestroyed.In1964,Theodore

RooseveltNationalPark(TRNP)provided12brucellosis-free

bisontoaugmenttheJacksonherd(Chevilleet al.1998).In

1968,theherdescapedfromtheprogressivelydeteriorating

enclosurefacility(Chevilleet al.1998;Williamset al.1993).From

thatpointtheparkallowedtheherdtoroamfreely.Thebison

herddiscoveredthefeedgroundattheNERin1980.Although

theherdwasapparentlyhealthywhenreleased,itissuspected

thatinfectedelkontheNERintroducedbrucellosistothe

Jacksonbison(Chevilleet al.1998).

SimilartotheYNPherd,thefree-rangingnatureoftheJackson

herdallowsforthepossibilityoftransmittingbrucellosis

todomesticlivestockinthearea,althoughsincetheNER

excludescattle,thereislimitedcontactbetweenJackson

bisonandcattleduringthewinterfeedingperiod(Chevilleet

al.1998).Thereispotentialforcontact,however,whenbison

moveamongprivate,USFS,GTNPandNERjurisdictions,

especiallyinsummer,whencattlearemaintainedongrazing

allotmentsinGTNP,privateranchlands,andadjacentUSFS

lands(Chevilleet al.1998;Keiter1997).

AnewbisonandelkmanagementplanfortheNERandGTNP

wasapprovedinApril2007.Anearlierbisonmanagementplan

approvedin1996,afterundergoingaNationalEnvironmental

PolicyAct(NEPA)process,wassubjecttolitigationbyananimal

rightsgroupthatquestionedtheinclusionofasporthuntto

managepopulationlevelsandtheexclusionofananalysisof

elkmanagementonthefederallandsinthedecisionprocess

(Cain,personalcommunication;USFWS-NPS2001).Thecourt

ruledthatdestructionofbisonforpopulationcontrolcouldnot

beconducteduntiltheinvolvedagenciesanalysedtheeffects

ofwinterfeedingonbisonandelkthroughanadditionalNEPA

process(USFWS-NPS2001).Thefeedinggroundsattract90%

oftheJacksonbisonand6,000-8,000elktoonesmallarea,

creatingzonesofhighanimaldensity,wheretransmission

maybeenhancedamongandbetweenelkandbison(Bienen

2002;USFWS-NPS2007).GTNPandtheNERdetermined

thatacombinedelkandbisonmanagementplanisneededto

addresstheinterconnectedissuesofthetwospecies,including

winterfeedinganddiseasemanagement.TheJacksonbison


andelkherdsmigrateacrossseveraljurisdictionsincluding

theNER,GTNP,YNP,Bridger-TetonNationalForest,Bureau

ofLandManagement,StateofWyoming,andprivatelands.

TheNPSandFWScoordinatedtheextensiveinvolvementof

theassociatedagencies,organisations,andprivateinterests

affectedbythisnewmanagementplanandEnvironmental

ImpactStatement(EIS).TheU.S.DepartmentofInterior

(USDOI)publishedarecordofdecisioninApril2007,selectinga

managementalternativethatemphasisesadaptivemanagement

ofelkandbisonpopulationswhilereducingtheirdependence

uponfeedgrounds.Theplanalsocallsforabrucellosis

vaccinationprogrammeforelkandbisonconductedbythe

StateofWyoming.Recenthuntingprogrammes,modification

offeedingprogrammesanddiseasemanagementhave

reducedthenumberofbisonto700animalsandthelong-term

managementofthisherdisnowprescribedinalong-termplan.

Severallegalchallengesweremountedandtheimplementation

oftheplanremainscontroversial.

5.3 An Occurrence of reportable Diseases in Wood Bison

WoodbisonherdsinandaroundWBNP,includingSRL,are

infectedwithBTBandbrucellosis(Gateset al.1992;Gateset

al.2001c).Thesediseasedherdsaccountforabout50%ofthe

totalwoodbisonconservationpopulation.JolyandMessier

(2001a)reportedthesero-prevalenceofthediseasestobe31%

forbrucellosisand49%fortuberculosis.Withtheexceptionof

free-rangingbisonintheWBNPandGYA,aggressiveeradication

programmesinboththeU.S.andCanadahavereducedthe

probabilityofbrucellosisandBTBindomesticcattleandbison

herdstoextremelylowlevels.Thewilddiseasedwoodbison

herdsinandnearWBNParetheonlyknownreservoirsof

BTBamongallbisonconservationherds(Gateset al.2001c;

Reynoldset al.2003;ShawandMeagher2000).

BTBandbrucellosiswerelikelyintroducedtowoodbison

populationswiththetransferofplainsbisonfromWainwright

BuffaloParkinthe1920s(Fuller2002).In1925,theCanadian

governmentimplementedaplantomove6,673plainsbison

fromtheovercrowdedWainwrightBuffaloParktoWBNP.The

transferproceededdespiteoppositionfrommammalogical

andbiologicalsocietiesintheU.S.andCanada,whowarned

oftransmissionofBTBtotheresidentwoodbisonpopulation

(Anonymous1925;Ogilvie1979).BTBwasfirstreportedin

WBNPin1937(Fuller2002;Gateset al.1992;Geist1996).

AlthoughitisnotknownwhetherBTBwasendemicamong

woodbisonpriortothetransfer(Reynoldset al.1982),

evidenceindicatesthatthediseasewasintroducedtowood

bisonwiththetransferofplainsbison(Fuller1962).Brucellosis

wasalsopresentintheplainsbisonherdandwasreportedin

WBNPin1956(Gateset al.1992).

ThepresenceofBTBandbrucellosisthreatenstherecovery

ofwoodbisoninseveralways.First,theinfectedanimalsare

subjecttoincreasedmortality,reducedfecundity,andincreased

vulnerabilitytopredation(Gateset al.1992;JolyandMessier

2001a).In1934,thebisonpopulationinWBNPwasestimated

at12,000animals(Soper1941).Thepopulationdecreasedfrom

approximately11,000in1970to2,151in1999(Joly2001).

Thisdecreasehasbeenattributedtotheinteractiveeffectsof

diseasesandpredation(Carbynet al.1998;Fuller1991;Jolyand

Messier2001a).Recently,theWBNPpopulationincreasedto

4,050,althoughthereasonsforthisincreaseareunclear(Bradley

2002,personalcommunication).

Second,thepotentialexistsfortheinfectedherdstotransmit

thediseasestohealthyherds,mostnotablytheMackenzie,

Nahanni,andHay-Zamaherds(AnimalPlantandFood

RiskAssessmentNetwork(APFRAN1999).Since1987,the

GovernmentoftheNorthwestTerritorieshasmanageda

39,000km2BisonControlAreasouthoftheMackenzieRiver

topreventmovementofdiseasedbisonintotheMBS(Nishi

2002).Recentanalysisandmodellingofbisonmovementson

thelandscapehavedemonstratedconsiderableriskpotential

fortransmissionofdiseasestohealthywoodbisonherdsand

bisonranchesinthevicinityofthediseasedherds(Gateset al.

2001a;Mitchell2002).TheGovernmentofAlbertaannounced

anewhuntingseasonfortheHayZamaherdin2008.The

purposeofthehuntistomaintainthewoodbisonpopulationat

approximately400andlimitdistributionoftheseanimalsuntil

thediseasedbisonissue,inandaroundWBNP,issuccessfully

resolved.Inparticularthehuntwillbeusedtocontrolexpansion

oftheHay-Zamaherdeastward,preventingcontactwithbison

emigratingfromWBNPthatmaybeinfectedwithbrucellosisor

BTB.Althoughpreliminary,resultsofserologicaltestsandpost

mortemexaminationofabout100bisonharvestedfromthe

Hay-Zamapopulationinthewinterof2008werenegativeforthe

twobovinediseases(D.Moyles,AlbertaSustainableResource

Development,personalcommunication).

Muchresearchanddebatehasbeenfocusedontryingto

resolvethediseasedbisonissueinnorthernCanada.In1990,

theFederalEnvironmentalAssessmentPanelreleasedits

reportonitsanalysisofthediseaseissues(FEARO1990).

Thepanelconcludedthateradicationofthediseasedwood

bisonpopulationsistheonlymethodforeliminatingtheriskof

transmissionofbrucellosisandBTBfrombisontodomestic

cattle,non-diseasedwoodbison,andhumans.Thepanelfurther

recommendedthathealthywoodbisonbereintroducedtothe

areafollowingdepopulationofthediseasedherds.Sourcesof

healthybisonforreintroductioncouldincludetheEINPwood

bisonherdandothercaptiveherdssupplementedbydisease-

freeanimalssalvagedfromtheNorthernBisonherds(FEARO

1990).Onesuchsalvageoperation,theHookLakeWoodBison

RecoveryProjectinFortResolution,NorthwestTerritories,was


attempted(Nishiet al.2002b),butfailed.In2006,after10years

ofisolationandrigorousdiseasetesting,BTB-infectedbison

weredetectedintheherd.

SeveralconstituenciesrejectedtheFEARO(1990)panel’s

recommendationtodepopulateWBNPherds.TheNorthern

BuffaloManagementBoard(NBMB)wasformedtodevelop

afeasibleeradicationplan(Chisholmet al.1998;Gateset al.

1992).TheNBMBrecommendedfurtherresearchintobison

anddiseaseecologybeforeplanningmanagementactions

fortheregion(RAC2001).In1995,theMinisterofCanadian

HeritageformedtheBisonResearchandContainmentProgram

(BRCP)tofocusondiseasecontainmentandecologicaland

traditionalknowledgeresearch(RAC2001).TheMinisterthen

createdtheResearchAdvisoryCommittee(RAC)tocoordinate

researchactivitiesundertheBRCP(Chisholmet al.1998).The

RACcomprisedaseniorscientistappointedbyParksCanada,

representativesfromtheAlbertaandNorthwestTerritories

governments,CanadianParksandWildernessSociety,and

fouraboriginalcommunities(Chisholmet al.1998).During

themandatedfiveyearperiod(1996-2001),theBRCPfunded

projectstoassesstheprevalenceandeffectsofthediseases

onnorthernbison(JolyandMessier2001a),andtoinvestigate

bisonmovementsandtheriskofdiseasetransfer(Gateset

al.2001a).TheRACproducedafutureresearchagendaand

budgetforminimumresearchstillrequiredundertheBRCP

mandate(RAC2001),buttheprogrammewasdiscontinuedin

2001.ManyoftheresearchneedsidentifiedbytheRACalign

withtherecommendationsoutlinedintheNationalRecovery

PlanforWoodBisonpreparedbytheWoodBisonRecovery

Team(Gateset al.2001c).Thereremainsconsiderable

disagreementbetweenfederalandprovincialgovernments

andaboriginalinterestsconcerningalong-termsolutionto

theWBNPdiseaseissue.Provincialgovernmentssupport

diseaseeradication,includingaggressiveinterventionto

achievediseaseeradicationwithinthenationalpark.Parks

Canadaisconcernedabouttheconservationandbiological

impactsassociatedwithaggressiveintervention.Atechnical

workshopwasconvenedin2005toexplorethefeasibility

ofremovingdiseasedbisonfromtheGreaterWoodBuffalo

NationalParkregionfollowedbyareintroductionofhealthy

bison(Shuryet al.2006),andtherewasunanimousagreement

amongstparticipantsthatthisoptionwastechnicallyfeasible.

Theonlysubsequentmanagementactionundertakenatthe

timeofwritingwastheimplementationofahuntingseason

fortheHay-Zamaherdin2008-2009,intended,inpart,to

testdiseasestatusandtoreducetheriskofinfectionwith

BTBandbrucellosisbyreducingpopulationsizeandlimiting

rangeexpansiontowardsinfectedpopulations(George

Hamilton,AlbertaSustainableResourceDevelopment,personal

communication).

5.4 Disease Management in perspective

Aprimaryconsiderationregardingdiseasemanagement

inwildpopulationsisdeterminingwhenadiseaseisa

conservationproblemandwhetherinterventioniswarranted

(GilmourandMunro1991).Itcanbearguedthatparasitism

bydiseaseorganismsisacrucialecologicalandevolutionary

forceinnaturalsystems(Aguirreet al.1995;Wobeser2002).

Classificationofapathogenasindigenousorexotictoahost

speciesorecosystemcaninfluencewhetheradiseaseshould

bemanaged(AguirreandStarkey1994;Aguirreet al.1995;

NationalParkService2000).BTBandbrucellosisarebelievedto

havebeentransmittedtobisonfromdomesticcattle.Therefore,

managementofthesediseasesinbisoniswarrantedbasedon

theirexoticorigins,aswellasthethreattheyposetodomestic

animals.However,manyotherpathogenshavecoevolvedwith

bisonanddonotwarrantveterinaryinterventionandshouldbe

managedinaccordancewithanaturalsystem.

Themostsignificantdiseasesinvolvingbisonaswildlifeaffect

atrinityofplayers(wildlife,humans,anddomesticanimals),

andinvolveatangleoftransmissionroutes(Fischer2008).

Managementofwildlifediseaseshasoftenbeenundertaken

tominimiseriskstohumansanddomesticanimals(Nishiet

al.2002c;Wobeser2002).Reportablediseasemanagement

foragriculturalpurposesistypicallybasedontheobjective

oferadicatingthediseasefromalivestockpopulation

(Nishiet al.2002c).Thepolicyandlegislativeframeworkfor

eradicatingreportablediseasesindomesticanimalsiswell

developed,however,whenappliedtowildlife,theprotocols

usedbyagriculturalagenciesareusuallynotcompatiblewith

conservationgoals(e.g.,maintaininggeneticdiversity,minimal

managementintervention)(Nishiet al.2002c).Increasingly,the

broaderconservationcommunityisexaminingwildlifedisease

issuesinthecontextoftheirimpactontheviabilityofwild

populations,conservationtranslocationprogrammes,andglobal

biodiversity(DaszakandCunningham2000;Deemet al.2001;

Wobeser2002).Creativedisease-ecologyresearchisneeded,

andanadaptivemanagementframeworkisrequiredforcoping

withdiseaseswithinaconservationcontext(Woodruff1999).

Anevaluationofthediseasemanagementmethodspresently

appliedtobisonpopulationsisneededandcouldassist

withdevelopmentofnovelconservation-appropriatepolicies

andprotocolsformanagingthehealthoffree-rangingbison

populations(Nishiet al.2002c).

Twoemergingpolicyconceptsbeingdiscussedtomanage

andcontrolthetransmissionordistributionofdiseaseatthe

domestic/wildanimalinterfaceincluderegionalisationand

compartmentalisation(CFIA2002;OIE2008).Regionalisation

offersonemeansofspatiallyidentifyingwherediseasecontrol

measureswilloccuronthelandwhilecompartmentalisation

separatesthecontrolprogrammesofwildanddomesticanimals.


Theseconceptsarebeingdevelopedandputintopracticeby

state/provincial,federal,andinternationalhealthagenciesto

addressthecomplicationsofmanagingintractabledisease

problemsinwildanimalsrangingonlargelandscapesthat

alsosustaindomesticlivestockindustriesandassociatedlocal

economies(Bengiset al.2002).

Nationalwildlifehealthstrategieshaverecentlybeendeveloped

inCanadaandtheU.S.inresponsetothemanydifficultdisease

issuessurroundingfree-rangingwildlife.Thedevelopmentof

nationalwildlifehealthprogrammesparalleledtheincreasing

profileofwildlifehealthissuesinsocialandpoliticalarenas.

Thesenationalstrategiesneedtoprovideclearguidancefor

coordinatedconservationactionandacountrywidelegislative

andpolicyframeworkthatwillinfluencebisonrestorationand

conservationeffortsinNorthAmerica.Itishopefulthatmounting

tensionbetweentheagriculture,human,andwildlifehealth

communitiescanbemitigatedbydevelopingacomprehensive

nationalwildlifehealthpolicy,supportivescientificresearch

programmes,broadstakeholderengagementindecision

processes,aconservation-sensitiveregulatoryframework,and

opensocialdiscussionaboutthediseaserisksfromwildlife.


Chapter6 GeneralBiology,EcologyandDemographics

Lead Authors: Peter J.P. Gogan, Nicholas C. Larter, James H. Shaw, and John E. Gross

Contributors: C. Cormack Gates and Joe Truett

6.1 General Biology

Anunderstandingoftheecologyandbiologyofbisonis

fundamentaltotheirsuccessfulmanagement,conservation,

andrestoration.Bisonhavethebroadestoriginalrangeof

anyindigenousungulatespeciesinNorthAmerica,reflecting

physiological,morphological,andbehaviouraladaptationsthat

permitthemtothriveindiverseecosystemsthatprovidetheir

dietofgrassesandsedges.Successfulpopulationmanagement,

conservationofgeneticdiversityandnaturalselection,modelling

andpredictingpopulationlevelresponsestohumanactivities,

andmanagingpopulationstructurealldependonunderstanding

thebiologicalcharacteristicsandecologicalrolesofbison.The

purposeofthischapteristosummarisewhatiscurrentlyknown

aboutthebiologyofbison;foranearliercomprehensivereview,

seeReynoldset al.(2003).

6.1.1 physiology

6.1.1.1 Metabolism

Bisonexhibitseasonalvariationinenergymetabolism.

Christophersonet al.(1979)andRutleyandHudson(2000)

observedthatmetabolisableenergyintakeandrequirementsof

yearlingmalebisonweremarkedlylowerinwinterthansummer.

Thiswasattributedtoareductioninactivityandacclimation.

Bisonarebetteradaptedtotemperatureextremesthanmost

breedsofcattle.Theyexpendlessenergyunderextreme

coldthandocattlebecauseofthegreaterinsulating

capacityoftheirpelage(PetersandSlen1964).

Coldtoleranceofhybridsbetweenbisonandcattle

isintermediatebetweenthetwospecies(Smoliak

andPeters1955).Toleranceofbisontoheathasnot

beenstudied,buttheoriginalcontinentalrangeof

thespeciesincludedthedry,hotdesertgrasslandsof

northernMexico,whereasmallpopulationofplains

bisonstillexiststoday(Listet al.2007).

Figure 6.1 Age-specific live-weights of male and female plains

bison at Wind Cave National Park, South Dakota, obtained at

fall roundups 1986–1989 and 1991–1999. Data courtesy D.

Roddy and B. Meunchau, Wind Cave National Park.

6.1.1.2 Growth

Birthweightsofintensivelymanagedplainsbisonhavebeen

reportedas25kgforfemalesand30kgformales(Agabrielet al.

1998;AgabrielandPetit1996;Rutleyet al.1997).Birthweights

(near-termfoetuses)offree-rangingplainsbisonrangefrom14

to32kg(McHugh1958;Meagher1986;Park1969).Goganet

al.(2005)estimatedthatthebirthweightoffree-rangingbison

calvesisonaverage10%lessthanthatofcaptivebison.Growth

fromcalfhoodtoadulthoodfollowedasimilarpatterntothatof

adults,withweightgainduringthesummerandlossduringthe

winter(P.J.Gogan,unpublisheddata).Weightgainamongcalf

andyearlingplainsbisonwasaffectedbytheinfluenceofthe

timingandmagnitudeofsummerprecipitationongraminoid

physicalstructure(Craineet al.2009).

Differencesinweightsofplainsbisoningeographicallyseparate

herdshavebeenattributedtodifferencesinclimate,nutritional

plane,andgeneticlineages(BergerandPeacock1988;Lottand

Galland1987).AtElkIslandNationalPark(EINP),femaleplains

andwoodbisonachievedasymptoticbodyweightbysixyears

andmaximumbodyweightat10years(Olson2002;Reynolds

et al.2003).FemaleplainsbisonatWindCaveNationalPark

(WCNP)reachedanasymptoticbodyandmaximumbodyweight

atfiveyears(Figure6.1).MaleplainsandwoodbisonatEINP

reachedanasymptoticbodyweightateighttonineyearsand

maximumbodyweightby13years(Reynoldset al.2003).Male

plainsbisonatWCNPcontinuedtogainweightthroughthe


2001;OlexaandGogan2007).PlainsbisonwithintheGreater

YellowstoneAreashowstrongfidelitytosubpopulations

(Christiansonet al.2005;Goganet al.2005;OlexaandGogan

2007)asdowoodbisonintheGreaterWoodBuffaloEcosystem

(GWBE)(Carbynet al.1998;2004;ChenandMorley2005;Joly

andMessier2004).Bisonwithinsubpopulationsshowstronger

cohesionandcoordinatedmovementsduringsummerthanin

winter(ChenandMorley2005;OlexaandGogan2007).

6.1.2.2 reproductive behaviour

Sexuallymaturemaleplainsbisonjoinmixed-sexandage

aggregationsduringtherut.Dominantbullsformso-called

“tendingbonds”withindividualcowsjustpriorto,orduring,

oestrus(Fuller1960;McHugh1958;Meagher1973).Thebullwill

typicallyattempttokeepotherbullsawayandtokeepthecow

neartheedgeofamixed-sexandagegroupuntilsheaccepts

copulation(BergerandCunningham1994;Lott2002;McHugh

1958).Maturemalesmoveawayfrommixed-sexandagegroups

attheendoftherut(BergerandCunningham1994;Lott2002).

Woodbisonalsoaggregateduringthesummer(JolyandMessier

2001;Komerset al.1992).Malewoodbisonbecomemore

solitarywithincreasingage,aremorefrequentlyaggressive,and

testfemalesforoestrusmorefrequentlythandoyoungerbulls

(Komerset al.1992).Duringtherut,maturemalesjoinmixedsex

andgroupstocompeteformatingopportunitiesandtemporarily

leavethesegroupstorecoverfromhighcostbreedingactivities

(Komerset al.1992).Intheexperimentalabsenceofmature

malesduringtherut,subadultmalesfedlessandinteracted

moreaggressivelythanwhenmaturemaleswerepresent

(Komerset al.1994).

6.1.2.3 Cow-calf behaviour

Femaleplainsbisonclosetoparturitionhavebeendescribed

asrestlessandexcitable(McHugh1958).Apregnantcowmay

firsteightyears(Figure6.1).Whiledifferencesamong

populationsinbodysizeandweightmaybeapparent

toanobserver,comparisonsmusttakeintoaccount

theannualcycleofweightgainandloss.

6.1.2 Behaviour

6.1.2.1 Social structure

Therearemanyhistoricalobservationsofhugeplains

bisonherdsroamingtheGreatPlains(Dary1989;

Hornaday1889;Isenberg2000;Roe1970).Observers

ofbothplainsandwoodbisonconsistentlyreport

adefinableherdstructurewherecows,calves,and

immaturemalesformunstablemixed-sexandagegroups,and

largebullsformseparate,smallergroupsthroughoutmuchof

theyear(Allen1876;BergerandCunningham1994;Komers

et al.1993;Meagher1973;Meltonet al.1989;Schuleret al.

2006).Seasonalvariationsingroupsizesareassociatedwith

abundanceordispersionofforage(Jarman1974;Schuler2006),

landscapefeatures(BergerandCunningham1994),breeding

behaviour(BergerandCunningham1994;Meagher1973;Melton

et al.1989;Komerset al.1993)andpopulationsize(Schuleret

al.2006).Thelargestaggregationsoccurduringthebreeding

seasonwhenmaturebullsjointhemixed-sexandagegroups.

MeangroupsizesduringtheAugustrutatBadlandsNational

Parkrangefromameanof157inflatterrainto79inbroken

terrain(BergerandCunningham1994).Meanmaximumgroup

sizesatYellowstoneNationalPark(YNP)increasedfrom140

inMaytomorethan250inSeptember(Hess2002).Groupsof

morethan1,000bisonhavebeenobservedduringtherutin

contemporaryOklahoma(Schuleret al.2006).Groupsizerapidly

diminishesduringautumninplainsbison(Hornaday1889)to

fewerthan30(BergerandCunningham1994;Schuleret al.

2006).Similarly,inwoodbison,typicalgroupsizeisgreatest

duringthepre-rutandrut,thendeclinesduringthefall(Komers

et al.1992).MeanmaximumgroupsizesatYNPdeclined

throughoutwinterfrommorethan250inDecemberto16inApril

astheareaoccupiedbybisonincreasedfrom1,000tomore

than1,200km2(Hess2002).

Malebisonformtemporary,unstablegroups,andexhibitalinear

dominancehierarchy,witholder,heavieranimalsdominantover

youngersmallermales(Komerset al.1994;Rodenet al.2005).

Dominanceisalsorelatedtoageinfemalebison(Rutberg1983).

Groupsofadultorsubadultmalesrarelyexceed10individuals

(BergerandCunningham1994).

Plainsandwoodbisonpopulationsubstructureoccursata

broadgeographicalscaleduetotraditionaluseofparticular

partsofarangebysegmentsofapopulation(JolyandMessier

plate 6.1 Plains bison bull tending a cow, Jackson Valley,

Wyoming. Photo: Cormack Gates.


rubbinganobject,typicallyashruborsmalltree,withits

head,horns,neck,orshoulders(CoppedgeandShaw1997).

Wallowinginvolvesabisonrollingindrylooseground(or

lessfrequentlyinwetground)andtearingattheearthwith

itshornsandhoovesasitrolls.Bisonprefertohornaromatic

shrubsandsaplings(CoppedgeandShaw1997;Edwards

1978;McHugh1958;Meagher1973),whichmayhaveinsect

deterrentproperties.Bisonhaveevenbeenobservedrubbing

ontreatedtelephoneposts(CoppedgeandShaw1997).Soper

(1941)observedthathorningandrubbingwereoftenassociated

withharassmentbyinsects.Likewallowing,horningmayalso

constituteaggressivedisplaybehaviour.

Bisonofbothsexesandallageclassesengageinwallowing

behaviourthroughouttheyear(Reynoldset al.2003),although

sexuallymaturemaleswallowmorefrequentlyduringtherut,

urinatinginthewallowbeforepawingandrolling(Lott2002;

McHugh1958).Wallowingbymaturemalesmaystimulate

oestrusinfemales(Bowyeret al.1998),andadvertiseamale’s

physicalconditiontoothermales(Lott2002).Plainsbison

mayalsowallowtocoolthemselvesduringthehotsummer

months,ortoachieverelieffrombitinginsects(McMillanet al.

2000;MooringandSamuel1998).Catlin(inHornaday1889)

describedbisoncreatingwallowsinareaswithahighwater

tableandrollinginthewallowasitfilledwithwater.Theresult

waspelagemattedwithmudandclay(CatlininHornaday1889).

Coatshedding,rut,andinsectharassmentoccursimultaneously

duringthesummer;thereforeintheabsenceofcontrolled

experimentation,itisnotpossibletodeterminetherelative

influenceofthesefactorsonthefrequencyofhorningand

wallowing(CoppedgeandShaw1997).

6.1.2.5 Movements

Plainsbisonfrequentlytravelinsinglefilealongwell-established

trailswhenmovingbetweenforagingpatches(Garretson

1938;Hornaday1889).Historically,plainsbisonundertook

leavetheherdpriortocalvingorgivebirthwithintheherd

(McHugh1958).Similarly,forwoodbisonintheMackenzie

BisonSanctuary(MBS),femaleshavebeenobservedcalving

inthemidstofherdsorinextremeisolationintheforestaway

fromanyotheranimals(N.C.Larter,personalobservation).

Birthingnormallyoccurswhilethefemaleislyingdown.The

mothertypicallyconsumesportionsoftheafterbirthasshe

freesthecalffromthemembranes(Lott2002;McHugh1958).

Thefemalelicksamnioticfluidfromthecalf’sfur(Lott2002).

Sucklingbeginsshortlyafterbirthandmaylastaslongas

10minutes(McHugh1958);althoughtherewasareportofa

woodbisonmotherattackingthenewborncalfduringsuckling

(CarbynandTrottier1987).Theclosecontactbetweena

cowandcalfbeginstodeclineafterthecalf’sfirstweekof

life(Green1992).Acalfistypicallyweanedbyseventoeight

monthsofage,althoughnursingmayextendbeyond12

months(Greenet al.1993).Thelongestassociationsamong

bisonarebetweencowsandtheirfemaleoffspring;whilemale

offspringmayremainwiththecowthroughasecondsummer,

femaleoffspringmayremainwiththecowthroughathird

summer(Greenet al.1989;ShawandCarter1988).

Thecowmayusequickchargesorsteadyadvancestodefend

acalfagainstthreats(Garretson1938;Hornaday1889;McHugh

1958).Anisolatedplainsbisoncowvigorouslydefendedher

calffromagrizzlybear(Ursus arctos),eventhoughthebearwas

ultimatelysuccessfulinkillingthecalf(VarleyandGunther2002).

Similarly,anisolatedcowvigorouslydefendedthecalffrom

wolves(Canis lupus)(C.Freese,personalcommunication).

Cowsandothermembersofmixed-sexandagegroupsmay

cooperativelyprotectcalvesfrompredators.Inresponsetothe

approachofagrizzlybear,amixed-sexandagegroupofadult

plainsbisonrespondedbyfacingthebearinacompactgroup,

withthecalvesrunningbehindtheadults(Gunther1991).Wolves

preferentiallyattempttopreyuponwoodbisonmixed-sexand

agegroupsthatincludecalves(CarbynandTrottier1987).During

wolfattacks,calvesmovedclosetothecow,or

tootherbison,ortothecentreofthebisongroup

(CarbynandTrottier1987;1988),althoughthis

defensiveresponsemaybreakdownwhenbison

groupsmovethroughforestedareasthatmay

impedethemovementsofthecalves(Carbynand

Trottier1988).

6.1.2.4 horning and wallowing

Allageandsexclassesofbisonengagein

behavioursreferredtoashorningandwallowing

(McHugh1958).Horninginvolvesananimal

plate 6.2 Wallowing modifies the landscape. Photos:

Dwight Lutesy (inset) and John Gross.


extensiveseasonalnorth-southmovementsfromsummerto

winterranges(Seton1929)onbothsidesoftheMississippi

River(Garretson1938;Roe1970)andfromtheprairiesintothe

Parkland(Campbellet al.1994).Largeherdsalsoremainedon

thenorthernprairiesthroughoutwinter(MalaineyandSherriff

1996).Rivervalleyswerecrucialtothesurvivalofbisonover-

winteringonthegrasslands(West1995).Plainsbisonalso

undertookseasonaleast-westmovementsfromtheprairies

tothefoothillsoftheRockyMountainsinwinter(Garretson

1938).Inferencesfromhistoricalreportsofseasonalmovement

patternsareconfoundedbythetimingoftheaccountrelativeto

theimpactsofmarkethunting,establishmentofpioneertrails,

andconstructionoftherailroads(Roe1970).Insummer,bison

ontheGreatPlainsmovedtowateronanalmostdailybasis,

andonoccasionmovedfrom80to160kilometresoverseveral

daystoaccesswater(Dary1989).

PlainsbisoncurrentlyoccupyingtheYNPspendsummerat

higherelevationsandmovetowinterrangesatlowerelevations

(Auneet al.1988;Gateset al.2005;Meagher1973;Olexaand

Gogan2007).Thesemovementsaremadeoveranetworkof

trails,geothermalfeatures,andalongthebanksofriversand

streams,oralonggroomedroadwaysalignedwithnaturaltravel

routes(BjornlieandGarrott2001).Adultmalesareoftenthefirst

topioneerpreviouslyunoccupiedareas,abehaviourthathas

beenobservedinbothwoodbisonandplainsbison(Gateset al.

2005).Yellowstonebisonhaveexpandedtheirrangeinresponse

toincreasedpopulationdensities(Taperet al.2000)exacerbated

byparticularlyseverewinters(Meagher1989).

WoodbisonatWoodBuffaloNationalPark(WBNP)annually

travelupto50kilometresmaximumfromacentreofactivity

(ChenandMorley2005),andindividualwoodbisonattheMBS

rangeoverareasof179to1,442km2(LarterandGates1990).

Woodbisonhaveslowlybeenexpandingtheirrangeinthe

northernborealforest.Rangeexpansionisgenerallyinitiated

bylargemaleswhothenseasonallyreturnfromtheperipheries

oftherangetojoinfemalesandjuvenilesduringtherut(Gates

andLarter1990;N.LarterandJ.Nishiunpublisheddata).

Subsequently,mixed-sexandgroupsmoveintotheexpanded

peripheralrange.Rangeexpansiontypicallyfollowsperiodic

highlocalpopulationdensities(GatesandLarter1990)andis

density-driven(Gateset al.2005).

6.2 Ecology

6.2.1 plains bison

6.2.1.1 Ecological role

MillionsofplainsbisonhistoricallyrangedoverNorthAmerica’s

grasslandsandfunctionedasakeystonespecies(Knappet al.

1999).Theysharedthislandscapewithavarietyofotherlarge

mammalsincludingpronghorn(Antilocapra americana),elk

(Cervus elaphus),deer(Odocoileusspp.),wolves,andgrizzly

bears.Atthelandscapelevel,bisonservedasecosystem

engineers,bothrespondingto,andcreating,heterogeneity.

Anestimated100millionbisonwallowshadamajoreffecton

surfacehydrologyandrunoff(Butler2006).Ephemeralpoolsof

standingwaterthatpersistedinwallowsformanydaysfollowing

springsnowmeltorrainstorms(Knappet al.1999)supporteda

varietyofwetlandplantspecies(CollinsandUno1983;Polley

andWallace1986).Similarly,bisonwallowsprovidedimportant

breedinghabitatfortheGreatPlainstoad(Bufo cognatus;Bragg

1940)andtheplainsspadefoottoad(Spea bombifrons;Cornand

Peterson1996).Bisondirectlyaffectvegetationcommunities

throughtheirgrazing,physicaldisturbance,andbystimulating

nutrientrecyclingandseeddispersal(McHugh1958).Such

activitieshelptomaintainmeadowsandgrasslandsonwhich

they,andmanyotheranimalandplantspecies,depend.

Intallgrassprairie,bisongrazingofgrassesincreasedsoil

temperature,lightavailability,andsoilmoistureavailabilityto

forbspecies(FahnestockandKnapp1993).Thenetresult

wasbeneficialtoforbsnoteatenbybison(Damhoureyehand

Hartnett1997;FahnestockandKnapp1993),andmaythereby

havebeenbeneficialforotherherbivoressuchaspronghorn.

Bisongrazingofshortandmixed-grassprairievegetation

increasedtheratesofnutrientcycling(DayandDetling1990),

modifiedplantspeciescomposition(CoppockandDetling

1986)andincreasedthenutritivevalueofgrasses(Coppock

et al.1983a;1983b;Krueger1986).Locally,bisonconsumed

forageresources(EnglandandDeVos1969;Hornaday1889)

andreducedforageheighttolevelsthatfacilitatecolonisation

byprairiedogs(Cynomysspp.;VirchowandHygnstrom2002).

Inturn,prairiedogactivitiesenhancedtheratioofplantlive:

deadmaterial,crudeproteincontent,anddigestibility(Coppock

et al.1983a;1983b)andtherebyencouragedfurthergrazing

bybisonovermorethan20%ofthenaturalshortandmixed

grassprairie(WhickerandDetling1988).Whilebisongrazing

wasindependentofpocketgopher(Geomyidae)activities,it

influencedgopherdistributionbymodifyingthedistributionand

abundanceofpatchesofforbsusedbygophers(Steuteret al.

1995).

Bisongrazing,frequentlyinconjunctionwithfireandwallowing,

enhancedthegrasslandheterogeneitynecessarytoprovide

suitablenestingsitesforavarietyofobligategrasslandnesting

birdspecies(Knappet al.1999).Bisongrazing,particularlyon

recentlyburnedareas,enhancestheabundanceofbreedingbird

species,suchasuplandsandpipers(Bartramia longicauda)and

grasshoppersparrows(Ammodramus savannarum),intallgrass

prairie(Fuhlendorfet al.2009;Powell2006).Similarly,anumber

ofbirdspeciesendemictotheshortandmixedgrassprairies

ofNorthAmerica,suchasthemountainplover(Charadrius

montanus)andMcCown’sLongspur(Calcarius mccownii),were


whichmaydiffermarkedlyfrompristineconditions(Fahnestock

andDetling2002).

Herbivores,includingbison,respondtogradientsinforage

qualityandquantity.Hornaday(1889)describedahighly

nomadicforagingstrategy,whereplainsbisonseemedto

wandersomewhataimlesslyuntiltheylocatedapatchwith

favourablegrazing.Abisonherdwouldthenremainandgraze

untiltheneedforwatermotivatedfurthermovement.This

accountcontrastswithmorerecentstudiesofbisonforaging,

whichhavefoundthatplainsbisonactivelyselectmore

nutritiousforages,andforageinahighlyefficientmannerthat

satisfiestheirnutritionalneedsandcomplimentsdietselection

bysympatricherbivores(Coppocket al.1983a;1983b;Hudson

andFrank1987;SingerandNorland1994;Wallaceet al.1995).

Spatialvariationinforagequalityandquantityresultsfrom

naturalgradientsinsoilmoisture,soilnutrients,fire,andother

disturbance,aswellasfromtheimpactsofforagingbybison.

Bisonexploitvariationsinforagequalityandquantityatall

scales;fromselectingsmallpatchesofhighlynutritiousforages

onprairiedogtowns,toundertakinglong-distancemigrationin

responsetoseasonalsnowfallordrought.

Thefollowingreviewofbisonhabitatinteractionsisbasedupon

NorthAmericanecoregionsidentifiedbyRickettset al.(1999)

andaggregatedbySandersonet al.(2008).

historicallydependentonacombinationofbisonwallowsand

prairiedogcoloniesfornestingsites.Theseareaswerealso

utilisedbyferruginoushawks(Buteo regalis)andlong-billed

curlew(Numenius americanus)(Knopf1996).Brown-headed

cowbirds(Molothrus ater),alsocalledbuffalobirds,occurred

inassociationwithbisonthroughoutcentralNorthAmerican

grasslandspriortotheintroductionoflivestock(Friedman1929).

Cowbirdsfeedoninsectsmovinginresponsetoforagingbison

(GoguenandMathews1999;Webster2005).Grasshopper

speciesrichness,composition,andabundancearestrongly

influencedbyinteractionsbetweenbisongrazingandfire

frequency(Joern2005;JonasandJoern2007).

Bisonfacilitateddispersaloftheseedsofmanyplanttaxaasa

resultoftheseedsbecomingtemporarilyattachedtothebison’s

hair(Berthoud1892;Rosaset al.2008)orviapassagethrough

thedigestivetract(Gokbulak2002).Peakpassagerateforseeds

was2daysfollowingingestion(Gokbulak2002).

Horningdamagetotreesalonggrasslandbordersiseffective

inslowinginvasionoftreesintoshrubandgrasslandplant

communitiesorinextendingtheexistinggrasslandintothe

forestmargin.BisonwithinYNPrubbedandhornedlodgepole

pine(Pinus contorta)treesaroundtheperipheryofopen

grasslandstotheextentthatsomewerecompletelygirdled

(Meagher1973).SimilarlyhorningbywoodbisonintheMBS

hasresultedincompletelygirdledwhitesprucestandson

theperipheryofmesicsedgemeadowsandwillowsavannas

(N.C.Larter,personalobservation).Severalauthors(Campbell

et al.1994;CoppedgeandShaw1997;Edwards1978)have

suggestedthatbison,incombinationwithotherfactorssuchas

fireanddrought,significantlylimitedthehistoricdistributionof

woodyvegetationontheGreatPlains.

Adecomposingbisoncarcassinitiallykillstheunderlyingplants,

butsubsequentlyprovidesapulseofnutrients,creatinga

disturbedareaoflimitedcompetitionwithabundantresources

thatenhancesplantcommunityheterogeneity(Towne2000).

Carrionfromdeadbisonisanimportantfoodresourceforboth

grizzlyandblackbears(Ursus americana)aswellasscavenging

birdssuchasbaldeagles(Haliaeetus leucocephalus),ravens

(Corvus corax),andblack-billedmagpies(Pica pica).

6.2.1.2 Contemporary habitat use, nutrition, and foraging

Thebisonisaruminantwithafour-chamberedstomachand

associationsofsymbioticmicroorganismsthatassistdigestion

offibrousforage.Onlowerqualityforage,suchasgrasses

andsedges,bisonachievegreaterdigestiveefficienciesthan

domesticcattle,butonhighqualityforagessuchasalfalfa,the

digestiveefficiencyofbisonandcattleconverge(Reynoldset al.

2003).Contemporarystudiesofplainsbisonhabitatselection

inNorthAmericangrasslandsarelimitedtoconfinedherds

artificiallymaintainedatvaryingdensities(Table6.1)—someofplate 6.3 Plains bison bull cratering in snow to forage. Photo:

Yellowstone National Park.


Ecoregion Location Seasonplant type

referenceGrasses (%)

Sedges (%)

Forbs (%)

Woody plants (%)

Others (%)

NorthernMixed

Grasslands

WindCaveNP,SD

Spring 81 7 9 3 Marlowet al.1984

Summer 79 9 10 2

Westfallet al.1993Autumn 77 12 6 5

Winter 79 12 2 7

Winter 59 37 4 WydevanandDahlgren1985

CentralShortgrass

Prairie

PawneeSite,CO

Lightlygrazed

Spring 98 2

Pedenet al.1974Summer 94 5

Autumn 99

Winter 94 4

Heavilygrazed

Spring 95 4

Pedenet al.1974Summer 96 4

Autumn 87 2 12

Winter 81 6 11

TallGrasslandsPrairieandSouthern

ShortgrassPrairie

WichitaMountainsNWR,OK

Spring&Summer 99 Buechner1950

TallgrassPrairie

Preserve,OK

Spring 60 39 1

Coppedgeet al.1998

Summer 88 11 1

Autumn 84 16 1

Winter 79 21 1

NorthernFescue

Grasslands

NationalBisonRange,

MTAnnual 90 1 2 1 McCullough1980

RockyMountainForests

YellowstoneNorthern

Range,WYWinter 53 441 1 1 SingerandNorland

1994

YellowstoneCentral

Range,WYSummer 55 37 <0.1 OlenickiandIrby

2004

NorthernForests

ElkIslandNP,AB

Spring 29 65 6 TelferandCairns1979Winter 18 82

PrinceAlbertNP,SK

Spring 35 65

Fortinet al.2002Summer 26 73 1

Autumn 17 63 20

Winter 34 59 7

table 6.1 Diets of plains bison at select locations within North American ecoregions.

1Includesrushes(Juncaceae)


6.2.1.2.1 Northern mixed grasslands

Intheabsenceoffire,bisonhavebeenobservedmaking

extensiveuseofprairiedogcoloniesinthenorthernmixed

grasslandsecoregion,wherecoloniesmayhavecovered2-

15%oftheshortgrasslands(Knowleset al.2002;Virchowand

Hygnstrom2002).Bisonutilisetheforb-dominatedcentresof

prairiedogcoloniesforrestingandwallowing,butfeedatthe

graminoid-dominatedperipheryofcoloniesratherthanatthe

colonycentre(CoppockandDetling1986;Krueger1986).Bison

useofprairiedogtownspeaksduringthesummeranddeclines

intheautumn(Krueger1986)whentheavailableforagebiomass

isloworthevegetationissenescent(Coppocket al.1983a;

1983b).Bisonuseofcolonysitesalsodeclineswhenrecently

burnedgrasslandsareavailable(CoppockandDetling1986).

Grassesandsedgeswerealmost90%oftheyear-roundbison

diet,andsedgesformed7to37%oftheseasonaldietinthe

northernmixedgrasslandecoregion(Table6.1).Bisonselected

foragingsitescontainingmorethan75%warmseason(C4)

grassesduringthesummergrowingseason(Steuteret al.

1995).C4grasseswereapproximately33%ofthedietinJune,

andamaximumof40%ofthebisondietinlatesummer,but

C4grasseswerelessinthebisondietinautumn,winter,and

spring(PlumbandDodd1993).Conversely,coolseasongrasses

formedapproximately50%ofthesummerdiet,butincreasedto

80%ofthedietinSeptember(PlumbandDodd1993).

6.2.1.2.2 Central shortgrass prairie

Inalightlygrazedsite,bisonalmostexclusivelyconsumed

grasses,butconsumedmorethan10%woodyplantsinthe

autumnandwinterataheavilygrazedcentralshortgrassprairie

sitesharedwithcattleandsheep(Table6.1).ThreeC4grasses

accountedfor65to75%ofthebisondiet(Pedenet al.1974;

SchwartzandNagy1976).

6.2.1.2.3 Tall grasslands prairie and

southern shortgrass prairie

Bisoninthetallgrasslandsprairieandsouthernshortgrass

prairieecoregionsutilisedonlyrecentlyburnedareasinspring,

butselectedareasburnedannuallythroughouttheyear(Shaw

andCarter1990;Vintonet al.1993).Bisongrazingandregrazing

canmaintainareaswithalowvegetativecoverandstanding

crop(CoppedgeandShaw1998;Vintonet al.1993).Areas

grazedbybisonwerecharacterisedbyalowerabundanceof

C4grasses,ahigherabundanceofC3grasses,andgreater

overallplantspeciesdiversity(Hartnettet al.1996).These

characteristicsweremorepronouncedinareasburnedannually

(Hartnettet al.1996),whichisconsistentwithgreaterbisonuse

ofannuallyburnedsites(ShawandCarter1990;Vintonet al.

1993).Bisongrazedlittlebluestem(Schizachyrium scoparium)

morefrequentlypost-burning,probablyinresponsetoremoval

ofstandingdeadtillersbyfire(PfiefferandHartnett1995).The

greateroverallplantspeciesdiversityinburnedareaswaslinked

toincreasednitrogencyclingandavailability(Bakkeret al.2003;

JohnsonandMatchett2001).

C3grasseswerethemostcommondietaryiteminwinter

(Coppedgeet al.1998).Dietaryquality,asmeasuredbyfaecal

nitrogen,peakedinMayandJune,coincidentwithapeakinC3

grassesproductivity(Postet al.2001).Upto39%ofthespring

dietwassedges(Coppedgeet al.1998).

6.2.1.2.4 Northern fescue grasslands

Understandingcontemporarytrophicecologyofbisoninthis

ecoregionisconfoundedsomewhatbyamanagement-imposed

rotationalgrazing,bywhichbisonaremovedthroughoutthe

NationalBisonRange(NBR)NationalWildlifeRefuge,Montana

(McCullough1980).Whenoccupyinglowerelevationareasofthe

NBR,bisonutilisedleveltoundulatingopengrasslands.Once

herdedtohigherelevationportionsoftherange,bisoncontinued

toutilisethemorelevelopenareasavailable(McCullough1980).

Theyear-rounddistributionofbisonwasawayfromhigher

elevationsteep-slopeareas.Bisonshowednoselectionfor

aspect,astheytendedtousethemorelevelareasavailable

throughouttheyear.Bisonfedalmostexclusivelyongrasses

(Table6.1;McCullough1980).

6.2.1.2.5 Rocky Mountain forest

InthehightopographicalreliefoftheRockyMountains

theheterogeneityofherbaceousproductivityandstanding

cropiscausedbythespatialdistributionofmoistureonthe

landscape.Herbaceousabovegroundnetprimaryproductivity

(ANPP)isinfluencedbysite-specifictopographicposition

relativetomoisturedistributionandaspect(Burroughset al.

2001).HerbaceousANPPisloweratlowelevationswithless

precipitationandatthehighestelevationsduetoashorter

growingseasonattributabletolowertemperaturesthanatmid-

elevations(Coughenour2005).Ingeneral,herbaceousANPP

occursasapulseofnitrogenrichvegetationthatsequentially

followsanelevationalgradientfromthelowerelevationwinter

rangestothehigherelevationsummerranges.Thispattern

ofANPPmakesyoungnutritiousandconcentratedforage

availabletobisonforuptosixmonthsofeachyear(Frank

andMcNaughton1992).Summermovementsofbisonto

higherelevationareasreducesvegetationutilisationatlower

elevationsandtherebyenhancestheavailabilityofvegetation

atlowerelevationsduringthenon-growingseason(Frankand

McNaughton1992).

BisononYellowstone’snorthernrangeforageonsedgeswithin

moremesicsitesinwinter(Meagher1973)totheextentthat

thewinterdietismorethan95%grasses,sedges,andrushes

(Table6.1;SingerandNorland1994).Similarly,bisonutilising

theYellowstonecentralrangeduringwinterprimarilyfeedon

sedgesalongtheedgesofthermallyinfluenceddrainagesand


atotherthermalfeatures(Meagher1973).Uplandsagebrush-

bunchgrasssitesareutilisedtoalesserextentinwinter

(Meagher1973).ThesummerdietofYellowstonebisonutilising

theHaydenValleywasmorethan90%graminoids,withone-

halfofthesebeingmesicgrasses,sedges,andrushes(Olenicki

andIrby2004).

6.2.1.2.6 Northern forests

BisonatEINParehighlyselectiveforuplandgrasslandsyear-

round,andtoalesserextent,selectsedgemeadowsinwinter,

andshrublandandaspenforestinspringandsummer(Cairns

andTelfer1980;TelferandCairns1979).Thebison’syear-

rounddietwasvirtuallyexclusivelyherbaceousvegetationwith

approximately80%ofthewinterdietand65%ofthesummer

dietsedges(Carexspp.;Table6.1;TelferandCairns1979).

PlainsbisonforagingatPrinceAlbertNationalPark(PANP)

selectedthesedgeCarex atherodes,andconsumedmore

sedgesthangrassesyear-round(Table6.1;Fortinet al.2002).

Theforagingstrategyfavouredshort-termenergygainoverlong-

termgainformostoftheyear(Fortinet al.2002).However,bison

alsoselectedCarexinspring,whenadietofmoredigestible

grasseswouldhaveenhancedshort-termenergygain(Fortinet

al.2002).Bisonmayavoidshiftsindiettofacilitatemaintaininga

consistentmicrobialrumenflora(Fortinet al.2002).

6.2.1.2.7 Arctic lowland taiga

IntroducedplainsbisonatDeltaJunction,Alaska,feedon

sedgesandfescuegrassesinwinter(CampbellandHinkes

1983).Incontrast,plainsbisonintroducedtothevicinityof

Farewell,Alaska,feedonwillows(Salixspp.)almostexclusively

insummer,andamixtureofwillowandshrubsintheautumn

(WaggonerandHinkes1986).Somepotentialexistsfor

competitionwithmoose(Alces alces)forwillowinriparian,

alluvialareas,althoughthetwospeciesselectshrubsofdifferent

sizes(WaggonerandHinkes1986).Thedrasticdifferences

betweenthedietofplainsbisonatDeltaJunctionandthoseat

Farewellaredirectlyrelatedtoforageavailability.TheFarewell

areaisalmostexclusivelyriparianwillowgrowthwithlittleinthe

wayofgraminoidsduetoadominantveryrockybraidedriver

substrate.Incontrast,theDeltaJunctionareaischaracterised

byextensivestandsofgrassesandsedgesanddomesticated

grains.Thesedifferencesunderscoretheimportanceofforage

availabilityininfluencingbisondiets.

6.2.1.3 habitat and dietary overlap

Originally,plainsbisonassociatedwithpronghorn(Allen1967;

Yoakum2004),elk(Miller2002)andmuledeer(Odocoileus

hemionus)throughoutmuchoftheirrange,andwithmoose

(Boer1997)alongthenorthernandhighelevationrange

limits.Ofthesympatricspecies,theseasonaldistributionsof

pronghornandplainsbisonweremostsimilar,buttheirdiets

weremostdivergent(SchwartzandNagy1976;McCullough

1980;Marlowet al.1984;WydevanandDahlgren1985;Singer

andNorland1994).Althoughthesetwospeciestendtohave

littledietaryoverlap,somecompetitionfortotalbiomassmay

occur(LovaasandBromley1972).Similarly,sympatricplains

bisonandmuledeermayoverlapinhabitatselectioninwinter

(CairnsandTelfer1980),buttheirdietsdiffer(McCullough1980;

WydevenandDahlgren1985;SingerandNorland1994).

Plainsbisonandelkexhibitextensiverangeoverlapinwinter

(CairnsandTelfer1980;Barmore2003),butlessinspringand

summer(CairnsandTelfer1980).Thedietsofbothspeciesare

predominantlygraminoidsfromautumnthroughspring,with

bisonfavouringsedgesandelkfavouringgrasses(Barmore

2003;SingerandNorland1994).Dietaryoverlapwithgrasses

continuesintothesummer(McCullough1980;TelferandCairns

1979),althoughthebison’sdietcontainsmoregrassandless

forbsandwoodyplantsthanthatofelk(Marlowet al.1984;

WydevenandDahlgren1985).

Plainsbisonanddomesticcattledietsweremostsimilarforgrass

consumptionduringtheautumnandwinteratalightlygrazed

shortgrasslandsite,andduringthespringatanearbyheavily

grazedsite(Pedenet al.1974).Bisonandcattlesummerand

autumndietsinashrub-stepperegionwerealmostexclusively

grasses(VanVuren1984;VanVurenandBray1983).Thedietsof

bisonanddomesticsheepweremostsimilarduringautumnata

lightlygrazedshortgrasslandsite(Pedenet al.1974).

6.2.2 Wood bison

6.2.2.1 Original distribution and ecoregions occupied

Zooarchaeologicalevidence,combinedwithdocumentary

recordsandoralnarrativesofaboriginalpeoplesinAlaska,

Yukon,andNorthwestTerritories,indicatethattheoriginalrange

ofwoodbisonincludednorthernAlberta,north-easternBritish

ColumbiaeastoftheCordillera,theNorthwestTerritoriessouth

andwestofGreatSlaveLake,theMackenzieRiverValley,and

largeareasofinteriorAlaska(Gateset al.1992;Lotenberg1996;

Stephensonet al.2001;vanZylldeJong1986).Theoriginal

distributionofwoodbisoninnorthernAlbertaandsouthern

NorthwestTerritoriescentredontheInteriorPlainsPhysiographic

Region,wheretheyrangedovertheinterconnectedand

overlappingglaciallakebasinsandmajorrivervalleys,where

soilconditionsareconducivetodevelopmentofsedge-grass

meadowplantcommunities(Gateset al.1992).Thetotalrange

ofwoodbisonwasmorerestrictedthanthatofplainsbison.

Contemporarywoodbisonherdsintheborealregionsexist

incomparativelynaturalsystems.Theyremainpartofafairly

diverse,largeungulatefauna,whichrepresentsthepreybase

forseveralpredators.Woodbisondistributionoverlapswith

thatofmoose,elk,borealandnorthernmountainecotypesof


woodlandcaribou(Rangifer tarandus caribou),white-taileddeer

(Odocoileus virginianus),muledeerandpossiblystonesheep

(Ovis dalli).Similarly,woodbisonareexposedtothefullsuite

ofpredatorsincludingwolf,grizzly,blackbear,wolverine(Gulo

gulo),cougar(Felis concolor),lynx(Felis lynx),andcoyote(Canis

latrans).Wolfpredationisanespeciallyimportantmortalityfactor

fornorthernbison(Carbynet al.1993;Larteret al.1994;Van

Camp1987).Furthermore,woodbisonmovementsaregenerally

notimpededbyfencesorotherlanduses.

6.2.2.2 Contemporary habitat relationships,

nutrition, and foraging

WoodbisonoftheNahannipopulationinthesouth-west

NorthwestTerritoriesmustcrosstheLiardRiverasitbisects

thebisonrangeforitsentirety.Animalsofbothsexesand

allageclassesfrequentlymakerivercrossings(Larteret al.

2003)makingthemsusceptibletogroupmortalityduring

springicebreakupandrapidsnowmelt.Bisonuseofsedges

associatedwithwetmeadowsandlakesinwinteralsomakes

themsusceptibletomassmortalitywhengroupsfallthrough

weakice.Atotalof177animalsdrownedintheMBSafter

breakingthroughthespringiceofFalaiseLake(Gateset al.

1991).AbnormallyhighJanuary2009temperatures(+12°C)

affectediceconditionswhichlikelycausedthedrowningof

upto13animalsoftheNahanniwoodbisonpopulation(N.C.

Larter,unpublisheddata).Springflooding,notablyatWBNP,has

causedthousandsofbisondeaths(Fuller1962).

Fire,especiallyinthenorthernborealregionmayimprove

foraginghabitatforbisonand,insomeareasoftheNorthwest

Territories,prescribedburninghasbeenusedasamanagement

toolforhabitatenhancement(Chownset al.1997).However,fire

mayplaylessofaroleinmaintaininglowlandmeadowsthan

sporadicflooding(Quinlanet al. 2003).

6.2.2.2.1 Northern forests

BisonatWBNPandSlaveRiverLowlands(SRL)utilisedmixed

woodlandsandaspenandpoplarstandsinterspersedwith

meadowsinsummer,anduplandmeadows,lowlandfloodplains,

anddeltamarshesinwinter(Soper1941).Theyfeedprimarily

ongraminoids(Table6.2)withtwogenera,sloughsedge(Carex

atherodes)andreedgrass(Calamagrostisspp.),makingupmost

oftheannualdiet(Reynoldset al.1978).Willowswere8%of

thesummerdiet(Reynoldset al.1978).Bisonselectivelygraze

standsofsloughsedgecharacterisedbyabiomasslevelthat

wouldprobablyminimisedailyforagingtime(Bergmanet al.

2001).

6.2.2.2.2 Subarctic boreal forests

Bisonexhibitsex-specificdifferencesinhabitatselectionwith

femalesfoundinmesicsedgemeadows55%ofthetime

inwinter(comparedtomales,38%)andwillowsavannas

77%ofthetimeinsummer(comparedtomales,48%),even

thoughthesetwoplantcommunitiescombinedconstitute

onlyabout5%ofthearea(LarterandGates1991;Matthews

1991).Bothsexesutilisedthemostabundantconiferous

forestinproportiontoitsavailabilityduringautumn(Larterand

Gates1991).Bisonfrequentareaswherefrozenlakes,ponds,

oxbowlakes,anddisturbedsitesprovidewinteraccessto

forage.Thebisondietvariedseasonallyfromamorediverse

combinationofgraminoidsandwoodyplantsorforbsinsummer

toapproximatelyone-thirdlichensandone-thirdgrassesin

autumn,toalmostexclusivelygraminoidsinwinter(Table6.2).

Suchfeedingpatternswereconsistentwithselectionforplants

withrelativelyhighavailablenitrogen(LarterandGates1991)

andtoenhanceshort-termenergyconsumption(Fortinet al.

2002).Thisfeedingpatternmayalsobeattributedtodedicating

timetoavoidinsectharassment,scanningforpredators,

maintainingthermalbalance,orsocialinteractions(Bergmanet

al.2001).

IntheNahannipopulationofsouth-westNorthwestTerritories,

bisonutilisehorsetails(Equisetum)insummer(LarterandAllaire

2007),aforagethatishighinnitrogen,butalsohighinsilica.The

highsilicacausesrapidtoothwear,resultinginteethwearingout

10yearsearlierthaninotherareas.

6.2.2.3 habitat and dietary overlap

Thereislittledietaryoverlapbetweenwoodbisonandthe

variousungulatespeciesthatshareitsrange.Competitionwith

moosemayoccurintheNorthwestTerritories,wherethebison’s

diethasahighbrowsecomponent.Bisonandborealcaribouin

NorthwestTerritories/YukonTerritorybotheatlichens,although

duringdifferentseasons.Caribouuselichenasadietstaple

inwinter,whereasbisonuseoflichenisinautumn,whenthey

disperseintothemoreforestedhabitats(LarterandGates1991).

FischerandGates(2005)concludedthatfoodcompetition

betweencaribouandbisonwaslowinwinter.

6.3 Demographics

Theabundanceofthefree-rangingpopulationsofplainsand

woodbison,soiconicforNorthAmerica,likelyfluctuated

considerablybylocationandthroughtime.Thesefluctuations

wereprobablydrivenbyasequenceofdensity-dependent

populationregulatoryfactors(Eberhardt1977;2002;Fowler

1981;1987;Gaillardet al.1998);reducedsurvivalofsubadults,

delayedageoffirstreproduction,declineinthereproductive

rate,andincreasedadultmortality.Thissequencewas

undoubtedlysetbackbydensityindependenteventssuchas

episodicdroughtsandseverewinters.Droughtsanddryseasons

ingeneralwerecharacterisedbywildfires,which,onoccasion,

killedbison(Dary1989;Isenberg2000).Winterswithdeep

snowandwarmingperiods,resultinginicecrustingontopof


thesnow,ledtomajordie-offsofbison(Dary1989).Thousands

ofbisonweredrownedinfloodsthatresultedfromthespring

meltingoflargesnowpacks(Dary1989).

Predationbywolvesmayhavebeenasignificantforce,taking

themostsusceptibleageandsexclassesatdifferenttimesof

year.Wolvesmayhavepreyedheavilyonbisoncalves(Flores

1991)andkilledoldersolitarymales(Dary1989).However,

predationmayhavehadlittleeffectonlargenomadicor

migratoryherdsofbison(Terborgh2005).Wolvesmaintaingroup

territoriesandbearaltricialyoung,traitsthatwouldhavemade

itimpossibleforwolfpackstosustainsufficientpressureona

wide-ranging,mobileprey(Terborgh2005).Grizzlybearskilled

somebison,occasionallyfromambush(Dary1989).

Priortotheavailabilityoffirearms,thesmallnumberofresident

humans,andtheirrelativelyineffectivehunting,limitedthehuman

tollonbison.Pedestrianharvestingwasmostlynon-selective

andinvolvedsurroundingordrivingofbisongroupsoverbison

jumps(Flores1991).However,bythelate17thcentury,firearms-

equippedtribesfromtheGreatLakesregionbeganmovingout

ontotheGreatPlains.Atthebeginningofthe19thcentury,

tribeswithhorseswerebeginningtoexertpressureonplains

bisonandselectforbreedingagefemales(Flores1991).Atthe

sametimeinter-tribalwarfareledtobufferzonesthatserved

asrefugiaforbison(Flores1991;MartinandSzuter1999).By

mid-1800s,anestimated500,000plainsbisonwerekilledfor

subsistence,andanadditional100,000werekilledfortheirhides

table 6.2 Diets of wood bison at select locations within North American ecoregions.

1Includesrushes(Juncaceae);2Lichens;3Equisetumspp.;4November/Decemberisearlywinter,January/Februaryismid-winter

annually(Isenberg2000).Bisonpopulationsbegantodeclineas

increasingnumbersofcattleandhorsesbegantocompetewith

bisonforforageandwater(Flores1991;Isenberg2000).

6.3.1 population structure

Bothplainsandwoodbisoncanbeclassifiedintosexand

ageclassesbasedonbodysizeandhornmorphology.Free-

rangingcalvesarereadilydistinguishablefromallotherage

classesbaseduponpelagecolourforthefirstthreemonths

oflife,buttheirsexescannotbedistinguished.Yearlingsmay

bedistinguishedfromadultsuntilaboutoneandahalfyears

old,baseduponbodysizeandconformation,whenexamined

atcloserange.Sexcanbedeterminedinanimalsmorethan

twoyearsoldonthebasisofhornmorphologyandhead

shape(BradleyandWilmshurst2005;Komerset al.1993),or

notingthepresenceorabsenceofapenilesheath,butagain

thisrequiresviewingfromcloserange(Carbynet al.1998).

Komerset al.(1993)describedcriteriafordistinguishing

betweensubadult(twotofouryearsold),mature,andold

bullsbasedonbodysizeandhornmorphology.Theresultsof

compositioncountsarefrequentlystandardisedasaratioof

selectedageandsexclassesper100adultfemales(Caughley

1977).Typically,withinpolygynousspeciessuchasbison,

adultfemalesarethemostabundantclassinapopulation

anddirectlydeterminethesizeoftheyoungestageclass

(McCullough1994).Thepresenceofnewcalvesinapopulation

issensitivetothetimingofthecountrelativetothecalving

Ecoregion Location Season

plant type

referenceGrasses (%)

Sedges1

(%)Forbs

(%)

Woody plants

(%)

Others (%)

NorthernForests

WoodBuffaloNPandSlaveLake,NWT

andAB

Spring 16 81 1 2

Reynoldsetal.1978,Reynolds.1976inReynolds

andPeden1987

Summer 24 59 8 8

Autumn 21 71 4 2

Winter 36 63 1

SubarcticBorealForests

MacKenzieBison

Sanctuary,NWT

Spring 6 68 1 26

LarterandGates1991Summer 11 53 2 28 62

Autumn 32 15 4 12 372

Winter 2 96 2

NahanniPopulation,

NWT

Summer 6 37 29 14 13

LarterandAllaire2007;Larter,N.C.unpublished

data

Autumn 19 58 7 12 43

EarlyWinter4 16 37 10 4 333

Mid-Winter4 2 89 4 3 23


season:WolfeandKimball(1989)reportedanincreaseinthe

percentageofcalvesfrom10.2%inlateMayto12.2%inlate

July(i.e.,counttooearlyandyoumaymisssome).

Similarly,segregationofageandsexclassesmayinfluence

estimatesofpopulationcomposition.Meagher(1973)reported

thatcalvesformed20%ofmixedageandmixedsexbison

herds,but11%ofthetotalYellowstonebisonpopulation.

Otherbiasesarealsopossible.Carbynet al.(1998)reported

anunweightedaverageof36calvesper100adultfemales

forbisoninDeltaAreaofWBNPfor1989-1996(Table6.3),

whileothersreportedbetween20and30calvesper100adult

femalesforthesameareaandduringthesametimeperiod

(BradleyandWilmshurst2005).Similarly,Carbynet al.(1998)

reportedanaverageof20yearlingsper100adultfemalesfor

thistimeperiod,whileothersreportedmorethan10yearlings

per100adultfemalesforonlyoneofthoseyears(Bradleyand

Wilmshurst2005).Thus,compositionestimatesneedtobe

interpretedwithconsiderablecautionandwouldbenefitby

inclusionsofconfidenceintervals.

Fewdatasetspermitevaluationofreproductivesuccessand

survivalofyounginrelationtopopulationdensities(Table6.4).

Thehigherratiosofcalvesandyearlingsper100adultfemales

intheMinkLakeareaofWBNPcomparedtoMBS(Table6.4)

reflectdifferencesbetweenincreasinganddecliningpopulations

(Larteret al.2000).Lowercalfandyearlingtoadultfemaleratios

werelinkedtoaperiodofpopulationdeclineatWBNP(Bradley

andWilmshurst2005).Reynoldset al.(2003)reporteddensity

dependentfecundityinbisonatEINP.

Informationontheagestructureoffree-rangingbison

populationsnotsubjectedtoregularcullingislimited.Wood

bisonattheMBSwereassignedtoageandsexclassesinJuly

1993:calvesandyearlingswerenotassignedtosexclasses,

allfemalestwoormoreyearsoldwereassignedtoasingle

category,andmalesmorethantwoyearsoldwereassignedto

oneoffouragecategoriesfollowingKomerset al.(1992).Here,

thepopulationagestructureispresentedwithanassumption

ofanequalsexratioincalvesandyearlings(Figure6.2).

Irrespectiveofthesex,therelativelylownumbersofcalvesand

yearlingssuggestalowrecruitmentrate(Figure6.2).

6.3.2 reproduction

Theageoffirstreproductionissensitivetonutritionalcondition

and,therefore,highlyvariable.Theproportionoffemalescalving

astwo-year-olds(conceivingasone-year-olds)rangesbetween

4-12%(Table6.5).However,femalebisontypicallyenteroestrus

astwo-year-olds,andgivebirthtotheirfirstcalfatthreeyears

(Table6.5).Maturefemalesinsomepopulationsreproduce

eachyear(Rutberg1984;ShawandCarter1989;Wolff1998),

althoughinotherpopulationsmaturefemalesmaynotbreed

insomeyears(Fuller1962;Green1990;Halloran1968;Soper

1941;VanVurenandBray1986;Wolfeet al.1999).Thisis

particularlytrueoffemalesbreedingastwo-tofour-year-olds

(Green1990).Fuller(1962)notedthatforwoodbisonintheHays

CampareaofWBNP,21%ofthefemalesmorethanthreeyears

oldatthetimeofparturitionwerelactating,butnon-pregnant,

whilethesamewastruefor9%ofthefemalesintheLakeClaire

areaofthepark.Thisproportionmayvarywithinthesame

populationatdifferentdensitiesofbisonandotherungulate

speciesrelativetoforageconditions(Halloran1968;Shawand

Carter1989).Theyoungborntofemalesfollowingayearofnot

breedingwerelargerandmorefecundthantheyoungoffemales

whobredthepreviousyear(GreenandRothstein1991).Females

continuetobreeduntilmorethan16yearsofage(Green1990).

Bisonaretypicallymonoparous,withtwinningreportedonly

occasionally(Reynoldset al.2003).

Malebisonmaintainedonsupplementalfeedarephysiologically

capableofbreedingasearlyas16monthsofage(Helbiget al.

2007),andthosenotreceivingdietsupplementsmaybreedat

twotothreeyearsold(MaherandByers1987).However,males

generallydonotbreeduntiltheyarefiveorsixyearsoldand

largeenoughtocompetewitholderandmoreexperiencedbulls

(Fuller1960;Komerset al.1994;Meagher1973;Rothsteinand

Griswold1991).

Theageoffirstsuccessfulreproductionmaybemodified

bydiseaseinbisonoftheJackson,YellowstoneandGWBE

populations.Morethan90%ofthefirstpregnancieswere

lostinbrucellosisinfectedcaptivefemalebison(Daviset al.

1990;1991).Infree-rangingbison,theimpactofbrucellosis

ontheageoffirstsuccessfulreproductionwillvarywith

theproportionoffirsttimebreedersinthepopulation,the

proportionofthosebreedersinfectedwithbrucellosis,and

theseverityoftheinfection(BradleyandWilmshurst2005).

Diseasesmayalsomodifyreproductiveperformanceofolder

females.AtWBNP,bothtuberculosis(BTB)andbrucellosis

mayimpactthereproductivesuccessoffemalesofallage

classeswithinselectpopulationsegments(JolyandMessier

2004;2005).Intwopopulationsegmentsofwoodbisonat

WBNP,infectionwithbrucellosisorBTBalonedidnotimpact

pregnancystatus,butinfectionwithbothdiseasesreducedthe

probabilityofpregnancyby30%(JolyandMessier2005).Ina

thirdpopulationsegment,infectionwithBTBalonereducedthe

probabilityofpregnancyby75%(JolyandMessier2005).

6.3.3 Mortality factors and survival

Proximatecausesofmortalityincontemporarywoodbison

herdsincludewolfpredationandtheexoticdiseasesbrucellosis

andBTB(Fuller1962;Calef1984;Carbynet al.1993;Jolyand

Messier2001,2004;2005;Wilsonet al.1995inBradleyand

Wilmshurst2005).Inaddition,somewoodbisonsuccumbto

irregularoutbreaksofanthrax(Bacillus anthracis)(Gateset al.


table 6.3 Ratios of select age classes:100 females among plains and wood bison populations.

Subspecies Locationperiod of

Observation

Adult Sub-adult Sub-adult

referenceMale Female Male Female Yearling Calves

PlainsbisonHenry

Mountains,UT

JulyorSeptemberweightedaverage

1977–1983

54 100 43 53VanVurenandBray

1986

Woodbison

SlaveRiver,NWT Summer1978 32 100 4 1 7 35

VanCampandCalef

1987

MackenzieBison

Sanctuary,

NWT

July1993 78 100 Gateset al.1995

MackenzieBison

Sanctuary,

NWT

July,unweighted

average1984–1998

100 22 41 Larteret al.2000

MinkLake,NWT

July,unweighted

average1989–1998

100 30 51 Larteret al.2000

WoodBuffalo

(DeltaArea),AB

Springunweighted

average1989–1996

100 20 36 Carbynet al.1998

1995).Wallowsmayserveasfocalareasforanthraxspores,

andmorefrequentwallowingbyadultmalesmaycontributeto

greatermortalityamongadultmalesthanadultfemalesduring

outbreaksofthedisease(Gateset al.1995).Bisonhavedied

fallingintohotpoolsandbogs.Accidentaldrowningofwhole

herdsofbisonbyfallingthrough

thiniceinspringandfallhasbeen

reported(Roe1970;Gateset al.1991).

Oncebisonbreakthroughlakeorriver

ice,theyaregenerallyunabletohaul

themselvesoutandbecometrapped

(Carbynet al.1993).

Droughtsandseverewinters,aloneor

incombination,haveledtoepisodic

over-wintermortalityintheabsence

ofwolfpredationinplainsbisonof

theYNPcentralherd(Chevilleet al.

Figure 6.2 Age and sex class structure of

wood bison at Mackenzie Bison Sanctuary,

Northwest Territories, assuming an equal

sex ratio among calves and yearlings (Gates

et al. 1995).

1998;Greenet al.1997).Episodicdroughtsreducelategrowing

seasonforagequalityandincreasetheprobabilityofwildland

firesthatreducetheamountofwinterforageavailable(Frank

andMcNaughton1992).Simulationsindicatethatover-winter

survivalofYNPnorthernrangebisonismoststronglyinfluenced


table 6.4 Age-specific reproductive rates (%) of female plains and wood bison at select locations. Ages are female ages at time of birth of offspring (so,

a female reported as pregnant at one year by necropsy is shown as giving birth at two years, her second birthday).

Subspecies LocationAge

reference2 years >2 years >3 years

Plainsbison

WichitaMountains,OK 13 52 67 Halloran1968

WichitaMountains,OK 12 72 ShawandCarter1989

FortNiobrara,NB 83 Wolff1998

HenryMountains,UT 52 62 VanVurenandBray1986

AntelopeIsland,UT 46 Wolfeet al.1999

NationalBisonRange,MT 86 Rutberg1986

KonzaPrairie,KS 66–79 Towne1999

Badlands,SD 4 67 BergerandCunningham1994

WindCave,SD 5 80 Green1990,GreenandRothstein1991

Yellowstone–NorthernHerd,

WY/MT40 Kirkpatricket al.1996

Yellowstone–CentralHerd,WY 52 Kirkpatricket al.1996

Yellowstone–mixed,WY 73 PacandFrey1991

Yellowstone–mixed,WY 79 MeyerandMeagher1995

Woodbison

WoodBuffalo–HaysCamp,NWT 4 53 Fuller1962

WoodBuffalo–LakeClaire,AB 12 76 Fuller1962

WoodBuffalo,NWTandAB

76*

70**JolyandMessier2004

WoodBuffalo,NWTandAB 43 Carbynet al.1993

MackenzieBisonSanctuary,NWT 70 GatesandLarter1990

*nodisease**infectedwithbrucellosisandbovinetuberculosis


bywinterseverityandtheareaofwildlandfires(Turneret al.

1994;Wallaceet al.2004).

Survivalofcalvestosixmonthsismorethan90%inplainsbison

herdsinprotectedareas,orthosethatareonlylightedhuntedin

theabsenceofpredatorsanddiseases(Table6.5).Thesurvival

rateforthefirstsixmonthsoflifeinthepresenceofwolvesat

WBNPwas47%(Table6.5;BradleyandWilmshurst2005).At

theSRLsurvivalratesforthefirstsixmonthsoflifeincreased

from6%to30%coincidentwithadeclineinwolfabundance

(Table6.3;Calef1984).Survivalthroughthefirstyearoflife,in

thepresenceofwolves,hasbeenestimatedat10%and41%for

bisonatWBNP(Table6.5;Carbynet al.1993;Fuller1962).Calf

survivalthroughthefirstyearoflifewas95%foranincreasing

SubspeciesLocation and

Years

Age

Comment reference<6 months

%

<1 year %

Adult %

Plainsbison

HenryMountains,UT 93 96 VanVurenandBray1986

Badlands,SD 98 BergerandCunningham1994

Jackson,WY 95 Femalesonly.Increasingpopulation. USFWSandNPS2007

WindCave,SD 99 1of153calvesborndied GreenandRothstein1991

Woodbison

WoodBuffalo,NWTandAB <10 Fuller1962

WoodBuffalo,NWTandAB 41 Calculatedfromlifetable Carbynet al.1993


92(mm)

94(ff)

Oneornodiseases.AverageofWilsonet al.1995andJolyand

Messier2001

BradleyandWilmshurst2005


<85(mm)

<87(ff)Bothdiseases

JolyandMessier2001,Wilsonet al.1995inBradleyand

Wilmshurst2005

WoodBuffalo,NWTandAB 47 33 BradleyandWilmshurst2005

MackenzieBisonSanctuary,NWT 95 Increasingpopulation.

Fewwolves. Calef1984

MackenzieBisonSanctuary,NWT 75

Increasingpopulation.Ad.Femalerange67–100:

Adultmalerange67–100Larteret al.2000

SlaveRiverLowlands,NWT

1974–19766 Calef1976inCalef1984

SlaveRiverLowlands,NWT

1976–197830 Followingwolfdecline VanCamp1978inCalef1984

table 6.5 Age-specific survival rates (%) of plains and wood bison at select locations (mm = male; ff = females).

herdattheMBS,whenwolfabundancewaslow(Table6.5;Calef

1984).Therearehighlyvariableestimatesonsurvivalpatternsin

thefirstyearoflife(Table6.5).

Adultsurvivalratesindisease-free,protected,orlightlyhunted,

populationsofplainsbisonaremorethan95%forsexes

combinedorfemalesonly(Table6.5).Survivalratesforboth

sexesinincreasingpopulationshaveaveraged75%forwood

bisonattheMBS,and95%fortheJacksonplainsbisonherd

(Table6.5;Larteret al.2000;USFWS-NPS2007).AtWBNP,

bisoninfectedwithbothbrucellosisandBTBexperience

lowersurvivalratesthandothoseinfectedwithonlyoneof

thetwodiseases,ornotinfectedatall(Table6.5;Bradleyand

Wilmshurst2005;JolyandMessier2001;2004;2005).


6.3.4 population growth rates

Therateofincreaseofapopulationisinfluencedbysexratio

andagestructure,forageandhabitatavailabilityandquality,

immigrationandemigrationcombinedwithreproductiveand

mortalityrates.Thehighestratesofincreaseoccurincaptive

bisonherds,intheabsenceofpredators,wherethesexratiois

skewedtowardsreproductiveagefemales,somesupplemental

feedingoccurs,andmost,orall,ofthepopulationisrounded

upannuallyand“surplus”bisonremoved.TheTallgrassPrairie

Preserve(Oklahoma)populationattainedarateofincrease

ofabout50%undersuchconditions(R.Hamilton,personal

communication).

Themaximumexponentialrateofincrease(rm)istherateat

whichapopulationwithastableagestructurewillgrowwhen

resourcesarenotlimiting(Caughley1977).Theobserved

Figure 6.3 Growth of the National Bison Range plains

bison population between 1909 and 1922 (14 years)

starting with 37 bison (upper panel), and the northern

Yellowstone National Park population between 1902 and

1915 (14 years) starting with 21 bison (lower panel).

exponentialrateofpopulationgrowthovertime(r̂ )may

approximatermforpopulationsintroducedintoareaswhere

resourcesareabundant(Caughley1977).Theobservedrate

ofgrowthmaybeexpectedtodeviatefromrmovertimeas

apopulationincreases,andpercapitaresourcesbecome

limiting.Thelengthoftimeforapopulationtodoubleinsize

maybecalculatedas(naturallog(ln)of2)/r̂ (Johnson1994).

Plainsbisonre-introducedtotheNBRin1909werepermitted

toincreasewithoutmanagementinterventionfor14years

(Roelle1977inFredin1984).Theobservedexponentialrate

ofgrowthofthepopulationinthisperiod,withastarting

populationof37,wasr̂ =0.2053(Figure6.3).Thepopulation

grewatarateof20.5%eachyear,withadoublingtimeof3.4

years,or,giventhebirth-pulsecharacteristicofbison,itwould

realisticallydoubleeveryfouryears.ThenorthernYellowstone

plainsbisonherdwasintensivelymanagedintheearly20th

century,withsupplementalfeedprovidedin

winter(Meagher1973).Numbersincreasedfrom

21in1902to239in1915,afterwhichbison

wereremovedfromtheherd(Meagher1973).

Theobservedexponentialrateofincreaseforthe

northernYellowstoneherdforthis14-yearperiod

wasr̂ =0.1787.Thepopulationdoublingtimeat

thisrateofincreasewasfouryears.

Plainsbison,allowedtobecomefreeranging

intheJacksonValley,Wyomingin1969,

experiencedminimalmanagementintervention

until1980,whentheseanimalsbeganutilising

supplementalforageintendedforelkatthe

NER(USFWS-NPS2007).Limitednumbersof

plainsbisonwerekilledbyagencypersonnel

orlicensedhuntersbetween1980and2002

(USFWS-NPS2007).Plainsbisonnumbershave

beenestimatedannuallybystaffofGTNP;S.

Cain;personalcommunication).Theobserved

exponentialpopulationgrowthrateforthe33-year

periodfrom1969to2002was0.129(Figure6.4).

Theobservedexponentialrateofincreaseforthe

Jacksonherdforthe14-yearperiodfrom1980to

1993,withastartingpopulationsizeof37,wasr̂

=0.1197.Attheseratesofincrease,apopulation

woulddoubleeverysixyears.


Thehighestrateofincreasereportedfora

bisonpopulationundernaturalconditions

wasfortheMackenziepopulationinthe

NorthwestTerritories.Itincreasedata

maximumexponentialrateof0.26,and

averagedanannualexponentialrateof0.21

duringthefirstthreedecadesfollowingits

establishment(Calef1984;GatesandLarter

1990).

Figure 6.4 Growth of the Jackson Valley plains

bison population in Wyoming between 1969 and

2007 (39 years) starting with 9 bison (upper panel)

and between 1980 and 1993 (14 years) starting

with 37 bison (lower panel).


Chapter7 NumericalandGeographicStatusLead Authors: C. Cormack Gates and Kevin Ellison

Contributors: Curtis H. Freese, Keith Aune, and Delaney P. Boyd

7.1 Introduction

The“GreatContraction”,atermusedbyFlores(1996)to

describethedestructionofbisoninNorthAmerica,hasbeen

chronicledbynumerousauthors(Dary1974;Isenberg2000;

Reynoldset al.;2003;Roe1970)andwassummarisedin

Chapter2ofthisdocument.Fewerthan300woodbisonand

perhapsonly200plainsbisonremainedattheturnofthe19th

Century.Thenumericalrecoveryofplainsbisonbeganwith

theeffortsofprivatecitizensintheU.S.andCanadatosave

afewremaininganimals(Freeseet al.2007).Governments

laterbecameinvolvedintheconservationofplainsandwood

bison.ProtectivelegislationwasimplementedfirstinCanadain

1877(Gateset al.2001).Thefirstlegislationprovidingspecific

protectionforbisonintheU.S.wastheNationalParkProtective

Act(LaceyAct)signedon7May1894byPresidentCleveland

(BoydandGates2006).Itimposedajailsentenceandfinefor

anyonefoundguiltyofkillinggameinYellowstoneNationalPark,

therangeofthelastfree-rangingplainsbison.

Between1900and1970,modestprogresswasmade,increasing

thenumberandpopulationsofbison,largelyinpublicherds.

Theninthemid-1980s,thecommercialbisonindustrybeganto

prosper(Freeseet al.2007;Reneckeret al.1989);thenumberof

bisoninNorthAmericaincreasedrapidlytomorethan430,000,

thevastmajorityofwhichareunderprivateownership(Boydand

Gates2006;Freeseet al.2007).However,numericalprogress

alonecannotbeequatedwiththesecurityofbisonasawildlife

species.Conditionsunderwhichprivatelyownedbisonare

raisedarecommonlymotivatedbymarketobjectivesandthere

arenoregulationsorgovernment-supportedguidelinesrequiring

privateownerstocontributetobisonconservation.Domestic

bison(thoseraisedforcaptivecommercialpropagation)maybe

subjecttosmallpopulationeffects,selectionfordomestication

andmarkettraitsincludingdocility,growthperformance,

conformationandcarcasscomposition,andintentionalor

unmanagedintrogressionofcattlegenes(Freeseet al.2007).

Althoughsomeprivateownersexercisetheirlegalpropertyright

tomanagebisonforconservationofthespeciesand/orfortheir

ecologicalrole,theconservationpracticesofsuchownersare

amatterofpersonalchoice,withnoguaranteeofpersisting

beyondtheowner’sinterestintheherd.Currentlythereare

nowell-developedregulatoryormarket-basedincentivesfor

managingprivatecommercialherdsforspeciesconservation

(e.g.,independentconservationmanagementcertification).

Unlesseffectiveprivate-sectorincentivesaredeveloped,bison

populationsmanagedinthepublicinterestaswildliferepresent

themostsecureopportunityfortheirconservation,adaptationin

theevolutionarysense,andviabilityofbisonasanecologically

interactivespeciesinthelongterm.

SomeNorthAmericanaboriginalcommunitiesandindividuals

alsoownbisonherds.Aswithotherprivatebisonpopulations,

themanagementofNative-ownedbisonisnotnecessarily

consistentwithconservationpolicies.Managementpractices

varyfromintensivemanagementforcommercialproductionto

semifree-rangingherdshuntedforsubsistenceandretentionof

culture.

Itwasbeyondthescopeofthisstatusreporttoevaluate

themanagementofindividualprivatelyownedherdsfor

theirconservationvalue,whetherownedbyaboriginalor

non-aboriginalpeople.TheIUCNBisonSpecialistGroup

acknowledgestheimportantopportunitythatAboriginal

Governments,theIntertribalBisonCooperative,andtheNative

AmericanFishandWildlifeSocietyhavetodevelopguidelines

forenhancingtheconservationvalueofherdsmanagedby

aboriginalpeoples.Similarly,thecommercialindustrycouldplay

arolebyprovidingstandardsandguidelinesanddeveloping

incentive-basedprogrammes,suchasindependentformal

certification,forconservationmanagement.

Contemporaryconservationisfocussedonensuringlong-

termpersistenceandmaintainingthepotentialforecological

adaptationthroughtheeffectsofnaturalselectionoperating

inviablepopulationsinthewild(Soulé1987;IUCN2003;

SecretariatoftheConventiononBiologicalDiversity1992).

Viabilityrelatestothecapacityofapopulationtomaintain

itselfwithoutsignificantdemographicorgeneticmanipulation

bypeoplefortheforeseeablefuture(Soulé1987).Inwild

populations,limitingfactors,suchaspredation,resource

limitationandmatecompetition,contributetomaintainingthe

wildcharacter,geneticdiversity,andheritabletraitsthatenable

aspeciestoadapttoandsurviveinanaturalsettingwithout

humaninterference(Knowleset al.1998).Therefore,viablewild

populations,subjecttothefullrangeofnaturallimitingfactors,

areofpre-eminentimportancetothelong-termconservation,

securityandcontinuedevolutionofbisonasawildlifespecies.

Weconsiderthethreeconservationbiologyprinciplesproposed

byShafferandStein(2000),resiliency,representation,and

redundancy,toberelevantforevaluatingthegeographicand


numericalstatusofbison.Beyondviability,resiliencyrefers

totheneedtopreserveindividualpopulationslargeenough

tohaveahighprobabilityofpersistingforextendedperiods

inthepresenceofminimalmanagement,andwhichpreserve

geneticdiversityandthepotentialforadaptationtochanging

conditions(minimumof1,000bison;GrossandWang2005).

Representationreflectstheneedtopreservepopulationsof

aspeciesacrossthefullestarrayofenvironmentsinwhichit

occurredoriginally.Redundancyreferstotheneedtopreservea

sufficientnumberoflargepopulationstosafeguardagainstlocal

catastrophes.

Here,weprovideasummaryofthestatusofwoodbisonand

plainsbisonpopulationsmanagedbynationalorstate/provincial

publicgovernmentsandnon-governmentalorganisationswhose

primarymissionisnatureconservation.Forsimplicity,these

populationsarereferredtoas“conservationherds”.Information

onthenumberofherdsandbisonundercaptivecommercial

propagationisalsoincluded.Displayherdsinzooswerenot

enumerated.Thefollowingsevencriteriawereconsideredfor

reviewingthestatusofconservationherds:numericalstatus;

geographicstatus;populationsizeclassdistribution;opportunity

formatecompetitionamongmaturemales;presenceof

wolves;thepresenceorabsenceofdiseasesthatcouldaffect

conservationstatus(seechapter5);andpresence,orlikely

presence,ofcattlegenesbasedonanalysisorstockinghistory.

7.2 Numerical Status

Numericalstatusreferstothenumberofbisonandnumber

ofpopulationsinNorthAmericainconservationherds.Where

possible,thereportednumberofbisonineachconservation

herdwasverifiedwithherdmanagersin2008,butthenumbers

reportedheremaydifferfromtheactualnumbersofanimals

presentbecausenotallherdsweresurveyedrecently,census

techniquesmaynotaccountforeveryanimal,herdsarenot

alwaysmanagedtoachieveaconsistenttargetnumber,and

herdsizeandproductivityvaryannually.

Figure 7.1 Locations and size classes of bison conservation herds in North America. Historic ranges of wood and plains bison were based on

Stephenson et al. (2001) and Sanderson et al. (2008), respectively.


Sixty-twoplainsbisonand11woodbisonconservationherds

wereenumerated(Figure7.1andAppendixA).Although

thenumberofplainsbisonconservationherdshassteadily

increasedovertime,thenumberofindividualsinconservation

herdshaschangedlittlesince1930(Freeseet al.2007).In2008,

weestimatedtherewere20,504plainsbisonand10,871wood

bisoninconservationherds.Amongplainsbisontherewere

9,227breedingagefemales(twoyearsoldandolder),4,121

maturemales(sevenyearsoldandolder)and1,230subadult

males(fourtosixyearsold).Amongwoodbisontherewere

4,892breedingagefemales,2,609maturemalesand652

subadultmales.

Sinceconservationeffortsbeganintheearly1900s,wood

bisonnumbershavefluctuatedindependentlyofthenumber

ofconservationherds(Figure7.2).Peakabundanceoccurred

fromthe1940stoearly1970sfollowingtheintroductionofmore

than6,000plainsbisonintoWoodBuffaloNationalPark(WBNP)

inthelate1920s.ThenumberofbisonintheGreaterWood

BuffaloNationalParkareadeclinedafter1971whenpredator

managementceased(Carbynet al.1993).Thenumberofwood

bisonconservationherdshasincreasedto11.However,there

arestillmorebisonintheWBNPandSnakeRiverLowlands

(SRL)metapopulation(6,141animals),whichisinfectedwith

bovinetuberculosis(BTB)andbrucellosis,thaninthenine

disease-freereintroducedpopulations(4,730animals).

Thenumberofbisonundercommercialpropagationhas

outnumberedthoseinconservationherdssinceabout1970

(Freeseet al.2007).In2006,therewere195,728bisonon1,898

farmsreportingintheCanadianNationalCensus(Statistics

Canada,www.statcan.gc.ca/daily-quotidien/080125/t080125b-

eng.htm,accessed4December2008).TheU.S.Departmentof

Agriculture’s2007CensusofAgriculturereported198,234bison

on4,499farms(http://www.agcensus.usda.gov/,accessed

10February2008).Thus,basedonthesenumbers,thereare

nearly400,000privatelyownedbisononaround6,400farmsin

CanadaandtheU.S.

7.3 Geographic Status

Theoriginalrangeofbisonextendedfromlowlandmeadows

ininteriorAlaskatodesertgrasslandsinMexico,andincluded

areasasfareastasNewYorkandasfarwestasCalifornia

(Listet al.2007;Reynoldset al.2003).Theoriginalrangeof

AmericanbisonspannedanareaestimatedbySandersonet al.

(2008)tobe9.4millionkm2,andencompassed22majorhabitat

types(derivedbySandersonet al.2008bycombiningsome

oftheeco-regionclassesmappedbyRickettset al.1999).In

assessinggeographicstatusofbisoninconservationherds,

weconsideredthreecriteria:representationofsubspecies

Figure 7.2 Numbers of herds and individual plains bison (upper panel)

and wood bison (lower panel) in North America, 1890-2008. Sources

for wood bison data: Novakowski 1978; Wood Bison Recovery Team

1987; Reynolds and Hawley 1987; Van Camp 1989; Larter et al.

2000; Gates et al. 2001; www.nwtwildlife.com/NWTwildlife/bison/

woodbuffalopark.htm accessed 15 January 2009, and 2008/2009 data

from agencies. Plains bison data follow Freese et al. 2007 and current

status data from agencies.

plate 7.1 Wood bison near the northern extent of their range in the

Yukon, Canada. Photo: Tom Jung.


populationswithintheiroriginalrangeandinmajorhabitattypes,

andthegeographicareaoccupiedby,orpotentiallyavailableto,

individualconservationherds.

Representation within and outside their original range:A

displacedpopulationofasubspecieswithintheoriginalrange

ofanothersubspeciesmayoccupyhabitatotherwiseavailable

fortherecoveryandconservationoftheindigenousform.

Eighty-sevenpercentof62plainsbisonconservationherdswere

locatedwithintheoriginalrangeofplainsbison(Figures7.1and

7.3).EightplainsbisonherdsresidinginCalifornia,northern

BritishColumbia,andAlaskaweredistinctlyoutsideplainsbison

originalrange.ThoseinAlaskaandnorthernBritishColumbia

occurintheoriginalrangeofwoodbison.Nineof11wood

bisonherdswerewithinoriginalrange.Thetwowoodbison

conservationpopulationsoutsidetheoriginalrangeinclude

onefree-rangingherdintheInter-LakeregionofManitoba

(originallytherangeofplainsbison)andafencedherdincentral

Alberta.ThelatterwoodbisonpopulationisCanada’snational

conservationbreedingherdatElkIslandNationalPark,which

alsosupportsaseparateherdofplainsbison.

Representation in major habitat types: Eighteenmajorhabitat

typesoccurwithintheoriginalrangeofplainsbison(Figure

7.4).Atleastoneconservationherdisrepresentedin14(78%)

ofthemand10(56%)majorhabitattypesholdtwoormore

conservationherds.Atleastonewoodbisonconservationherd

isrepresentedinfour

(57%)ofsevenmajor

habitattypesintheir

originalrange,and

fourhabitattypeshave

twoormoreherds.

Withtheexceptionof

WBNPandtheadjacent

SRLbisonherds,

geographicseparation

ormanagementofother

populationsprecludes

inter-population

movements.

Available area: Thearea

availableforaherd

representsthepotential

forsupportingalarge

resilientpopulation

andopportunitiesfor

bisontobehaveasa

“landscapespecies”,

interactingwithspatially

variableresourcesandavarietyofothernativespecies.On

smallpastures,bisonmaybeunabletosegregateintosocial

units(maturebulls,maternalandnon-maternalherds)ortomove

inrelationtoresourcedepletionandabundancegradients.In

addition,thelargertheareaavailable,thegreaterthenumberof

bisonthatcanbesupportedsustainably.Landscapeareaisan

importantfactorinconsideringtheconservationstatusofbison.

Theareaofrangeavailabletobisonconservationherds

wasclassifiedintofourcategories(metricconversionsare

approximate):smallareas(lessthan20km2;5,000acres);

mediumareas(morethan20km2andlessthan200km2;more

plate 7.2 Plains bison near the southern extent of their range near Janos, Chihuahua, Mexico. Photo: Rurik List.

Figure 7.3 Numbers of plains and wood bison populations within and

outside their original range.


than5,000acresandlessthan50,000acres);largeareas(more

than200km2andlessthan2,000km2;morethan5,000acres

andlessthan500,000acres);andverylargeareas(morethan

2,000km2;morethan500,000acres).Abouthalfofplainsbison

conservationherdsoccuronsmallrangesandonly10%of

herdsareonverylargeranges(Figure7.5).Incontrast,37%of

woodbisonherdsoccuronverylargerangesandnoneoccuron

smallranges.

7.4 population Size Distribution

Usingasimulationmodel,GrossandWang(2005)demonstrated

thataminimumpopulationofabout400animalswasneeded

toretain90%ofselectivelyneutralvariationwitha90%

probabilityfor200years.Allelicdiversitywasmoresensitive

tomanagementtreatmentsthanaverageheterozygosity.On

average,ahighproportionofalleleswithaninitialfrequency

oflessthan0.05werelostwhenherdshadfewerthan400

animals.Differencesingenerationtimeaccountedforabout

Figure 7.4 Representation of plains and wood bison conservation herds in original ranges and major habitat types in North America. Habitat types were

based on Sanderson et al. (2008).

75%ofvariationinretainedheterozygosityforpopulationsof

200-800bison.Aspopulationsizeapproached1,000,theeffects

ofpopulationmanagementongeneticvariationweresmall.

Therefore,weconsideredpopulationsexceeding1,000tobe

moreresilientthansmallerpopulations.

Sandersonet al.(2008)definedthefollowingsizeclassesfor

rankingcontributionsofbisonherdstoecologicalrestoration:

smallcontribution,fewerthan400animals;modestcontribution,

400-1,000animals;largecontribution,1,000-5,000animals;

exceptionalcontribution,morethan5,000animals.The

frequencydistributionofconservationpopulationsize(Figures

7.1and7.6)illustratesthatsmallpopulations(fewerthan400

animals)arethemostcommonpopulationsizeclassamong

bothplainsandwoodbison(74%and55%,respectively).Five

plainsbisonandthreewoodbisonherdsexceed1,000animals.

Onlytwopopulationshaveencompassed5,000animalswithin

theirrecentrangeofsizevariability(GreaterYellowstoneArea

andGreaterWoodBuffaloParkarea).


7.5 Mate Competition

Thesexandagestructureofsmallpopulationsaresometimes

manipulatedtoreducetheriskofescapes,removeaggressive

animals,compensateforunequalmatingbymales,alter

fecundity,ortoincreasetherateofpopulationincrease.A

commonpracticeforbothsmallconservationherdsand

commercialherdsistoremovemalesbeforetheybecome

morphologicallyandbehaviourallymature(sixtosevenyears

oldandolder),whentheymaybecomedangeroustopeopleor

otheranimalsandproperty(e.g.,fences).Furthermore,thesex

ratiomaybemanipulatedtomaintainonlysufficientyoungmales

toensurefecundity(e.g.10males:100females).Incontrast,in

non-manipulatedwildherdsthematuremale:femaleratiocan

exceed50:100(Gateset al.1995)andmatecompetitionamong

malesisassured.

Thebisonisapolygynousspeciesinwhichmaturemales(six

orsevenyearsold)competevigorouslyformatingopportunities

(Komerset al.1992).Intheabsenceofmaturemales,juvenile

andsubadultmalesarecapableofbreedingsuccessfully,but

thereislittlecompetitionamong

themformatingopportunities

(Komerset al.1994a,b).We

consideredthatthepresenceof

twoormorematuremalesindicates

thepotentialformatecompetition.

Sixteenpercentofplainsbison

conservationherdsdidnotcontain

maturemales.Incontrast,twoor

morematuremalesweremaintained

inallwoodbisonconservation

herds,thusprovidingopportunityfor

matecompetition.

7.6 presence of Wolves

Keyspecies,suchasbison,have

adisproportionateinfluence

onthepatternsofoccurrence,distribution,anddensityof

otherspecies.Wherepresent,bisoninfluencethestructure,

composition,andstabilityofplant(Campbellet al.1994;Knapp

et al.1999)andanimalcommunities(Bogan1997;Roe1970;

Truettet al.2001).Grazerslikebisonalsoenhancemineral

availabilityandnutrientcyclingthroughfaecesandurine

deposition,andcarcassdecomposition(AugustineandFrank

2001;Towne2000;WallisDeVrieset al.1998).Thepresenceof

wolves,theonlyeffectivepredatorofbison(asidefromhumans),

isanindicatorthatthemaximumnumberofinteractionsis

possiblebetweenbisonandotherspeciesinanecosystem.If

wolvesarepresentweassumedthatallothernaturallimiting

factorswouldlikelybepresentintheecosystem.Wolvesare

associatedwithonly10%ofplainsbisonconservationherds(6

of62)incontrastto82%ofwoodbisonherds(9of11).

7.7 presence of reportable Diseases

Althoughdiseasesmaylimitbisonpopulationgrowthand

productivitytheyareunlikelytocauseextirpation.However,

thepresenceofdiseasesreportable

underfederalorstate/provincial

statutesmayleadtomanagement

interventionsthatimpact

conservation(Chapter5).The

typeofinterventionvarieswiththe

diseaseandjurisdiction(Chapter5).

Forexample,captiveconservation

herdsthattestpositiveforBTB

orbrucellosiswouldnormallybe

depopulated,whilelessserious

interventions(suchastheuseof

Figure 7.5 Area classes of ranges available for existing bison conservation herds.

Figure 7.6 Number of bison conservation

herds in four size classes.


Figure 7.7 Results of tests for cattle gene

introgression in conservation herds.

controlareas)maybeappliedforinfectedwildpopulationsin

largewildernessareas.Thepresenceofreportablediseasesmay

precludetranslocations.Managementinterventionsarepossible

tocontrolsomediseases(anthrax,BVD,JD).Reportable

diseaseswerepresentin5of62(8%)ofplainsbisonherdsand

3of11(27%)woodbisonherds.

7.8 Cattle Gene Introgression

Themolecularlegacyofhistorichybridisationbetweenbison

andcattleisaseriouschallengeforbisonconservation

today(HalbertandDerr2007).Forcedhybridisationhas

leftalegacyofcattleDNAthatis

widespreadamongcontemporary

bisonpopulations(Chapter4).The

implicationsforbisonconservationare

justbeginningtobeunderstoodand

appropriateinterventionsconsidered.

Availabletechnologyallowstesting

ofpopulationsforthepresenceof

markersforthecattlegenomeand

mitochondrialDNA(MtDNA),butall

conservationsherdshavenotyet

beentested(Figure7.7).Amongthose

tested,introgressionwasdemonstrated

insevenplainsbisonconservation

herds,butnoneofeightwoodbison

herds.Basedonstockingsources,

introgressionislikelyin17plainsbison

herdsandnowoodbisonherds.

plate 7.3 Male plains bison sparring. Photo: Dwight Lutsey.

7.9 Conclusions

Originally,theAmericanbisonrangedfrom

northernMexicotoAlaska.Plainsbison

occurredfromNorthernMexicotocentral

Albertaandwoodbisonoccurredfromcentral

AlbertatoAlaska.Thecontinentalpopulation

underwentadramaticdeclineduringthe

19thcentury,causedbyoverhunting,buthas

sincepartiallyrecovered.Approximately93%

ofthecontinentalpopulationismanagedfor

privatecommercialpropagation;veryfewof

theseherdsaremanagedprimarilyforspecies

conservation,andnonearemanagedinthe

publicinterestforconservation.Bisoncurrently

occupylessthan1%oftheiroriginalrange,

andconservationherdsoccupyasmallfraction

ofthat1%.Thenumberofconservationherds

hasincreasedsince1930,butthenumbersof

individualsinpopulationsmanagedprimarilyforconservation

haschangedlittlesincethen.Thereare62plainsbisonand

11woodbisonconservationherds(managedforconservation

inthepublicinterest).Conservationherdsaretypicallysmall

(fewerthan400animals)andpopulationsarewidelydispersed

withonlyonesituationthatprovidesgeographicconditions

fornaturalmovementsbetweenpopulationunits.Thecurrent

numberoflargepopulationsisfiveplainsbisonandthreewood

bisonherds.Theestimatednumberofbreedingfemalesin

conservationpopulationsis9,227plainsbisonand4,892wood

bison.Theircurrentrangeisrestrictedbylanduseandwildlife


managementpoliciesinthesouth,andbywildlifeandreportable

diseasemanagementpoliciesinthenorth.

AmongNorthAmericannations,thespeciesismostlimitedin

Mexico,whereaninternationaltrans-boundarywildherdrecently

occurred,butisnowlimitedbymanagementtoaprivateranch

inNewMexico(U.S.),wheretheyareclassifiedaslivestock.

Severalincreasingherdsornewprojects(AmericanPrairie

Reserve,Montana;BrokenKettleGrasslandReserve,Iowa;San

LuisValley,Colorado;PANP,Saskatchewan,Canada;Janos

Grassland,Chihuahua,MexicoandadjacentNewMexico;Yukon

Flats,MintoFlats,andlowerInnokoRiverareasinAlaska)have

thepotentialtodevelopresilientpopulationsonlargelandscapes

therebyadvancingthelong-termsecurityofbisonaswildlife.

TheAmericanbisonnearlyqualifiesforlistingasVulnerable

C2a(i)underIUCNcriteriaandiscurrentlylistedasNear

ThreatenedontheIUCNRedListinlightofitsdependence

onongoingconservationprogrammesandaverylimited

numberoflargeresilientpopulationsinthewild(Gatesand

Aune2008).Futureprogressontheconservationandrecovery

oftheAmericanbisonwilldependonsignificantchangesin

itslegalstatusandmanagementaswildlifebyfederaland

state/provincialagencies,harmonisationofpoliciesand

activitiesamongagenciesatmultiplelevels,cooperationwith

landednon-profitorganisations,andpossiblythroughthe

creationofvoluntaryformalconservationstandardsforprivate

commercialherdsandpopulationsmanagedbyNativeAmerican

governments.


Chapter8 LegalStatus,PolicyIssuesandListings

Lead Authors: Keith Aune and Rick Wallen

Contributors: C. Cormack Gates, Kevin Ellison, Curtis H. Freese, and Rurik List

8.1 Introduction

ThebisonisaniconicNorthAmericanwildlifespeciesthat

symbolisesthewildandopenwesternprairieandborealforest

landscapesoftherecentpast.Althoughtheirdecline,and

subsequentrecovery,isfrequentlyrecountedinconservation

circles,theecologicalrecoveryof“wild”bisonwasneverreally

considered,andconsequentlytheirrestorationhasneverbeen

fullyaccomplished(Sandersonet al.2008).Mostplainsbisonin

NorthAmericaarefoundonfarmsandranches(about400,000)

whilerelativelyfew(about30,000)arelocatedonprovincial/

state,federal,andnon-profitconservationreserves(seeChapter

7).Fewpopulationsaredistributedbroadlyonnativelandscapes

insuitablehabitat,andmostdonotenjoyequallegalorpolicy

statuswhencomparedtootherimportantwildlifespeciessuch

aselk(Cervus elaphus),deer(Odocoileus spp.)orpronghorn

(Antilocapra americana).Woodbisonaremanagedmore

commonlyaswildlifewithintheirhistoricrangethanplainsbison,

butsufferfromfragmenteddistributionanddiseaseissuesthat

complicatetheirmanagement.

Thepurposeofthischapteristoevaluatethehistoricandcurrent

legalstatusofbisoninNorthAmericaandidentifylegaland

policyobstaclesrelevanttoconservationeffortsforthisspecies.

Duetoahistoricalparadigmthatviewedbisonaslivestock,

andpastconservationmeasuresthattreatedtheminamanner

similartolivestock,bisonhavenotachievedalegalorpolicy

statuscommensuratewithapremierkeystoneherbivorenativeto

prairieecosystems.Duringthegreatrestorationperiodofwildlife

management,bisonwereroutinelyclassifiedandmanagedby

state/provincialandfederalagenciesacrossNorthAmericaasa

formoflivestock,whileotherwildlifewereclassedandmanaged

asfree-roamingwildanimalsconsistentwithwildlandscapes.

8.2 history of protection and Conservation

8.2.1 Early legal and policy efforts by governments to protect plains and wood bison

8.2.1.1 Early policy development in the United States

Outcriesduringthe19thCenturytohaltthedestructionof

bisonintheU.S.werelargelyignored.In1820,MajorStephens

expressedconcernabouttheexcessivekillingofplainsbison

andadvocatedalawtopreventwantonslaughter(Dary1989).

In1843,JohnJ.Audubonissuedwarningsagainsttheslaughter

ofbison(Dary1989).Despitetheirpleas,noconservation

policyorprotectivelegislationwasenactedforseveralmore

decades.Numerousbillstoprotectplainsbisonwereintroduced

bymembersoftheU.S.Congressbetween1871and1876;

nonewaspassedintolaw.Althoughtherewerenosuccessful

federalinterventionstohalttheslaughter,severalstatesenacted

legislationontheirown.Between1864and1872,thestates

ofIdaho,Wyoming,andMontanaimplementedstatutesto

reducethekillingofgame,includingbison.Althoughtheselaws

reflecteddeepconcernfortheconservationofwildlife,they

werelargelyineffectiveowingtolimitedenforcement.In1872,

PresidentUlyssesS.GrantestablishedYellowstoneNational

Parktoprotectallresources,includingbison,withinitsborders.

The“ActtoProtecttheBirdsandAnimalsinYellowstone

NationalParkandtoPunishCrimesinSaidPark”wassignedby

PresidentGroverClevelandinMay1894,providingthemeans

necessarytohalttheextirpationofthelastfree-rangingplains

bisonpopulationinNorthAmerica(Gateset al.2005).Despite

theseefforts,by1902,fewerthan25free-rangingplainsbison

remained,andthesewerelocatedintheremotePelicanValley

ofYellowstoneNationalPark(YNP)(Meagher1973).Afewwood

bisonmayhavepersistedintothe20thCenturyinAlaska,but

weresoonextirpated(Stephensonet al.2001).

8.2.1.2 Early policy development in Canada

InCanada,earlyconservationeffortsbeganin1877withthe

passingoftheBuffaloProtectionAct(Hewitt1921).In1883,

theOrdinancefortheProtectionofGamewaspassed,butit

wasnoteffectiveowingtopoorenforcement(Ogilvie1979).

PlainsbisonwereextirpatedfromthewildinCanadabythe

1880s(COSEWIC2004),butwoodbisonpersistedinasmall

populationinwhatisnowWoodBuffaloNationalPark(WBNP).

Thenationalparkssystemfirstbecameinvolvedinplainsbison

conservationin1897,whenthreeanimalswerepurchased

fromCharlesGoodnightinTexas.However,thefirstsignificant

contributionbytheGovernmentofCanadawasmadein1907

whenitpurchasedtheprivatelyownedPablo-Allardherdin

Montana.ThegovernmentofCanadaenactedtheUnorganised

TerritoriesGamePreservationActin1894,partlyasaresponse


tothedeclineofwoodbison.The1922OrdersinCouncilunder

theForestReservesandParksActestablishedWBNPinan

attempttosavewoodbisonfromextinction(Boyd2003;Gates

et al.2001a;2001b;Soper1941).

8.2.1.3 policy development in Mexico

Historically,bisonwerepresentinfivestatesinnorthernMexico,

butuntilrecentlyexistedinthewildonlyintheborderlands

betweentheJanosregionofChihuahuaandsouth-western

NewMexico(Listet al.2007).Mexicofirstincludedbisonon

itsred-listofendangeredspeciesin1994.Themostrecent

version(SEMARNAT2002)specificallylistsbisonintheJanos-

Hildagoherdas“endangeredwildlife”.Althoughthepopulation

isaffordedlegalprotectioninMexico,itisconsideredlivestock

whenitrangesintoNewMexico.Seesection8.5.5.3formore

detailsonthisherd.

BisonconservationinMexicohasprimarilybeenimplemented

throughfederalprogrammes;statushasnotyetbeen

establishedunderstatelegislation.TheNationalMinistryof

Environment(SEMARNAT2002)managedbisonformany

years.Recentlytheresponsibilityforpriorityspecies,including

bison,wastransferredtotheNationalCommissionofProtected

NaturalAreas.TheInstituteofEcologyoftheNational

UniversityofMexicoisadvocatinglegalprotectionoftheherd

inbothcountries,includingprotectionunderinternational

treatiesonmigratorywildlifespeciesbetweenMexicoand

theU.S.TheIUCNBisonSpecialistGroup(BSG)strongly

encouragesthisprotectiveactionandothereffortstorestore

plainsbisontotheChihuahuanDesertgrasslands.

8.2.2 plains bison conservation by the private sector

Privatesectorconservationeffortscanbecategorisedintotwo

non-exclusivegroups:1)privatecitizensinterestedprimarily

incommercialproductionofbisonandsecondarilyinbison

conservation;and2)privateconservationgroupsinterested

inconservingbisonaswildlife.Theformerdonottypically

haveformalconstitutionsmandatingconservation,whilethe

latterinstitutionstypicallydo.Legislation,regulations,rules,

andpoliciesaffectingcaptiveherdsownedbythesesectors

aresimilartodomesticlivestock,focusingontransport,trade,

export,import,animalhealth,anduseofpublicgrazinglands.

Notably,TurnerEnterpriseshasbeeninvolvedinthedevelopment

ofproductionherdson14largeranchesintheU.S.,thelargest

numberofplainsbisonownedandmanagedbyasingleowner.

Bisonaremanagedwithlowmanagementinputssimilarto

manypublicconservationherds.Notably,theCastleRockherd

onVermejoParkRanchinNewMexicoisderivedfromstock

translocatedduringthe1930sfromYNPandshowingnoevidence

ofcattlegeneintrogression.Althoughsomeprivatelyowned

herdsmaybevaluableforconservation,thereisnoprecedentfor

assessingtheirlong-termcontributiontoconservationofbison

aswildlife.Recently,theWildlifeConservationSocietydeveloped

anevaluationmatrixthathelpsidentifythekeycharacteristics

andpossiblemanagementadjustmentsthatwouldbenecessary

forprivatelyownedherdstocontributetobisonconservation

(Sandersonet al.2008).Thismatrixisstillevolvingandwas

recentlytestedamongasmallproducergrouptorefineand

improveitsapplication.Populationandgeneticmanagement

guidelinespresentedearlierinthisdocumentmayalsobeuseful

forguidingprivateproducerstowardmanagingtheirherdsin

supportofconservation.However,asystemforcertifyingherds

forconservationmanagementwouldberequiredtoensurethat

guidelinesarefollowed.

Severalnon-governmentalorganisations(NGO),particularly

TheNatureConservancy(TNC),theNatureConservancyof

Canada(NCC),AmericanPrairieFoundation(APF),andthe

WorldWildlifeFund(WWF)havebeenactiveindeveloping

conservationherds.Moreinformationontheirinitiativescanbe

foundinsection8.5.5.4.

8.2.3 Conservation efforts by tribes and First Nations

ManyNorthAmericanNativePeopleshavestrongcultural,

spiritual,andsymbolicrelationshipswithbison(Notzke1994;

Zontek2007).Sometribesbelievethatbecausetheanimals

oncesustainedtheirIndianwayoflife,they,inturn,musthelp

thebisontosustaintheirplaceontheearth.Theconservation

ofwildbisonincludestheintangiblevaluesthesetribesholdfor

bison.Valuesvarygreatlybetweentribes,andinsomecases,

evenbetweenmembersofthesametribe.Sometribalpeople

believethatthestatusofthebisonreflectsthetreatmentofNorth

AmericanIndians.Interestinpreservingtheculturalsignificance

ofbison,andinrestoringculturalconnectionstothespecies,can

beimportantincentivesforNativegovernmentsandcommunities

toparticipateinbisonconservation(Notzke1994;Zontek2007).

Sometribalbisonmanagersconsiderallbisonaswildanimals

regardlessofthesourceofstock,geneticintrogressionfrom

cattle,ordomesticationhistory.Thiscanbethebasisforconflict

withconservationbiologistswhoapplybiologicalcriteriawhen

evaluatingtheconservationmeritofaherd.Tribalgovernments

commonlyoperateunderchallengingcircumstances.Political

viewscanvarybetweensucceedingtribaladministrations,

creatingunstablepoliciesthatcanaffectbisonmanagementand

conservationpractices.NumerousNativeTribesownorinfluence

themanagementofasignificantlandbasethathasthepotential

tosustainlargebisonherds.However,therehasyettobea

systematicsurveyofthenumberofherdsorthedistributionof

bisonunderNativemanagement—ataskofsufficientmagnitude

andcomplexitytoexceedthescopeofthisreview.

Thepotentialfortribestoparticipateinbisonrestoration

isimprovingwiththedevelopmentoftribalgameandfish


administrations,andtheincreasingcapacitytoimplement

modernwildlifemanagementforwildlifeontriballands.Some

tribeshavedevelopedindependentbisonprojects.Others

havejoinedtheIntertribalBisonCooperative(ITBC)toobtain

guidanceandsupport.TheITBCwasformedin1990withthe

missiontorestorebisontoIndianNationsinamannerthatis

compatiblewiththeirspiritualandculturalbeliefsandpractices

(ITBCwebsite:http://www.itbcbison.com/).Incooperationwith

theNativeAmericanFishandWildlifeSociety,theITBCwas

abletosecureU.S.congressionalsupportforbisonrestoration

in1991.In1992,tribalrepresentativesmetandtheITBC

becameanofficiallyrecognisedtribalorganisationintheU.S.

TheITBCisanon-profit501(c)(3)organisationgovernedbya

BoardofDirectorscomprisedofatribalrepresentativefrom

eachmembertribe.Currentlythereare57membertribesthat

collectivelymanagemorethan15,000bison.TheroleofITBC

istoactasafacilitatorforeducationandtraining,developing

marketstrategies,coordinatingtransferofbisonfromfederal

ownershiptotriballands,andprovidingtechnicalassistance

totribalmemberstoencouragesoundmanagement.TheITBC

doesnothaveapresenceinCanada,noristhereanequivalent

organisationthere.Asummaryoftribalbisonconservation

initiativesisinsection8.5.5.5.

8.3 Important policy and regulatory Considerations

8.3.1 Legal status and listings of bison

8.3.1.1 International and global status

TheConventiononInternationalTradeinEndangeredSpeciesof

WildFaunaandFlora(CITES)isamultilateralagreementamong

nationstoensurethatinternationaltradeinspecimensofwild

animalsandplantsdoesnotthreatentheirsurvival.Species

listedinAppendixIarethosethreatenedwithextinction,while

specieslistedunderAppendixIImightsoonbeiftradeisnot

controlled.WoodbisonweretransferredfromCITESAppendix

ItoAppendixIIin1997basedonCanada’sabilitytosatisfy

the“precautionarymeasures”ofResolutionConf.9.24(Annex

4,paragraphsB.2.b.iandii).Althoughbisonareindemand

fortrade,theyaremanagedaccordingtotherequirementsof

ArticleIV.ItwasdeterminedthatCanadamaintainsappropriate

enforcementcontrolstopreventtheunauthorisedtakingofwild

bisonforcommercialfarming,andthatthetransfertoAppendix

IIwasconsistentwiththegoalsofthegovernment’srecovery

plan,andwouldnothamperprogresstowardtherecovery

ofwoodbisoninthewildwithintheiroriginalrange.Import

andexportofwoodbisonisregulatedunderpermitbyCITES

authoritieswithinmembernations.Plainsbisonarenotlisted

underCITES(http://www.cites.org/).

Americanbisonwererecentlylistedas“NearThreatened”inthe

IUCNRedListofThreatenedSpecies(GatesandAune2008).

AtaxonisNearThreatenedwhenithasbeenevaluatedagainst

thecriteria,butdoesnotqualifyforCriticallyEndangered,

EndangeredorVulnerablenow,butisclosetoqualifyingfor,oris

likelytoqualifyfor,athreatenedcategoryinthenearfuture.No

distinctionismadebetweenwoodandplainsbisonintheWorld

ConservationUnion(IUCN)RedBook.

NatureServeisanon-profitconservationorganisationand

internationalnetworkofbiologicalinventoriesknownasnatural

heritageprogrammesorconservationdatacentresoperatingin

all50U.S.states,Canada,LatinAmericaandtheCaribbean.

ItassignedanoverallconservationstatusranktoAmerican

bisonofG4(ApparentlySecure),meaningtheyareglobally

common(morethan100occurrences)generallywidespread,

butmayberareinpartsoftheirrange,andalthoughtheyare

secureintheirglobalrange,theremaybeaconcernfortheir

securityinthelongterm(NatureServe2006).Thewoodbisonis

rankedbyNatureServeasG4T2Q,where“T”referstoitbeing

anintraspecifictaxon(trinomial),“2”meansimperilled,and“Q”

referstoquestionabletaxonomy.Theplainsbisonisrankedas

G4TU,where“U”meanscurrentlyunrankableduetoalackof

informationorsubstantiallyconflictinginformationaboutstatus

ortrends.

8.3.1.2 Status in North America

ThewoodbisonwasdesignatedbyCanadaas“Endangered”

in1978.Owingtoprogressmadetowardsrecovery,itwas

downlistedto“Threatened”in1988.Thisdesignationwas

re-evaluatedandaffirmedinMay2000.Thewoodbisonis

protectedundertheCanadianSpeciesatRiskAct(2003),but

huntingisallowedinAlberta,theNorthwestTerritories,andthe

Yukon,subjecttoconservationstrategiesandmanagement

regulation.InJune1970,thewoodbisonwaslistedunderthe

U.S.EndangeredSpeciesAct(ESA)as“EndangeredinCanada”

toreflectitsstatusinCanadaatthattime.CanadaandtheU.S.

areundertakingeffortstoharmonisethenationallistingsofthis

subspecies(Gateset al.2001b).Arecentpetitiontodownlist

woodbisonfromendangeredtothreatenedintheU.S.was

submittedandthedecisionisunder90-dayreviewbytheU.S.

FishandWildlifeService(USFWS).

AlthoughplainsbisonarecurrentlynotlistedintheU.S.

orCanadaunderspeciesatriskofextinctionlegislation,

considerationofalistingstatusisbeingundertaken(COSEWIC

2004).In2004,COSEWICrecommendeddesignatingplains

bisonasThreatenedundertheSpeciesatRiskActinCanada

(WilsonandZittlau2004).Theproposedchangewaslisted

forcommentonthepublicregistryin2005.Criticismensued

fromcommercialbisonproducersconcernedwiththe

impactontheirindustryandinternationaltrade,andthere


wasalackofsupportbyAgricultureandAgri-FoodCanada

andtheprovincialgovernments.InJuly2006,TheFederal

MinisteroftheEnvironmentrecommendedthatplainsbison

notbelistedbecauseofpotentialeconomicimplicationsfor

theCanadianbisonindustry(http://canadagazette.gc.ca/

partII/2005/20050727/html/si72-e.html).

Thereareseveralpotentialcomplicationsthatwouldaccompany

theprocessoflistingplainsbisoninNorthAmerica.One

complicationregardingthelegalstatusofbisonistheissue

ofhybridisationwithcattle.Thereisconsiderableuncertainty

concerningif,andhow,endangeredspeciesstatusshouldbe

appliedtohybridsinCanadaandtheU.S.(BoydandGates

2006;CamptonandKaeding,2005).Hybridsareexemptfrom

theEndangeredSpeciesAct(ESA)whenpropagatedincaptivity,

andwhentheyaretheprogenyfromonelistedandonenon-

listedparent(O’BrienandMayr1991).Asecondcomplicationis

theconsiderationofcommercialbisonproductioninevaluating

thenumericalstatusofthisspecies.Athirdcomplicationisthe

legaldistinctionandstatusofwildandcaptivebisonshould

listingbeconsideredforthewildform(Boyd2003).

BisonoftenenjoyprotectedstatusinCanadianandU.S.

nationalparksasaresultofthelegalstatusofthehabitat.

TheCanadianNationalParksActprotectsbisonandtheir

habitatinnationalparks.InCanada,provincialandterritorial

governmentscanalsousethefederalWildlifeTradeActto

controlthemovementofbisonacrosstheirborders.Inthe

U.S.,enablinglegislationattachedtoeachnationalparkwhen

itwasestablished,typicallyprotectbisonaswildlifeunless

theyarenotconsiderednativetotheregion.Wheretheyare

notconsiderednativetoaregion,orareknowntobecattle

hybrids,nationalparksoftenconsidertheminvasiveandmay

considerremovaloreradication.

TheUnitedStatedForestService(USFS)classifiestheAmerican

Bisonas“NotSensitiveinRegion2andNotofConcern”by

itsSpeciesConservationProgramassessment(USDAForest

Service2009).Therationaleforthisclassificationisthat

populationsandhabitatsarecurrentlystableorincreasing.

ThisUSFSreviewsuggeststhatwhilethespeciesmaywarrant

restorationasanecologicalkeystonespecies,itdoesnot

warrantsensitivestatus.

ConservationandrestorationprogrammesforAmericanbison

areconfoundedbysocioeconomicchallengesresultingfrom

theconfusinglegalstatusforthisspecies.Thelegalstatusof

bisonrangesfromdomesticlivestocktowildlifeamongvarious

federal,state,andprovincialjurisdictionsacrossNorthAmerica

(Table8.1).Thelegalrecognitionofbisonaswildlifeisoften

impededbytheirhistoric,orinmanycasesdual,classification

asdomesticlivestock.Wheretheyhaveattainedtheirstatus

aswildlife,theyareroutinelymanagedwithinfencedpreserves

wheresome,ifnotall,naturalselectiveprocessesarecurtailed.

TenstatesintheU.S.,fourprovincesinCanada,andone

stateinMexicoclassifybisonaswildlife(Table8.1).Allother

statesandprovinceswithintheiroriginalrangedesignate

bisonsolelyasdomesticlivestock.Plainsbisonaredesignated

andmanagedaswildlifeinAlaska,Arizona,Utah,Montana,

Wyoming,BritishColumbia,Alberta,Saskatchewan,and

Chihuahua.Fourotherstatesconsiderbisonaswildlife,butdo

nothavefreerangingpopulationstomanage;Idaho(extreme

rarity),Missouri(extirpated),NewMexico(nolongeroccurring),

andTexas(extirpated).Plainsbisonarelistedandmanaged

aswildlife,butareconsideredextirpated,inAlbertaand

Manitoba.Wildbisonarepreserved,asapublictrustresource,

managedtoprotectnaturalselectionprocesses,andhunted

asfreeroamingwildlifeinAlaska,Arizona,Utah,Montana,

Wyoming,BritishColumbia,Alberta,andSaskatchewan.Wood

bisonaredesignatedandmanagedaswildlifeunderprovincial

statutesinManitoba,Alberta,BritishColumbia,Yukon,andthe

NorthwestTerritories.Woodbisonenjoyprotectedstatusinall

oftheseprovinces.Therearelegalrestrictionsonhuntingand

otheractivitiessuchascaptureandharassment.Subsistence

huntingbyaboriginalpeoplesisallowedunderstrictregulation

inNorthwestTerritoriesandYukon.

UnderMexicanlaw,wildlifebelongstothenation.However,

Mexicohasonlyrecentlydevelopedawildlifeconservationand

managementsystemthatentitlesalandownertoberegistered

intheprogramme(Unidades de Manejo y Aprovechamiento)

and toreceivethebenefitsofharvestandcommercialuse

ofwildlife.Thisprogrammehasdoubledthelandscape

availableforwildlifeprotectioninMexico.In1995,thefederal

governmentestablishedabureaumanagedbytheSecretary

oftheEnvironment.Withinthisorganisationisadepartment

fortheadministrationofwildlifeconservationprogrammes.In

2007,theconservationofthreatenedspeciesisbecomingthe

responsibilityoftheNationalSystemofProtectedNaturalAreas.

Thereisonlyalimitedstateorlocalwildlifemanagement

infrastructuretosupportfederalwildlifeconservationefforts

inMexico.Localcommunitiesareonlynowbeginningto

acceptandappreciatethevalueoffree-rangingwildlifeon

landscapesthattheyownandmanage.Untilabroaderlegal

andpolicyinfrastructureisestablished,federallawandpolicy

willcontinuetodirectwildlifemanagementconservation

inMexico.Federalpolicyisprimarilyaimedatdeveloping

partnershipswithlandownersandcooperativelyidentifies

conservationmeasuresacceptabletoindividuallandowners.

Inaddition,federalconservationlawandpolicydrivesthe

protectionoflandtoestablish“NaturalProtectedAreas”to

conservespeciesassociatedwiththoselandscapes.Public

interesthasincreasedindevelopingwildlifeprogrammesfor

economicandconservationpurposes.Interestinconservation

continued on page 73


table 8.1 Current legal status of plains and wood bison (Excluding portions of bison range where large landscapes are no longer available).

Country/State

province

Animal Classification protected and/or

Wildlife Status

Long term Conservation

Goal

Key Statutes or policies Affecting

Conservation

proposals for

restoration

Major Legislative and/or policy

ObstaclesWildlife Domestic

UnitedStates Yes Yes

PlainsbisonpetitionedunderESAbutdenied;

ManagedascaptivewildlifeonUSFWS

Refuges;USFSR-2classifiesbison

asnotsensitive;Managedaswildlife

(captiveorfree-ranging)inseveral

NationalParks.Recentpetitionto

downlistwoodbisonto“Threatened”is

under90-dayreview.

Nocomprehensivestrategy;Activity

limitedtoandfragmented

amongNGOs,veryfewstates,

NationalParksandUSFWSRefuge

System.

NEPA;NationalRefugeAct;EachNationalParkhasitsown

organiclegislation-Interpretedbyeach

ParkSuperintendent;WoodbisonarelistedasEndangeredunderESA;AnimalHealthProtectionAct(7#U.S.C.8301etseq.).

No

Absenceofstrategicplanning;Multiplejurisdictionsand

coordinationofagencies;

Managementincaptivityunderrefuge

policy;Diseasetransmissionto

livestock;Limitedinvolvementand

interestbymanystatewildlifeagencies;

Confusedregulatorystatusinmanystates.

Alaska Yes Yes

4introducedplainsbisonherdsare

“Wildlife”;OneplainsbisonherdonPopof

Island;Maintainahuntingprogramme

bypermitonly.

Long-termgoalsbeingestablished

forwildwoodbisoninStateWildlifeAction

Plan(SWAP)andreintroductionprogrammes;Management

planningforthe4introducedplains

bisonherds.

ESA10(J)statusforwoodbison-MintoFlatsintroduction;

Title16inAlaskastatestatutesdesignates

bisonaswildlife;DeltaBisonMgt.Plan;Wood

bisonConservationPlaninprogress;

Livestockmanagescaptivebisonunder

Title3inAlaskastatestatutes.Domestic

bisongovernedundersamerulesas

domesticcattle.

Yes;YukonandMintoFlatsWood

BisonRestorationisunderway.

Plainsbisonoutsidetheiroriginalrange;Aboriginalhunting

rights;USFWSinterpretationsoflegalstatusofwoodbison

underESA.

Arizona Yes Yes

Bisonarewildlife,specificallybiggame,andare

managedbyAGFDontwostatewildlifeareas(HouseRock

andRaymondRanch).

Yes,inSWAP.

Title12,R12-4-401GameandFish

CommissionRulesforLiveWildlife;R-12-4-406RestrictedLive

WildlifeSectionB9.dexemptsrestrictions

onpossessingcaptivebison(permitnot

requiredtopossess);A.R.S17-101A22defineswildlifeand

101Bdefinesbisonasagameanimal.

No

Arizonaisattheedgeofbisonoriginal

range;Currentstrategicplanlimitsconservationtotwoexistingpopulations;

HouseRockpopulationhybridisedwithcattle;Agricultureandforestryconflicts.

Colorado No Yes

Bisonareexemptfromtherequirementsof

wildlifecommissionregulations.Today,

captiveherdsaredesignatedaslivestock.

ConservationherdsexistintwoDenver

Cityparks,oneUSFWSRefugeandoneTNCpreserve.

Yes;OnTwoUSFWSRefuges

andoneTNCpreserve.

Chapter11,Section406-8Wildlife,Parks

andUnregulatedWildlife;Wildlife

CommissionRegulation#1103

exemptsbisonfromallwildlifecommission

regulations,asdomesticanimals

NoAgricultureand

forestryconflicts;Regulatorystatus.


Country/State

province


Wildlife Status


Goal


Conservation

proposals for

restoration



table 8.1 (continued)

Country/State

province


Wildlife Status


Goal


Conservation

proposals for

restoration



Idaho Yes Yes

IdentifiedasS1speciesinwildlife

commissionstatusreport.S1=criticallyimperilledspeciesathighriskbecauseof

extremerarity.

No

Livestockregulationschapter210section

01.a;NotmentionedinSWAP.

No

DiseaseStatusinYNP;Agricultureand


Illinois No Yes ConsideredextirpatedinIllinois. No

ManagedaslivestockunderstatestatuteChapter225part

650/1;NotmentionedinSWAP.

No

Agricultureandforestryconflicts;Smallparcelsofpublicorprivate

conservationland;Regulatorystatus.

Iowa No Yes

ConsideredextirpatedinIowa;

FoundonlyononesmallNational

WildlifeRefuge.

Yes;OnlyononeNationalWildlife

Refuge.

Managedaslivestockunderstateanimal

healthstatutes.Bisonstatutescombinedwiththoseofcattle;

Notmentionedanywhereinwildlife

regulationsorwildlifeconservation

strategies.

No



Kansas No Yes

Consideredextirpatedpriorto1900;Designateddomesticunderbeefrules;State

wildlifedepartmentmanagesbisonon

twosmallgameranges;TNChas

twoadditionalpreserves.

Yes;OnlyonTNCandstate

preserves

IdentifiedinSWAPasnotmeetingcriteria

forspeciesofgreatestconservationneed;Chapter60section4001inlivestock

regulations.

No



Louisiana No YesAllbisonareconsideredlivestock.

No

LouisianaCodeofregulations7:XXI.11705;No

mentionofbisoninSWAPorinwildlife

regulations.

No



Minnesota No Yes

Wildbisonareconsidered

extirpatedinMN;Foundonlyonacoupleofsmall

preserves

No

Minnesotastatutesforlivestock(17A.03);Bisonnotmentioned

inSWAP.

NoAgricultureand


Missouri Yes Yes


extirpatedinMissouri.

No

Identifiedasclass1wildlifeintitle3CodeofStateRegulations(CSR)10;Identifiedaslivestockintitle2

CSR30.

No


conservationland.



Country/State

province


Wildlife Status


Goal


Conservation

proposals for

restoration



Montana Yes Yes

Gameanimalstatus;Tier1

speciesinSWAP;Speciesinneed

ofmanagementinYNP;ManagedinhabitatsadjacenttoYNP.OnNBR;

OwnershipofNBRisindispute;American

PrairieReserve(APF).

YesinSWAP;NationalBison-

RefugePlan;Yellowstone

InteragencyBisonManagement

Plan;APFBisonReintroduction

andConservationPlan.

MontanaEnvironmentalPolicyAct(MontanaCode

Annotated(MCA)75-1-102);Legislativeauthoritytomanage

wildbisoninMontana(MCA81-2-120;MCA

87-2-130);SWAP;InteragencyBisonManagementPlan-

EIS,2000.

Yes;CharlesM.Russell

RefugePlan

Agricultureandforestryconflicts;

DiseasestatusinYNP.

Nebraska No Yes


extirpatedinthestate;Bisonaredefinedas

livestock;Foundonlyonseveralsmall

preserves.

Yes;OnlyonNationalWildlifeRefugeandTNC

preserves.

BisonfoundonlyintheDepartment

ofAgricultureregulations.Title23and54;Section54definestherequiredhealthregulationsfor

cattleandbison;

No;Possiblytribalefforts.

Agricultureandforestryconflicts.

NewMexico Yes Yes

Classifiedasgameanimalsin

1978;Identifiedinwildlifedatabaseas“apparentlynolongeroccurring”

butnotidentifiedasextirpatedorextinct;IncludedinSWAP.

Yes

Title17-2-3NewMexicoAdministrative

Code(NMSA)1978classifiesbison

asgameanimalsexceptwhereraised

incaptivityforcommercialpurposes;

Title19(Wildlife)chapter31describesthelegalweaponsfor

takingofbisonyettherearenohuntingregulationsforbison(19.31.10.16);Title19Chapter26describeslivestock(andnamesbison)asdomesticanimalsraisedona

ranch(19.26.2.7);Title21(agricultureandranching)hasmanyreferencestoward

managementofbison.

No

Agricultureandforestryconflicts;Lackofsuitable

habitat.

NorthDakota No Yes

Classedasnon-traditionallivestock;

BisonarefoundonlyinTheodore

RooseveltNationalParkandmanaged

asdomesticlivestockoutsidethe

NationalPark.

Yes;OnlyontwofederalandoneTNCpreserves.

Unabletofindanyreferencetobisonin

agricultureregulations(Title4)orwildlife

regulations(Title20).

NoAgricultureand



Country/State

province


Wildlife Status


Goal


Conservation

proposals for

restoration




Country/State

province


Wildlife Status


Goal


Conservation

proposals for

restoration



Nevada No Yes


extirpatedinNevadaandarenot

classifiedbytheNevadaDept.of

Wildlife;BisonareclassifiedbyNevadaDept.ofAgriculture.

No

Bisonnotreferencedinwildliferegulations(NevadaAdministrativeCode(NAC)502,503or

504);Regulationsnotethatpossessionof

bisondoesnotrequireapermit;Regulationspertainingtodomesticbisonaredescribedin

NAC571.

NoAgricultureand


Oklahoma No Yes

Classifiedasdomesticated

animals;Protectedontwopreserves

(onefederalandoneprivate).

OnlyforWichitaMountains

NationalWildlifeRefugeandTNC

preserve.

Therearenoreferencestobison

intheGameandFishregulationsinTitle

29;Title800-25-25-3listsspeciesofwildlifeexemptfromwildlifepermitsorlicense;

Regulationspertainingtodomesticbison

aredescribedinTitle2(Agriculture)andTitle4(Animals)ofOklahomaCode.

No



SouthDakota

Yes,partially Yes

Identifiedas“Wildlife”onlyintheconfinesofNationalParkSystem;BisonarecontainedwithinCusterStatePark.

Yes;OnlywithintheStateandNationalPark

SystemandoneTNCpreserve.

SouthDakotastatutesTitle41donotmentionbisonanywhereinthe

wildliferegulations;Statelawsidentifybisonaslivestock.

No

Statusofbisonislivestockoutsidethe

NationalParkSystem;Managementundercaptivity;Agricultureandforestryconflicts;

Regulatorystatus.

Texas Yes Yes

TexasParksandWildlifeDepartmentconsiderswildbison

extirpated;FoundonlyinCaprock

StateParkandononeTNCpreserve.

Onlywithinonestateparkandone

TNCpreserve.

NolongerconsideredagameanimalinTexas-Parksand

WildlifeCodeChapter43;TexasAgricultureCode(chap.2.005)recognisesbisonaswildanimals

indigenoustothestatebutcanberaisedforcommercialpurposes

topreservethespecies.

No

Agricultureconflicts;Smallparcelsofpublicorprivate


Utah Yes Yes

Freeroamingpopulationsare

foundintheHenryMountainsandonAntelopeIsland;

Utahjustcompletedareintroductionto

theBookCliffs.

Herdmanagementplanbeing

developedfortheHenryMountainspopulationand

BookCliffs.

Wildbisonaremanagedunder

regulationsinTitle23ofUtahCode;

RegulationspertainingtodomesticbisonaredescribedinTitle4of

UtahCode.

Yes;Recentintroduction

toBookCliffs.

Agricultureconflicts.



Country/State

province


Wildlife Status


Goal


Conservation

proposals for

restoration



Wyoming Yes,partially Yes

“Wildlife”withinnationalforestandnationalparksofParkandTeton

countiesintheGYA;Areclassifiedas

domesticanimalsintheremainderofthe

state.

Yes,inNERandGTNP

ManagementPlanandEIS;Yellowstonepopulation

conservedthoughInteragencyBisonManagementPlan

withMontana

WY(WyomingFishandGame

Commissionregulation)11-6-32viclassifiesbison

aslivestockunlessotherwisedesignated

byLivestockBoardandWildlifeCommission;WY

23-1-302xxvigivesauthoritytodesignate

individualbisonorherdsaswildlife;

ManagementPlanandEISforbisonandelkonNERandGrandTetonNationalPark.

Yes;Northern

Arapahore-introduction

totheWindRiver

Reservation.

Statusofbisonoutsideofdesignatedareasinstatute(ParkandTetonCounties);

DiseasestatusofYNPandJackson-GrandTetonbison;

Agricultureandforestryconflicts;RegulatorystatusoutsideofParks.

Canada Yes Yes

TheGeneralStatusofSpeciesforplainsbisonis

Sensitive;Plainsbisonpetitionedforendangeredstatus

denied-CurrentStatusThreatened;

WoodbisonarelistedasThreatened;Bothsubspeciesaremanagedasnative

wildlifeonsomeCanadianParksandinsomeprovinces

No,plainsbison;Yes,woodbison,

inNationalRecoveryPlan.

1996AccordfortheProtectionofSpecies

atRiskinCanada;SpeciesatRiskAct,

2002;COSEWICdesignatedplains

bisonthreatenedinMay2004;Wood

bisonwereclassifiedasendangeredin1978moveduptoThreatenedin1988

(COSEWIC);CanadaNationalParks

Act(2001);WoodbisonareonThe

RecoveryofNationallyEndangeredWildlife(RENEW)prioritylist.

Yes,inBanffNational

Park,Waterton

LakesNational

Park,Grasslands

NationalParkfor

plainsbison;NationalRecoveryPlanfor

woodbison.

Absenceofstrategicplanningforplains

bison;Multiplejurisdictionsandcoordinationof

agencies;Agricultureandforestryconflicts;Diseasetransmissiontocattle:Diseased

statusofsomeexistingwildbison;

Managementincaptivity

Alberta

Yesforwoodbison;Noforplainsbison.

Yes

Considerplainsbisonasextirpated;

PlainsbisonarenotlistedundertheAlbertaWildlifeAct;Plainsbisonlistedatriskin2000statusreport;Listswood

bisonasendangeredintheHay-Zama

woodbisonprotectionareain

NWAlberta.

Noforplainsbison;Yesfor

woodbison,withNationalRecovery

Plan.

1985PolicyfortheManagementof

ThreatenedWildlifeinAlberta;AlbertaWildlifeAct(1998)

2000StatusofAlbertaWildSpecies.

Yes;inBanffandWatertonNationalParks.

Legalstatusofplainsbisonis“livestock”;

Agriculturalandforestryconflicts;

Conservationstatusofhybridbisonin

WBNP.

BritishColumbia Yes Yes

ForplainsbisontheGeneralStatusof

Species=Sensitive.GeneralClassis“BigGame”and“Wildlife”;ListedasVulnerable;

WoodbisonareontheProvincial

RedList-Imperiledsubspecies.

Noforplainsbison;Yes–for


Plan

BritishColumbiaWildlifeAct(1996)GeneralStatusof

SpeciesinCanada(CESCC2001);

ProvincialBlueListandProvincialRed

List(BritishColumbiaConservationData

Centre2000).

Noforplains

bison;Woodbisonunder

NationalRecovery

Plan.

Agriculturalandforestryconflicts;

Plainsbisonoutsidetheiroriginalrange.


Country/State

province


Wildlife Status


Goal


Conservation

proposals for

restoration




Country/State

province


Wildlife Status


Goal


Conservation

proposals for

restoration



Manitoba

Yesforwoodbison;Noforplainsbison.

Yes

ProvincialHeritageStatus-S1-

SusceptibletoExtirpation;Listedas“atRisk”byCESCC;Plainsbisonarenotlistedas“Wildlife”butareclassedasLivestock;Wood

bisonareprotectedintheChitekLake

area.

Noforplainsbison;Yesfor


Plan.

ManitobaWildlifeAct(2004);ManitobaAgriculture,FoodandRuralInitiatives(2003).

Noforplains

bison;Woodbisonunder

NationalRecovery

Plan.

Statusofplainsbisonas“livestock”;

Agriculturalandforestryconflicts.

Sas-katche-

wanYes Yes

ProvincialHeritagestatus

-S3=Vulnerable;CESCCstatusas“maybeat

risk”;Bisonare“Wildlife”but

therearenoopenhuntingseasons;

DepartmentofNationalDefenseoffersprotectionduetoprohibitionoftrespassexcept

byColdLakeFirstNations;

FirstNationshaveaboriginalhuntingrights;protectedinBuffaloPoundProvincialPark,

PrinceAlbertandGrasslandsNational

Parks;NatureConservancyof

Canada(NCC)OldManonHisBack

ConservationArea.

NoforplainsbisonexceptinNational

orProvincialParks;Yesfor


Plan.

SaskatchewanWildlifeAct(1998);The

WildlifeRegulations,1981;SaskatchewanGameFarmPolicy

1998includescaptivebison;RangeAccessAgreementbetween

CLFNandDND(2002);Saskatchewan

ParksAct(1997);CooperativeInter-JurisdictionPlains

BisonManagementStrategy.

Plainsbisonin

GrasslandsNational

Park;Woodbisonunder

NationalRecovery

Plan.

Agricultureandforestryconflicts;Limitedsuitable

habitat.

NorthwestTerritories Yes Yes

Bothplainsandwoodbisonare“Wildlife”;Wood

bisonaredesignatedasindangerof

becomingextinct;Someregulatedhuntingofwood

bisonisallowedindesignatedherds;

Importationofplainsbisonprohibited.

Yes,woodbisoninNational

RecoveryPlan;Bisonharvestis

regulatedunderaco-managementprocess;Hook

Lakeismanagedunderaspecific

HookLakeRecoveryPlan.

NorthwestTerritoriesWildlifeAct(1964)designatedwoodbisonaprotectedspecies;Agency

policiespreventplainsbisonranchesor

introductiontothewild.

No

Conservationstatusofhybridplains/woodsbisonin

WBNP.

Yukon Yes Yes

Bothplainsandwoodbisonare“Wildlife”;Wood

bisonareaprotectedspecies;

Importationofplainsbisonprohibited.

Yes,woodbisoninNationalRecovery

Plan;Bisonaremanagedonasustainedyieldbasisunderacooperative

managementplan.

YukonWildlifeAct(2002);Agency

policiespreventplainsbisonranchesor

introductiontothewild.

No



andprotectionoftheJanos-Hildagobisonherdisanexampleof

thisrisingconservationinterest.Bisoninthisspecificpopulationare

protectedbyendangeredspeciesstatusunderfederallaw.Allother

bisoninMexicoareprivatelyownedandmaintainedonfenced

privateproperty.

Over93%ofthebisoninNorthAmericaareprivatelyownedand

managedforcommercialproduction(Chapter7).Bisoncanbekept

asdomesticlivestockinalloftheU.S.Thesebisonareprivately

ownedandtypicallymanagedformeatproductionorbreeding.In

Alberta,Saskatchewan,andManitoba,wherebisonareregulatedas

livestock,individualsintheprivatesectormayownbison.InBritish

Columbia,bisonmaybeproducedcommercially,butagame-

farminglicenseisrequired.Commercialherdsownedbyindividuals,

corporations,orNGOsaremanagedindependently,subjectto

marketforces,andregulationsgoverninganimalhealthandtrade.

IntheYukonandNorthwestTerritories,existingpolicypreventsthe

establishmentofplainsbisonranchesortheirintroductionintothe

wild.Thereisnounifiedconservationeffortorregulatoryframework

thatencouragesorfacilitatesconservationofcommercialbisonas

wildlifeatnational,stateorprovinciallevels.The“laundering”ofwild

animalsthroughcaptive-breedingoperationsandfarmshasnotbeen

detectedinCanadaortheU.S

8.3.2 Disease status

Earlyinthehistoryofbisonrestoration,diseaseswerenotconsidered

veryimportantandrestorationeffortsproceededwithlimitedconcern

forthetransferofpathogens.Asaresultofsignificantfailuresto

guardagainstdiseasetransferandcontrolduringtranslocation,bison

restorationprojectstodayhavetoovercomesomehistoricbaggage.

Withthedevelopmentofanextensiveandaggressivedomestic

animaldiseasecontrolprogrammeinNorthAmericaduringthemid

tolate1900s,theimplicationsofdiseasestowildliferestoration

hasincreased(Friend2006).Furthermore,withthesuccessful

restorationofmanywildlifespecies,andthesubsequentincreasein

theirdistribution,thesesamediseasesarenowveryimportanttothe

wildlifecommunity(Wobeser1994).Finally,increasedglobalisation

andthehighmobilityofsocietyareincreasingthelikelihoodof

pathogentransferacrosscontinents,therebyincreasingthevigilance

ofdiseasecontrolprogrammes(Friend2006).Asaresult,efforts

toconductbisonrestorationwillhavetoconsiderthesignificance

ofdiseasesinrestorationprojects.Foracomprehensivereviewof

diseasessignificanttobisonconservation,thereadershouldrefer

toChapter5ofthisdocument.Unfortunately,diseaseissuesoften

trumpconservationinterests,especiallywhentheconservation

actionsarelikelytocomeindirectconflictwithpowerfulagricultural

industries.Thiswillnecessitatethecarefulselectionofsource

Country/State

province


Wildlife Status


Goal


Conservation

proposals for

restoration



Mexico Yes Yes

Appearedasextirpatedin1994;

In2002red-listJanosbison

werelistedasendangered.

Notofficially,howevernon-governmental

conservationisemergingandproposinga

long-termvisionforconservation

preserves.

SecretariadeDesarrolloSocial,1994-NOM-059-

ECOL-1994.SecretariadeMedioAmbienteyRecursosNaturales-NOM-059-

ECOL-2001.

Yes,DevelopingaNationalRecovery

Plan.

Agricultureconflicts;Lackofsuitablehabitats:Small

propertiesavailable;Economicandmarket

obstacles;Lackofpublicinterest:Adevelopingwildlife

conservationprogramme;Varied

statusoftheJanosbisonattheinternationalborderwithNewMexico

TribalandFirstNations

Yes Yes

VariesbytribeorFirstNation;Mosttribeswithstrongculturalhistoriesprotectbisonfortribaluse;The

IntertribalBisonCooperativehas57membertribesthat

areactivelypursuingbisonmanagement

forculturalandcommercialinterests.

Yes,dependingupontribal

conservationprogrammes;Sometribes

aredevelopingadvancedgame

codesandsophisticated

speciesrestorationandmanagement

plans.

VariesbutgenerallydeterminedbyTribal

CouncilandmanagedbyTribalFishand

GameCommissions;IntertribalBisonCooperativewas

formedtoencouragetherestorationof

bison;Culturalconsiderationis

primarydriverforlegalandpolicy

considerationsbyeachtribe.

Yes

Variabilityoftribalgovernment

structureandfunction;Agriculture

conflicts;Variablewildlifeconservation

andmanagementinfrastructure.


stock,extensivetestingandscreeningofsourceherds,health

monitoringofherds,andregulatoryinvolvementintheprocess

oftranslocation(Table8.2).

Successfulrestorationprojectswillneedtonavigatetheanimal

healthregulatoryprocessnecessarytopermittranslocationof

bisonandtoaccomplishtheeventualestablishmentofhealthy

conservationherdsinNorthAmerica(seeChapter5).The

keydiseasecategoriesthatneedtobeconsideredinbison

restorationare:ForeignAnimalDisease(FAD)events,regulatory

diseases(acrossinternationalboundariesandwithincountry

jurisdictions),anddiseasesofsignificancetolivestock,but

notregulated.Aforeignanimaldiseasewillcausesignificant

impacttobisonrestorationandagriculturalactivitiesinany

jurisdiction.Asignificantresponsenetworkisalreadyavailable

toaddressFADswithincountries,states,andprovinces.This

responsenetworktypicallyinvolvesfederal,state,andprovincial

agriculture,wildlife,andpublichealthagencies.Anysuchevent

involvingsourcebison,oronarestorationlandscape,would

haltarestorationprojectandstopmovementofindividualsfrom

aninfectedsourcestock.Abisonconservationeffortisatrisk

whenabovineFADarrivesinanycountry,andasubsequent

federalresponseisrequiredtoimmediatelystopmovementof

allaffectedanimals.Regulatorydiseasesontheotherhandare

typicallymoremanageable,withregulatorystepsrequiredto

allowmovementafterhealthstandardsaremet.Althoughthey

aresignificant,thereareestablishedprotocolstotest,manage,

andevencontrolmanyofthesediseases.Eachdiseasehasits

owncharacteristicsandsubsequentlythechallengesofdisease

testing,managementandcontrolvary.Therehavebeenmany

historicefforts,somesuccessfulandsomenot,tocontroland

eliminatethesetypesofdiseasesinbison.Thishistoricrecordis

agoodplacetogotoseewhatworksandwhatdoesnot.

Thesciencebehindwildlifediseaseissuesisimproving,but

moreworkisneeded(Friend2006).Considerableresearchis

neededtoestablishquarantineandtestingprotocolsrequired

toensurethesafemovementofanimals.Tobecertainthat

restorationprojectswillnotintroducenewdiseases,or

exacerbateexistingdiseases,itisimportanttoaccuratelyand

reliablyestablishthehealthbackgroundofsourceherdsandof

thewildanddomesticanimalswithinrestorationareas.There

willbemanyagriculturalinterestsexaminingbisonrestoration

efforts,soduringarestorationproject,utmostattentionshould

begiventocommunicatingthehealthpreventionmeasures

taken,andtestinginformationobtained.Itislikelythat

agriculturalconflictswillbeoneofthemajorimpedimentsto

restoration,butembracingmodernapproaches,withcareful

monitoringofpopulationhealthandintegratingregulatoryhealth

officialsintotheprojectsfromthebeginning,canmitigatemost

diseaseissues.Restorationeffortsshouldestablishandmaintain

regularcommunicationwithstate,provincial,andfederalanimal

healthregulatorsandotherappropriatepublichealthagencies.

Generalcommunicationsshouldalsobeestablishedwithkey

animalhealthorganisations,suchastheU.S.AnimalHealth

AssociationorWildlifeDiseaseAssociation,toensurethatthe

besthealthinformationisbeingopenlydiscussed

andsharedwithaffectedgroupsandindividuals.

Restorationprojectsthatinvolveinternational

transportofbisonaresubjecttoadditionallegal

andpolicyconsiderations.Forexample,increased

animaldiseaseregulationsduetoanydiscovery

andcontrolofbovinespongiformencephalopathy

(BSE)acrosstheU.S.–CanadianorU.S.–Mexican

borderswillundoubtedlycomplicatetrans-

boundarymovementofbison(APHIS,USDA

2007).Untiltheserestrictionsareeasedtherewill

belimitedopportunityforinternationalmovement

ofbisondespiteanyevidencethatthisdisease

actuallyexistsinAmericanbison.Restoration

planningwillneedtoincludeathoroughsearch

ofcurrentinternationalborderrestrictionsrelated

todiseasecontrol.Earlydiscussionswithanimal

healthregulatorswillbeessentialtoidentify

anydiseaseregulationsandspecifictesting

requirementsfortransportofbisonacrossan

internationalboundary.

Diseaserestoration is prevented

Significant Impediment

Medium Impediment

Locally Significant

AnyFAD* x

Anthrax x

BovineTuberculosis x

BSE** x

Brucellosis x

MCF*** x

JD**** x

RespiratoryDiseases(e.g.BVD,IBR,BRSV,PI3,Bacterial)

x x

Endoparasites x

Ectoparasites x

OtherBacterial/Viralinfections

x

table 8.2 Some diseases that will or may have implications to bison restoration.

*Foreignanimaldisease**Bovinespongiformencephalopathy***Malignantcatarrhalfever****Johne’sdisease.


8.4 Legal and policy Obstacles hindering Conservation of Bison

Bisonconservationandrestorationintersectsdirectlywithmany

laws,rules,andpolicieswithinacomplexsocial-economic-

ecologicalmatrix.Isenberg(2000)detailedthehistorical

relationshipsofsocialandeconomicchangetopreservationofthe

bisonattheturnofthecentury.Bisonwerecaughtinavortexof

social,economic,andecologicalchangeontheGreatPlains,and

werenearlyexterminated(Isenberg2000).Thesechangesremain

thecentralthemesforanongoingmodernGreatPlainsdrama.

Thecontinuedexpansionofthehumanpopulation(exceptinrural

areasoftheGreatPlains,whereitisdeclining),thedominantuse

ofprairiegrazinglandsfordomesticlivestock,andtheconversion

ofnativeprairietocropland,haveledtopersistentcompetition

betweenwildbisonandhumansforprimaryuseofgrassland

habitats.However,intermixedamongtheseagriculturaland

urbanisinglandscapesarerelativelyintactislandsofsuitableprairie

habitatwithpotentialforbisonrestoration.Theseremainingintact

landscapesaretypicallyamixofprivateandpubliclandandare

characterisedbyamosaicoflandownership,landmanagement

regimes,socio-economicinterestsandlandusepolicies.Excluding

diseasestatusofbison(seeabovesection),wehaveidentifiedsix

principleobstaclesthataremajorimpedimentstoconservationof

bisonwithinthissocial-economic-ecologicallandscape.Although

therearemanyotherminorobstacles,mostofthesearesite

specificinnatureandcanbeaddressedwithouteffortstoshape

law/policyorpublicattitudesinarangewidescale.

Themostsignificantlegalandpolicyobstaclestowildbison

restorationareindirectlyderivedfromsocio-economicconcerns

andpersistenthistoricalparadigmsofbisonmanagement.The

greatestimpedimentissocialintoleranceforalargegrazing

bovidthatisperceivedtocompetewithotherinterestsadjacent

to,orwithin,prospectiveprairielandscapessuitableforbison

restoration.Asaspecies,thebiology,behaviouralplasticity,and

wideecologicalscopeofbisonprovideunlimitedopportunity

forrestorationeffortswithahighprobabilityofsuccessin

recolonisingavailablegrasslandhabitats.

8.4.1.1.1 Confusing legal classification and status

Therearerelativelyfewstatesandprovinceswhereconservation

bisonherdsarelegallyclassifiedaswildlife(seeTable8.1).Other

states/provinceshavemixedstatusforbisonandthereissome

confusionrelativetothelegalauthorityorpoliciesofotherbison

herds.Manystates/provinceswithintheoriginalrangeofbisonhave

classifiedbisonasdomesticlivestockandmanagementauthorityis

vestedwithinagriculturalagencies.Inaddition,manyconservation

herdsaremanagedbyfederalagencies,suchastheNationalPark

Service(NPS)orU.S.FishandWildlifeService(USFWS)Refuge

System,addingafederallayeroflawsandpoliciesuponbison.This

confusinglegalclassificationandstatusincreasesthedifficultiesin

conservingthespeciesinacomprehensivemanner.

Privatelyownedbisonherdsdonotenjoylegalstatusaswildlife.

Somebisonownedbyprivateproducersmayhaveconservation

value(e.g.,goodgenetics),butmanagementisprincipally

productionoriented.Severalprivatelyownedbisonherds

managedbyNGOsaremanagedinanecologicallyrelevant

manner,butarealsonotlegallyclassifiedaswildlife.InAlaska,

woodbisonwerenotconsiderednativewildlifeformanyyears

bytheUSFWS,butplainsbisonherdswereestablishedbythe

StateofAlaskaandmanagedaswildlife.Federallyownedbison

herdsaretypicallymanagedaswildlife,althoughbehindhigh

fences,buttheyareusuallynotrecognisedasnativewildlifeby

stateauthorities.Thisconfusioninthelegalstatusofbisonis

probablythesinglemostimportantobstacleimpedingecological

restorationandhinderinganationwideconservationstrategyfor

thisspecies.

8.4.1.1.2 Historical management policies

Addingtotheconfusedlegalstatusofbisonistheconsistent

policyofestablishingandmanagingbisonbehindhighfences

bystateandfederalagencies.Thismanagementparadigm,

establishedintheearly1900stoprotectthespecies,has

persisted,furtherconfusingthemanagementpolicyframework

andpublicattitudetowardbisonasawildlifespecies.This

confusingmanagementapproachtobisonisnotconsistentwith

otherwildlifeandhasproducedthesecondmostsignificant

obstacletoecologicalrestoration.Fewagenciesormembersof

thepublicidentifybisonasnativewildlifedeservingthesame

statusasotherfree-rangingwildlife.Apublicrecognitionforthe

needtomanagebisonaswildlife,inanecologicallysensitive

way,isessentialtosuccessfulrestoration.Ecologicalrestoration

ofbisonwillbehindereduntilthismanagementparadigmshifts

andsocialtoleranceisdevelopedtoallowfree-rangingbisonon

nativeprairiehabitats.

8.4.1.1.3 Complex partnerships needed to

manage large landscapes

Bisonpopulationsmanagedonpubliclandsareconsidered

asthecoreofthewildherdsbeingmanagedtoconserve

thespeciesforthefuture(Boyd2003;Knowleset al.1997).

However,fewpubliclandmanagementagencieshavea

sufficientlandbasetomanagebisonpopulationsinamanner

thatallowsfornaturalselectionprocesses.Bisonneed

largelandscapestoallownaturalmovementsandexpress

appropriateecologicalfunction.Unfortunately,mostwildbison

arebeingmanagedassmallpopulationsonrelativelysmall

areasbysingleagenciesortribes.Forgingthepartnerships

tomanagepopulationsacrossmultiplejurisdictionsonlarge

landscapesseemstolimitexistingconservationefforts.Building

partnershipstomanagewildbison,asapublictrustresource

byacoalitionofprivateandpublicinterests,whiletheoretically

feasible,hasbeenlimitedinpractice.


Theproblemsofgovernanceandscalehavebeenwell

discussedintheliterature(WestleyandMiller2003;Wilkeet

al.2008).Theretypicallyisawiderangeofactorsassociated

withtheconservationoflargelandscapesandspecieswith

largespatialneedssuchasbison.Itiseasytounderestimate

thecomplexityofownershippatternsonlargelandscapes

andtomissidentifyingkeyactorsonthisconservationstage.

Furthermore,differentkindsofactorswillhavedifferentrights,

interests,andcapacities,andwillneedtobeapproachedin

differentways(Wilkeet al.2008).Thechallengeofforming

complexpartnershipsattheappropriatescaleisformidableand

oftendiscourageseffortstoconsiderlarge-scaleinitiatives.

8.4.1.1.4 Defining the social and economic value

of wild bison

Manylegalandpolicychangesnecessaryfortheecological

restorationofbisonarelinkedtosocialandeconomicfactors.

Agenciesandconservationistsneedtoidentifytheeconomic,

social,andecologicalbenefitsofrestoringwildfree-ranging

bison,whileprotectingexistingculturalandeconomicinterests

(Geist2006).Thevalueofrestoringwildbisonmustbe

expressedinamannerthatdoesnotnecessarilydiminishthe

economicvalueofexistinglivestockandcommercialbison

marketsmanagedunderanagriculturalparadigm.Thismaytake

creativeapproachesinvolvingpolicyadjustmentsandparadigm

shiftsamongcooperatingagencies/privatesectorsthatoptimise

complimentarylandusestrategiesandmitigateidentified

conflicts.Thisprocesscouldbesupportedbytaxincentives,

paymentforenvironmentalservices,ecotourism,incentivesfor

landownercooperation(e.g.,Colorado’sRanchingforWildlife

Program),extensionservices,andtrainingforanewgeneration

oflandownersandmanagers.

8.4.1.1.5 Coordination of policies, rules, and

regulations by government

Coordinationofmanagementpolicies,rules,andregulations

(orthelackthereof)byvariousgovernmentshasalsohindered

bisonconservationefforts.Becausenosinglegovernment

agencyownsormanagessufficientlylargeblocksoflandto

sustainfree-rangingbison,cooperationbetweenagencies

isneededforrestorationandconservationplanningand

implementation.Manyagencies’missionsarenotreadily

compatiblewithcooperativemanagementstrategiesneeded

forconservationofbisonatlargescales.Furthermore,many

landmanagementagencieshavedirectedmissionsandgoals

thatmaynotimmediatelysupportthetypesofpolicychanges

requiredtomanagefortheconservationofbison.Inaddition

tocoordinationamonggovernmentagenciesthereisoftena

compellingneedtocoordinatewithandamongTribalandprivate

landsinfluencedbyotherpoliciesandmanagementobjectives.

8.4.1.1.6 Agricultural conflicts among

mixed land ownership

Themostsignificantconflictsassociatedwithrestoringwildfree-

rangingbisonarelikelytobewithagriculturalneighboursliving

nearconservationreserves.Establishingfree-rangingwildbison

herdsinNorthAmericawillundoubtedlyleadtoconflictsfrom

cropdamage,foragecompetitionwithlivestock,mixingwith

livestock,possibleinterbreedingwithcattle,diseaseissues,and

damagingprivateproperty.Theseagriculturalconflictsarenot

entirelyuncommonwithotherlargeherbivores.

Thesesixpolicyobstaclesarequitecommonacross

international,state/provincial,andpublic/privatejurisdictional

boundarieswithintheoriginalrangeofbison.Bisonrestoration

mustoccuratsufficientlylargelandscapescalesthatfew,if

any,individualagencieswillbeabletoimplementaneffective

managementprogrammeontheirown.Coordinationofagency

missionstoconservewildbisonmustinthelongrunbea

negotiatedprocesstoensurejointconservationgoalscanbe

establishedandimplementedwithinthelegalframework.In

addition,conservationgoalsmustbeestablishedtoencourage

privatelyownedpopulationsofwildbison(asdefinedelsewhere

inthisdocument)tobemanagedoverthecourseofmanyyears

inamannerthatallowsrancherstobuildnewmarketsthat

provideeconomicbenefitsforconservingthecharacteristicsof

ancestralbisonherds.

Otherobstaclestorestorationinclude:longtimescales,

institutionalresistance,funding,andconservationmission

creep.Mostlarge-scaleconservationprojectsforlong-lived

mammalsneedtoplayoutacrosslongtimescales.Itiseasy

forconservationpartnerstofatigue,andforshiftingpolitical

andsocialclimatestomakeextendedtimescalesproblematic.

Institutionalresistanceisinevitablewithinandamongthe

cooperatingagenciesandprivatesectorpartnersinvolvedin

abisonrestorationproject.Withinagenciesandorganisations

thereislikelytobesomeinternalresistancetovariousaspects

oftheproject,socarewillbeneededtobuildreasonable

consensus.Althoughmanyagencyorprivategroupsmay

supporttheconceptofrestoration,thereisafundamentalneed

forfundingandcontributionfromallcriticalpartners.Finally,with

everyconservationprogramme,theimplementationcancreep

offtargetormovebeyondintendedgoals.Thishasatendency

todismantlesocialandpoliticalsupportforaprojectbycreating

adifferenttypeofmanagementorobjectivethanwasoriginally

identifiedandagreeduponbystakeholders.Forexampleas

landscapesbecomelarger,andsomemeasureofsuccessis

achieved,theremaybeatendencytomovetheconservation

focus.Conservationandrestorationstrategiesandplanning

effortsneedtoclearlyarticulatetheconservationgoalandbe

abletomeasureprogressandidentifycriticalbenchmarksfor

meetingthosegoals.


8.5 Overcoming Obstacles to the Ecological restoration of Bison

8.5.1 Disease management considerations

Animalhealthanddiseaseissuescanpresentsignificant

obstaclestobisonrestorationefforts.Thepresenceofregulated

diseasesinbisoncanpreventthetransportofbisonacross

jurisdictionalboundariesandlimitsaccesstosourcesofbison.

Potentiallyimportantsourcesofgeneticallyreputablebison

forrestorationfromWBNPandYNParedeemedunsuitable

becauseoftheirdiseasestatus.However,recentresearchefforts

areexploringmethodsofquarantiningbisonfromthesesources

todetermineifdiseasefreestatuscanbeestablishedforanimals

passingthroughstrictquarantineprocedures(Nishiet al.2002b;

2006).Theuseofeffectivequarantinetoreleasethesegenetic

sourcesofbisoncouldbeextremelyhelpfulforenhancing

accesstoabroadersourcegenepoolforrestoration.

Beforeanimalscanbetranslocatedforrestoration,eachstate

orprovinceandinternationalborderthatwouldbecrossedby

bisonwillrequirespecifichealthtests.Whendesigningspecific

restorationprojects,itisessentialtocontactState/Provincial

orFederalVeterinarianssothatrequireddiseasetestingisa

clearlyarticulated.Appropriateregulatoryveterinarian(s)havethe

expertisetoestablishwhichdisease(s)requirescreening,and

whichapprovedtestprotocolsanddiagnosticlaboratoriesare

acceptable/requiredforhealthclearanceforspecificjurisdictions.

Thesehealthapprovalsneedtobeobtainedbeforetransporting

anybisonacrossjurisdictionallines.Goodhealthmonitoringof

thesourceherdcanprovideimportantinformationtosupportthe

testingcarriedoutpriortotransport.Agoodhealth-monitoring

programmewillidentifyexistingdiseasescirculatingamongthe

sourceherd,andincludebackgroundinformationregardingthe

presenceorabsenceofregulateddiseases.

Infectiousdiseaseisanemergingthreatthatconservationists

maybeillequippedtomanage(Woodroffe1999).Despitethese

limitationsthereareseveraldiseasemanagementmodelsacross

theglobethatcouldhelpsupportdiseasemanagementplanning

inbison(Osofskyet al.2005).Throughcarefulplanning,and

researchofexistingdiseasemanagementmodels,thisissuecan

besubstantiallyreducedinscopeandimpact.

8.5.2 Legal status and policy considerations

Inordertoaddressobstaclestoecologicalrestorationof

bison,itisimportanttoidentifythestrategiccomponentsofa

continentalconservationplan.TheIUCNBSGhasprovidedthis

strategicframeworkandassociatedtechnicalguidelinesfor

bisonconservationtohelpagenciesandthepublicaccomplish

ecologicallyrelevantconservationprojects.Thisframeworkcan

assistinresolvingissuesofinternationalstatusandovercome

legal/policyobstaclesfromastrategicperspective.Whilethis

continentalwidestrategyshouldbeusefulinadvisingsome

oftheoverarchinglegalandpolicychangesnecessaryto

achieveconservationmissions,federalstate/provincialand

localauthoritieswillneedtobeinvolved,andsupportiveof

significantlocalchangesinpolicies,sothatrestorationprojects

canbeaccomplished.

Formostbisonrestorationprojectstoadvance,changesin

lawsandpolicywillbenecessary,buttheymustbedesigned

toencouragebisonconservationinanecologicallyrelevant

mannerwithdueconsiderationofthepotentialsocio-

economicconsequencestocountries,state/provincesorlocal

communities.Laws,rules,andpoliciesofgovernmentscan

impedeconservation.However,theymaybetransformedinto

supportiveframeworksifthereissocialacceptanceanda

highvalueassociatedwithrestorationgoals.Comprehensive

policiesandlawsneedtobedevelopedthatpromoteecosystem

conservation,withoutbeingoverlyprescriptive.Therewillbe

aneedfornegotiation,compromise,andcooperationinthe

processofchanginglawsandpolicies.Suchprocessesare

interdisciplinaryinnature,requiringintegrationofthedisciplines

ofeconomics,law,ecology,andsociologytobesuccessful

(Wilkieet al.2008).

8.5.2.1 role of the non-governmental organisations

NGOscanplayakeyroleadvocatingforthenecessary

changestolaws,rules,andpoliciesthathinderrestoration.

NGOscanactivelylobbyfornecessarylegal/policychangesby

federal,stateorprovincialgovernmentstoovercomeidentified

obstacles.Theycanprovideandsecureorsupportgovernment

fundingforconservation.CoalitionsofNGOsandgovernment

agenciescanbeformedtoadvocateforspecificconservation

efforts.NGOscouldalsosupporttheecological,economic,

culturalandspiritualinterestsofindigenouspeopleswithan

interestinbisonconservation.Theycanaidlocalcommunity

groupsinnegotiationsandhelpthesecommunitiesinfluence

stewardshipofnaturalresourcesintheirarea(Fraseret al.2008).

Finally,someNGOscouldhelptoresolveinternationalissues

relatedtostatusandlegal/policyobstaclesassociatedwith

individualprojects.Whilemanyagenciesmustoperatewithin

jurisdictionalboundaries,NGOscantranscendtheselimitations

andbrokercommunicationandcooperationamongagencies.

ThehistoricmodeloftheAmericanBisonSociety(ABS),asa

consortiumofindividualsandgroups,isanexampleofhow

conservationorganisationscanplayapowerfulroleinspecies

restoration.TheABSadvocatedfortheformationofbison

preservesinthewestandsupportednewwildlifepolicyand

legislationtopreserveaspeciesatthebrinkofextinction.

TheRockyMountainElkFoundationisanexcellentexample

ofaNorthAmericanNGOemployinglandpreservationand

activeadvocacytosupportconservationpoliciesthatcreate


suitablelandscapesforwildelk.TNCisanotherconservation

organisationthathasworkedeffectivelywithprivatelandowners

andgovernmenttoprotectbiodiversityandestablishprotected

areasthroughtheuseoflandpurchaseandeasements.TNChas

incorporatedbisononseveraloftheselandscapesasameansof

providingecosystemservices.

8.5.2.2 State/provincial and federal governance

Itisvitalthatgovernments(bothelectedofficialsand

governmentagencies)beengagedinpolicymakingand

legislationthatsupportbisonconservation.Government

agenciestypicallyestablishprocesseswithintheirstatutory

authoritytoevaluateandapproveappropriatepolicychanges,

andrecommendcongressionalandlegislativechanges,

necessarytoconductconservation.Itwillbenecessaryto

employalloftheinstrumentsandprocessesofgovernments

tomodifypoliciesorlegalstatutesaffectingbisonconservation

atstate,provincial,andfederallevels.Governmentagencies

canalsodirectpublicfundingandstaffresourcestosupport

implementationofarestorationproject,anddevelopthe

necessaryinteragencyagreementstoachieveconservation

goals.Itisnecessarythatelectedofficials,asrepresentatives

ofthepeople,approverelevantpolicies,andtodevelop

alegislativeframeworkthatsupportsbisonrestoration,

byempoweringtheappropriateagenciestoimplement

managementstrategiesforconservingbisonaspubliclyowned

wildlife.Forexample,opportunitiesforbisonrestorationcould

beincreasedbylinkingthemtoexistingpoliciesforlanduse

planningforecologicalintegrity.Thiswillrequirebuildingpublic

supportforpolicychangesandacceptancebyrespective

constituenciesthatthesegovernmentsserve,byusing,for

example,extensiveoutreach,publicadvocacyandeducation.

Itwillalsorequireeducatingandinfluencingkeypoliticiansand

governmentofficialswithcriticaldecisionmakingroles.

8.5.2.3 the private sector

Thereissubstantialevidenceofamassivechangeinland

ownershipandshiftingeconomiestakingplaceintheGreat

PlainsandWest,aswellassomemultiple-generationranchers

whoareentrepreneurialandreadyforchange(Powers2001).

Thisshiftinlandownership,economies,andvisionsbrings

opportunitiestocreateanewparadigmformanagingrangelands

ofhighconservationvalue.Privatelandownerscouldhavea

strongvoiceinfluencingelectedandagencyofficialsofthe

needforpolicychangesthatprovideincentivesfor,andremove

barriersto,bisonconservationonprivatelands.Therefore,there

iscurrentlyasubstantialopportunitytoengagelandownersto

petitiongovernmentforchange.

Privatelyownedbisonmanagedonprivatelyownedland

typicallypresentfewerregulatoryobstaclesthanencountered

inrestoringwildbison.However,privateherdsaretypically

managedunderaprivatepropertydecisionframework,which

maynotleadtoabisonherdofconservationvalue.Itis

difficulttoblendprivatepropertyrightswiththepublictrust

frameworkforwildlifewithoutnegotiationandcompromise.

Foreffectivecooperation,privateownersofbison,orbison

habitat,wouldhavetobewillingtosacrificecertainrightsand

submittopublicreviewandscrutinyofoperations.Government

partnerswouldalsoneedtobesensitivetoprivateproperty

rightsandtheeconomicvalueofthoserightsforindividualsor

corporationswillingtoengageinbisonconservation.Effective

cooperationshouldincludecreativeincentives,financialor

other,toencouragetheprivateentrepreneurtoengagein

bisonconservation.Forexample,conservationeasements

compensateland-ownersfortransferringspecificproperty

rights.Asnotedearlier,asystemforcertifyingproducerswho

followconservationguidelinesinmanagingtheirbisonherds

mayalsoprovideanincentive.

Toincreaseopportunitiesforlarge-scaleconservationofbison,

thereisaneedforfederalandstatepolicyprogrammesthat

fosterthecreationofprivate(for-profitornon-profit)protected

areas(PPAs).PPAsareoneofthefastestgrowingformsof

landandbiodiversityconservationintheworld(Mitchell2005).

However,unlikeAustraliaandmanycountriesinsouthern

Africa,theU.S.andCanadianfederalgovernmentsandstate

andprovincialgovernmentsdonotgenerallyhavepolicies

specificallysupportingthecreationofPPAs.TheIUCNhas

developedguidelinesfor,andexploredpoliciesandprogrammes

thatsupport,thecreationofPPAs(Dudley2008).Thedanger

isthatprivatebisonreservesmayquicklyshiftawayfroma

conservationmissionanddevolveto“privategamefarms”for

privatelyownedwildlife,forwhichmoststateshavepolicies

andregulations.Inaddition,privatenaturereservesmaybe

vulnerabletochangeofownershipandsubsequentshiftsintheir

missionunlessclearlegalinstrumentsareinplacetoprotect

conservationvalues.Clearguidelinesformanagementand

accountabilityforthelong-termsecurityofprivateprotected

areasisessential(Dudley2008).

8.5.2.4 Indigenous peoples

Manyprotectedlandscapesandseascapeswouldnotexist

withoutthedeeplyrootedculturalandspiritualvaluesheld

bythepeoplethatoriginallyinhabitedtheseplacesandwho

oftencontinuetocareforthem(Mallarach2008).Mallarach

(2008)pointsoutthatsafeguardingtheintegrityoftraditional

culturalandspiritualinteractionswithnatureisvitaltothe

protection,maintenance,andevolutionofprotectedareas.

Hence,protectedlandscapesandseascapesarethetangible

resultoftheinteractionofpeopleandnatureovertime.In

recentyearstherehavebeenmanyimportantdevelopments

inconservationandprotectionofimportantlandscapeson


indigenouspeoples’land(Dudley2008).Withintheoriginal

rangeofbison,thereareextensiveNative-ownedgrasslandand

mountainfoothillslandscapessuitableforbisonrestoration.

Thesetriballandspresentgreatopportunitiestorestorebisonin

aculturallysensitiveway,protectingtherightsandinterestsof

traditionallandowners.IUCNhasidentifiedbasicprinciplesof

goodgovernanceastheyrelatetoprotectedareasoverlapping

withindigenouspeoples’traditionallands.Inaddition,thereis

onegroup,theITBC,whosedefiningmissionisrestorationof

bison.Cooperationoftribesandtribalorganisationsisessential

totheconservationandrestorationofbisoninNorthAmerica

andshouldbeencouraged.GovernmentsandNGOsinNorth

Americashouldexamineandthenmodifycurrentpolicyand

legislationtosupportthetraditionalandculturalinterestsof

indigenouspeoplesrelevanttobisonrestoration.

Thereissignificantvariationinjurisdictionalpowersover

triballandscapes,rangingfromsovereigntyovertheland

toco-managementwithothergovernments.Itisimportant

tounderstandindigenouspeoples’rightsandtheirlevel

ofauthorityoverlandscapeswhendesigningrestoration

andconservationplansforbison.Itisequallyimportantto

understandtheculturaltraditionsandspiritualconnections

betweenindigenouspeopleandbison.Someofthis

informationistraditionalknowledgethatcanonlybeacquired

throughconversationwitheldersandtriballeaders.

8.5.2.5 Local communities and economies

Onekeyingredientofsuccessfulbisonconservationis

activestakeholderparticipationinthedevelopmentand

implementationofconservationprogrammes.Stakeholders

includeallpeopleorgroupsofpeoplewhoareaffectedby,or

canaffect,theconservationprogramme.Onpubliclandsitis

particularlyimportanttohavelocalsupport(individuals,adjacent

landownersandcommunities)forpolicychangesandnew

legislation,andtoavoidbacklashfromthetypesofregulatory

protectionthatmightbenecessaryforasuccessfulconservation

initiative(Merenlenderet al.2004).Forlandscapeswithmixed

jurisdiction(publicandprivate),itwillbenecessarytoengage

stakeholdersbydevelopingcriticalrelationships,buildingmutual

understandinganddesigninganappropriateco-management

framework.

Restoringbisontomixed-uselandscapeswillinvolveaddressing

conflictswithneighbouringlandowners.Theseneighboursneed

someassurancethatwhenconflictsarisetheywillbeaddressed

asrestorationprojectsareimplemented.Comprehensive

restorationandmanagementplanswillberequiredtoclearly

articulatepopulationgoalsandtoidentifyhowagricultural

conflictsaregoingtoberesolved.Ranchlandneighbours,

livingonagriculturelandsnearrestorationprojects,posea

greatchallenge,butmayalsoprovideasignificantopen-space

bufferessentialtothesuccessoflarge-scaleconservation

efforts.Measuresmustbedesignedtoappropriatelymanage

thedistributionofbisonandaddressanytrespassconflicts

thatarise.Otherconceptstoconsiderincludetheideaof

wildlifedamageinsurance,economicincentives,andcreative

conservation-incentivestoencourageandrewardtolerance

(Muchapondwa2003).

Ecosystemserviceshavebeendefinedas“the process by

which the environment produces resources that we take for

granted such as clean water, timber, pollination of plants,

and habitat for fish and wildlife”(Dalyet al.1997).Bison

restorationandconservationprogrammesshouldconsider

assessingthevalueofecosystemservicesassociatedwiththe

developmentofaconservationstrategyforbison.TNChas

madesignificantinvestmentsinpursuingthevaluationand

marketingofecosystemservicesasaconservationstrategy

andfinancingtool(Groveset al.2008;Nelsonet al.2009).

TNC,incollaborationwithStanfordUniversityandWWF,has

developedaNaturalCapitalProjecttobetterunderstand

theeconomicvaluesassociatedwithnaturalsystems(www.

naturalcapitalproject.org).Thisprojectdevelopedatoolknown

asInVEST(IntegratedValuationofEcosystemServicesand

Tradeoffs)forquantifyingecosystemservicesfortheirinclusion

innaturalresourcedecision-making.Theyalsoestablished

a“SwatTeam”ofecosystemmodellersandmapperswho

useInVESTtobringthevaluationsystemintopolicyand

decision-makingforconservationprojects(Groveset al.

2008).Approachessuchasthismaybeusefulinthevaluation

ofecosystemservicesassociatedwiththeconservation

oflargegrasslandlandscapesandtheroleofbisonasa

keystoneherbivoreonthoselandscapes.Werecommend

furtherexplorationoftheseemergingvaluationtoolsandtheir

applicationtotheconservationandrestorationofbisonin

NorthAmerica.

Inanothernovelprogramme,acoalitionofNGOs,stateand

federalagencies,ranchers,andresearchershasbeendeveloping

aPay-for-EnvironmentalServices(PES)programmeinFlorida

(Bohlenet al.2009).Thisprogrammecompensatescattle

ranchersinFlorida’snorthernevergladesforprovidingwater

storageandnutrientretentiononprivatelands.Keychallenges

tothisprogrammeinclude:identifyingabuyeranddefiningthe

environmentalservice;agreeinguponapproachestoquantifythe

service;reducingprogrammecostsinlightofcurrentpolicies;

andcomplexregulatoryissues.DesignofaPESprogramme

requiresnavigatingthroughacomplexregulatorymazecreated

bymultiplestateandfederalagencies(Bohlenet al.2009).

ThisnewmodelmayprovideanexamplefordevelopingaPES

onbisonlandscapes.Inthecaseofbisonrestorationitwillbe

challengingtomeettheneedsofmultiplestakeholders,andto

findthefirstentrepreneuriallandownerwillingstartanewtrend

byparticipating.


Significantchallengeslieaheadfortheformulationoflawsand

policiesaboutecosystemservices(Ruhl2009).Somecritical

stepssuggestedbyRuhl(2009)includebetterdefinitionof

propertyrights,policiesthatprimethemarketsforecosystem

services,designingbettergovernanceinstitutionsand

instrumentsforthesemarkets,andcreativeresearchtomeet

policyneeds.Somegovernmentsarealreadyengagedinthis

typeofwork,sointerestedreadersareencouragedinvestigate

programmeandpolicyinitiativesintheirregion(Freeseetal.

2009).Furthermore,conservationorganizationsareencouraged

tocoordinatetheiractivitieswithevolvinggovernmentinitiatives

tomoreefficientlyadvanceecosystem-basedconservation.

8.5.3 Coordination of agency missions, goals, regulations, and policies affecting bison conservation and restoration

Theremaybeaneedfornewgovernancesystemsthatwill

allowlocalcommunities,tribes,andgovernmentstoco-manage

grasslandreserveslargeenoughtosustainbison.Political

boundaries,agencypolicy,andlegaljurisdictionsneedtobe

creativelyblendedtocreateacooperativeatmospherefor

thesuccessfulestablishmentandco-managementofnewor

expandedbisonpopulationsinthefuture.Accomplishingthe

coordinationnecessarytoconducteffectiveconservationand

ecologicalrestorationwillbeformidable.However,therewardsfor

sucheffectivecoordinationwillgobeyondthebenefitofplacing

bisonontheland,andcouldencouragemuchmoreopportunity

toconserveotherspeciesassociatedwiththeselandscapes.

Itislikelythatsometypeofstandingco-managementcouncil

orcommitteemaybenecessarytocoordinatemanagementof

largelandscapeswithcomplexlandownershipandaffected

localcommunitiesandeconomies.Thiscommitteeshouldbe

structuredandfunctiontofacilitateandmaintainpartnerships

amongthevariousgovernmentagencies,NGOs,landowners,

andsportsmenorconservationgroupsthathaveinterestin

theprojectarea.Aco-managementcommitteecanencourage

efficienciesinfundingandcoordinaterestorationactivitiesofthe

variousstakeholders.Acommitteeshouldincluderepresentatives

fromlocalstakeholderswhoareaffectedbythecoordinated

managementeffort.ArecentannouncementbytheU.S.

DepartmentoftheInterior(USDOI)ofanewBisonManagement

FrameworkestablishedaUSDOIBisonWorkingGrouptohelp

coordinatebisonmanagementamongtheagencies.Theworking

groupprovidesaninitialefforttocoordinatemanyofthekey

federalagenciesinvolvedinbisonconservation,butdoesnot

includenon-governmentpartners.Thisworkinggroupcould

becomeanewmodelformanagingbisononmultiplesmall-scale

reservesasthoughitwereonelarger-scalepopulation,creating

aneffectivepopulationofsufficientsizetoprotectgeneticand

ecologicalintegrity.InMontana(NorthernYellowstoneEcosystem)

andWyoming(SouthernYellowstoneEcosystem)interagency

bisonorbison/elkmanagementplanswerecreatedthatdefined

aco-managementstrategytranscendingstateandfederal

jurisdictionalboundaries(Seechapter5).Publicparticipationin

theseprocesseswasachievedthroughnumerouspublicmeetings

wherestakeholderswereprovidedopportunitiestocomment

onandinfluenceaproposedco-managementdesign.Through

thisprocess,informationwasprovidedtothestakeholders,

andsomedegreeofacceptanceforproposeddecisionswas

negotiated.TheestablishmentoftheSturgeonRiverPlainsBison

CouncilinSaskatchewanisanotherco-managementexample

developedbylocalstakeholdersaffectedbybisonmanagement

onneighbouringfederallands.Theseexamplesrepresent

contrastingmodelsoftopdownversusbottomupapproaches

tobisonconservation.Bycombininglocal(bottom-up)and

national(top-down)approaches,betterformsofgovernancecan

evolve,naturalresourcesmaybemoreeffectivelymanaged,and

livelihoodscanbeimproved(Fraseretal.2008).

Detailedproject-specificplanningforecologicalrestoration

(seeChapter10ofthisdocument)shouldbecompletedby

agencies,NGOsandprivatepartnersinvolvedintheproject

areapriortoimplementinganybisonconservationproject.

Thesuccessfulcompletionoftheenvironmentalevaluations

requiredundernational,stateorprovincialenvironmentallaw

willbecriticaltotheadvancementofanybisonrestoration

projectinvolvingpublicland.Theseenvironmentalevaluations

willrequireapublicinvolvementprocessandshouldgather

inputfromallaffectedstakeholdersinameaningfulprocess.

Inadditiontopublicinvolvementsignificantpubliceducation

andoutreachshouldbedevelopedandimplementedduringall

phasesofarestorationproject.

Technicalsupportfromsciencegroups,suchastheIUCNBison

SpecialistGroup,canprovidethenecessarytechnicalguidance

forsciencebasedconservationstrategiesatthelocal,state/

provinceandcontinentalscale.Guidancefromthistechnical

groupcanidentifybestmanagementpractices,andrecommend

policyandlegislativechangesnecessarytosupportsound

conservationandrestorationinitiatives.Additionalguidancefor

ecosystemrestorationeffortscanbefoundthroughotherIUCN

publications(Clewellet al.2005;IUCN1998;Chapter10).

8.5.4 recommendations

Somefundamentallegalandpolicychangesrecommendedto

enhancebisonrestorationinclude:

1) Wheresocialacceptanceforwildbisoncanbeattained,

establishthelegalstatusofbisonasanativewildlife

speciesthroughworkingwithstate/provincial/federal

jurisdictions.

2) Modifycurrentpoliciesthatpreventpartnershipsandco-

managementamongagencies,privatesector,andtribes.


3) Developoutreachtostateandfederallandmanagement

agenciesencouraginglandmanagementagencies

toconsiderbisoninagencyplanningandpolicy

development.

4) Reformcurrentpoliciesgoverningsuitablebison

landscapestoprotectthecorehabitatconservationvalues

asdefinedinSandersonet al.(2008)andthisdocument.

Thisistoprotectthecorevalueoftheselandscapesfor

futureecologicalrestorationpendingsocio-economic

shiftsfavourabletobisonrestoration.

5) Developoutreachmaterialsidentifyingsocialand

economicbenefitsandecosystemservicesassociated

withrestorationofbisonandprairieconservationeffortsfor

localcommunities,theprivatesectorandgovernments.

6) Createadecisionframework,suitableforprivate

conservationefforts,thatencouragesrestorationstrategies

withanecologicalemphasis.

7) Createpoliciesoreconomicandconservationincentives

thatrewardprivatelandownerswhomanagefor

biodiversityincludingbison.

8) Establishnecessarystateandfederalregulationsandlegal

instrumentstosupportvaluationandcompensationfor

ecosystemservices.

9) Workwithanimalhealthorganisations(IUCNWildlife

HealthSpecialistGroup)andregulatoryagenciesto

encouragebisonfriendlyhealthregulations.

10) Identifyandsupportnecessaryresearchandmonitoring

tocultivateascience-basedbutadaptiveprocessfor

ecologicalrestorationofbison.

11) Encourageeconomicandpowerstructuresthatsupport

sustaininglocalcommunitiesandlifestyles.

12) Makeeffortstoreformpolicyandlegislationthatimpede

theinterestsandrightsofindigenouspeopletomanage

bisoninaculturallysensitivemanner.

8.5.5 recent initiatives to conserve and restore bison

Sandersonet al.(2008)presentacollectivevisionforthe

ecologicalrestorationofbisoninNorthAmerica.Froma

seriesofmeetingswithvariousconservationorganisations,

governmentagencies,indigenousgroups,bisonranchersand

privatelandownersa“VermejoStatement”wasjointlywritten

thatdescribeswhatecologicalrestorationofbisonmightlook

like.Fivekeyattributeswereidentifiedinthisstatementthat

createbothopportunitiesandchallengesforbisonrestoration,

suchaslargescale,longterm,inclusive,fulfilling,andambitious

efforts.Sandersonet al.(2008)exploredasharedvisionforwild

bisonrestorationwith20,50,and100-yeartimelines.Specific

initiativeswerenotdescribed,butarange-wideprioritysetting

methodologyresultedinascorecardmatrixwithwhichto

evaluatetheconservationvalueofpublicandprivatelyowned

bisonherdsandamapofpotentialrestorationareas.Significant

changesinthelandscapewherebisononceroamedarecreating

possibilitiesforbisonrestorationwherefewexistedbefore

(Freeseet al.2007;Sandersonet al.2008).

8.5.5.1 United States

IntheU.S.,therearenospecificfederaleffortsproposedto

protectplainsbisonbeyondtheboundariesofexistingnational

parks,monumentsorwildliferefuges.TheU.S.ForestService

(USFS)recentlyconductedanassessmentofitsmanagementof

nationalgrasslandsinMontana,NorthDakota,Nebraska,South

Dakota,andWyominganddismissedaproposedalternativeto

restorefree-rangingbison(USDAForestService2001).

TheU.S.SecretaryoftheInteriorrecentlyannouncedanew

managementframeworkforimprovingtheadministration

ofthevariousbisonherdsonFederalWildlifeRefuges.The

strategywillconsidertreatingthevariouspopulationsasalarger

metapopulation,lookingatwaystocreateandmaintaingene

flow,aswellasprotectingprivateallelesamongthesesmall

populationsbyimprovinggeneticmanagementstrategies.This

frameworkalsocommittedUSDOIagenciestoexpandingherd

sizeifpossible,andbuildingcooperationwithpartnersforthe

conservationofbison.Inaddition,comprehensiverefugeplans

arebeingreviewedtoconsiderthefeasibilityofattemptingbison

restorationonlargerefugelandscapes,suchastheCharlesM.

RussellNationalWildlifeRefuge.

UtahjustcompletedareintroductionofbisonintotheBookCliffs

areaofEastCentralUtah.ThisisajointeffortbetweentheState

ofUtahDepartmentofWildlifeResourcesandtheUteIndian

Tribe.BisonweremovedontothislandfromtheUtetribalbison

herdandtheHenryMountains.Thesebisonarelegallyclassified

aswildlifeandwillbemanagedasavaluedwildliferesourcein

Utah.Aherdmanagementplanhasbeenapprovedwherehunting

programmeswillregulatebisonpopulationsizeanddistribution.

PublicinterestinwoodbisonrestorationinAlaskahasgrown,

andthereiswidespreadstate,national,andinternationalsupport

forrestoringoneormorepopulationsinthestate.Thereisalso

supportamonglocalcommunitiesintheareasbeingconsidered

forwoodbisonrestoration.AWoodBisonRestorationAdvisory

Groupcomprisedofrepresentativesofvariousstateandnational

interestshasrecommendedthatAlaskapursuethereintroduction

ofwoodbisonatthethreesites,whichincludetheMintoFlats,

YukonFlats,andlowerInnoko/YukonRiverareasininteriorand

westernAlaska.Theseareashavesufficienthabitattosupport

from500to2,000ormorebisoneach,dependingonthelocation.

In2008,woodbisonweretransportedfromElkIslandNational

Park(EINP)toatemporaryholdingfacilityinAlaska,where

theyarebeingquarantinedfor2yearspriortoreleaseinthewild.


AlaskaDepartmentofFishandGameandUSFWSare

developingaspecialrulethatwilldesignatewoodbisonin

Alaskaasanonessentialexperimentalpopulation(NEP)under

section10(j)oftheU.S.EndangeredSpeciesAct,whichlists

woodbisonasendangered.Thefederalrulewillremove

manyoftheregulatoryrequirementsthatnormallyapplyto

endangeredspecies,allowingahighdegreeofmanagement

flexibilityandprovidingprotectionagainstpossibleregulatory

burdensandeffectsonotherlanduses.NEPstatuswill

helpmaintainandenhancepublicsupportforwoodbison

restoration.Analternativeproposaltodelistbisonfromthe

ESAisbeingconsidered,whichwouldobviateconcernsby

theoilandgassectoraboutimpactsofanewlistedspecies

ondevelopmentopportunities.WoodbisoninAlaskawillbe

legallyclassifiedaswildlifeand,afterpopulationsreachlevels

thatcansupportasustainableharvest,theirnumberswillbe

regulatedinpartthroughahuntingprogrammeasoutlinedin

cooperativemanagementplansthatwillbedevelopedforeach

areapriortoeachreintroduction.

8.5.5.2 Canada

TherehavebeenseveralCanadiannationalparkproposalsand

publicdiscussionstoincludeplainsbisonintheirnativespecies

managementplans.Theseincludemanagementplansfor

Banff,Waterton,andGrasslandsnationalparksinAlbertaand

Saskatchewan(Boyd2003;seealsoChapter7).Watertonpark

determinedthattherewasinsufficientlandscapeavailablefor

free-rangingbisonwithinthepark.PrinceAlbertandGrasslands

nationalparksalreadyhaveestablishedplainsbisonherds.

Bisonintheseherdsareclassifiedasfederallymanagedwildlife

andcouldbeallowedtoexpandtheirrangeifcoordinated

managementagreementscanbenegotiatedwithpublicand

privatelandownersborderingtheseparks.

Canadahasseverallargemilitaryreserveswithsuitablebison

habitat.Restorationonmilitarypreservesisbeingdiscussed,but

fewdetailedplansarecurrentlyavailable.Bisonareprotected

onDepartmentofNationalDefenceColdLake/PrimroseAir

WeaponsRangebyvirtueofprohibitingtrespass,exceptfor

theColdLakeFirstNations,whocanhuntwithpermission.

CanadianForcesBase(CBF)Suffieldisa2,600km2military

reservelocatedintheDryMixedGrassNaturalSub-region

ofAlberta.Itisusedasatrainingareaformilitaryground

manoeuvresanditisamostlyintactnativeprairielandscape.

CFB-Suffieldhasfree-rangingpopulationsofallindigenouslarge

herbivores,exceptbison,forwhichthebiologicalpotentialfor

restorationishighlyfavourable.

Canada’sNationalWoodBisonRecoveryTeamwasformed

in1973andincludesmembersfromallrelevantfederal,

provincial,andterritorialgovernments,aswellasacademia.

Thedraftnationalrecoverystrategy(H.Reynolds,personal

communication,1March2009)providesthefollowing

populationanddistributionobjectives:1)establishand

maintainatleastfivegeneticallydiversepopulationsofgreater

than1,000animalsineachherd,2)establishandmaintain

smallerfree-ranging,disease-freeherdswherepossible,and

3)establishandmaintainatleasttwopopulationsineach

originallyoccupiedecologicalregion.


speciesforeducationaloutreach.Anotherpotentiallyimportant

areafortherecoveryofbisoninMexicoistheColumbiaValley,in

theStateofCoahuila,whereaprivatelyownedherdmovesover

averylargeareaandisminimallymanaged.Bisonwerenativeto

thestateofCoahuilauntilthesecondhalfofthe19thCentury.

8.5.5.4 Non-governmental organisations

TNCandNCChaveplayedaleadroleinNorthAmericain

developingconservationprogrammesinvolvingbison.TNC

(eightherds)andNCC(oneherd)alreadymanageninebison

herdsongrasslandpreservesinU.S.andCanadarespectively.

TNCisprincipallyusingbisonasanativegrazerandis

consideringaddingbisontoadditionalpreservesintheU.S.

andCanada.Specifically,theNCCisimplementingarestoration

strategyfortheOldManonHisBackConservationAreain

Alberta,withaherdalreadyestablishedwithbisonfromEINP

(Freeseet al.2007).

In2005,APFandWWFimplementedaprivatelyfunded

conservationeffortrestoringbisontotheAmericanPrairie

ReserveinsouthernPhillipsCounty,Montana.Plainsbison

wereobtainedfromWindCaveNationalPark.UnderMontana

regulations,theyarecurrentlyclassifiedasprivatelyowned

livestock,however,theFish,WildlifeandParksCommission

hasauthority,underMontanalaw,toclassifythesebisonas

8.5.5.3 Mexico

Sincetheoriginalrangeofbisonextendedonlyashortdistance

intothenorthernportionofMexico,therearefewsuitable

locationswheretheywouldbeexpectedtosuccessfullyre-

coloniseavailablehabitatsintheirformerrange.Thelarge

grasslandsoftheJanos-CasasGrandeinnorth-westernMexico

isthebestlocationforbisonconservationefforts,andalarge

biospherereserveisproposedforthisareatoprotectfree-

rangingplainsbison.

Arecentseriesofstakeholderandscienceworkshopsheld

inthisboundaryareahaveidentifiedconservationneeds

andpotentialstrategiesforadvancingbisonrecoveryinthis

boundaryareaofMexico,includingreintroducingaplainsbison

conservationherdinMexico.InNovember2009,23plains

bisonweretranslocatedfromWindCaveNationalParkinSouth

DakotatoTNC’sRanchoElUnoEcologicalReservelocatedin

theJanosBiosphereReserveinChihuahuaState.Theproject

ispartofanationalprogrammeforrecoveryofpriorityspecies

inMexicoandaninternationalcollaborationonwildlifeand

habitatconservationinNorthAmerica.TheU.S.NationalPark

ServicedonatedthebisontoTheWorkingGroupforRecovery

ofBisoninMexico(ElGrupodeTrabajoparalaRecuperación

delBisonteenMéxico),whichisledbytheNationalCommission

ofProtectedNaturalAreas(laComisiónNacionaldeÁreas

NaturalesProtegidas).Thesebisonarethefoundationstock

forabreedingherdthatwillbeusedtorepopulateotherareas,

withtheultimategoalofrestoringtheecologicalroleofbison

inthegrasslandsofnorthernMexico.Thebisonwillprovide

opportunitiesforecologicalresearchandwillserveasafocal

Plate 8.1 Plains bison were reintroduced to the arid grasslands of the

Janos Valley in northern Chihuahua State, Mexico in November 2009.

The bison reside on Rancho El Uno Ecological Reserve, a property of The

Nature Conservancy. Photo: Rurik List.


wildlifeifAPFagreesandifthereispublicsupportforsuch

legalaction.APFintendstopurchaseupto405,000hectares

(onemillionacres)oflandforagrasslandpreserveupon

whichwildbisonwouldbeallowed.Inaddition,theAmerican

PrairieReserveleasesadjacentBLMgrazingallotmentsand

recentlymodifiedthesetochangetheclassoflivestockfor

theseallotmentsfromcattletobison.Similarly,theUSFWS

hasauthoritytoestablishbisonontheCharlesM.Russell

WildlifeRefugeadjacenttotheAmericanPrairieReserve.

Thecombinedeffortsofthesetwoagencies,andother

conservationpartners,couldresultinbisonrestorationona

verylargenativegrasslandhabitat.

8.5.5.5 tribal initiatives

ManytribalinitiativesarealsounderwayacrossNorthAmerica.

TheITBCwasformedin1990andhas57membertribes

managingover15,000bison(http://www.itbcbison.com/index.

php).ItsstatedgoalistorestorebisontoIndianNationsina

mannerthatiscompatiblewiththeirspiritualandculturalbeliefs

andpractices.Congressappropriatedfundingfortribalbison

programmesinJuneof1991,andhasapprovedappropriations

forITBCannuallysincethen.Thisactionofferedrenewedhope

thatthesacredrelationshipbetweenIndianpeopleandthe

“Buffalo”mightnotonlybesaved,butwould,intime,flourish.

SpecificinitiativesincludetheCheyenneRiverSiouxTribe,

whichhasstartedan8,900-hectareTribalWildlifeRefuge.The

RosebudSiouxTribehasofficiallyendorsed“TheMillionAcre

Project”developedbytheGreatPlainsRestorationCouncil

centredonthePineRidgeIndianReservationinSouthDakota

(Freeseet al. 2007).Anotherpotentialinitiativeisidentifiedina

strategicplanbeingdevelopedbytheLowerBruleSiouxTribe

inSouthDakota(LowerBruleSiouxTribe10yearstrategicplan;

LowerBruleSiouxDepartmentofWildlife,Fish,andRecreation).

TheWindRiverReservationinWyomingisworkingona

managementplanthatwouldrestorewildfree-rangingbison

toavailablehabitatonthattriballandscape.TheFortBelknap

ReservationinMontanahasrequestedYellowstonebisonfrom

thestate/federalquarantinefacility.Acomprehensiveevaluation

oftherestorationpotentialofNorthAmericantribal/firstnation’s

landscapesandcontinentalconservationpriorityassessments

forthoselandscapeshasnotbeencompleted.


Chapter9 ConservationGuidelinesforPopulation,Genetic,and

DiseaseManagementLead Authors: John E. Gross, Natalie D. Halbert, and James N. Derr

Contributors: Keith Aune, Joel Berger, Brett T. Elkin, C. Cormack Gates, Peter J.P. Gogan, David

Hunter, Damien O. Joly, Duane J. Lammers, Nicholas C. Larter, Daniel Licht, Rurik List, Robert L.

Paulson, Jenny Powers, Robert O. Stephenson, Joe Truett, Rick Wallen, and Margaret Wild

9.1 Introduction and principles

Thischapterprovidesmanagementandpolicy-relevant

guidelinestofosterbisonconservationandfullrecovery.

Conservationimpliesretainingdesirableecological,cultural,and

geneticcharacteristicsthatcurrentlyexist,whilefullrecovery

impliesabroadervision—bisonpopulationsinhabitingareas

thatpermitfullexpressionofnaturalbehavioursandecosystems

functioninginwayssimilartothoseofthepast.

Wefocusonguidelinesandprinciplesthatarebroadly

applicable,andweavoidedhighlyspecific,prescriptive

recommendations.Thisapproachrequiresmanagersandothers

tounderstandthebasisforourguidelines,andtoevaluate

carefullyhowaguidelinecanbestbeimplementedinaparticular

situation.Weprovideonlybriefreviewsofthescientificbasisfor

guidelines,andreadersshouldrefertochaptersfour,five,and

sixinthisvolumeformorecomprehensiveinformationonbison

genetics,disease,andecology.

Asmallsetofoverarchingprinciplesisthefoundationformost

oftheguidelinesinthischapter,andtheyprovideaframework

fordevelopingandassessingconservationactions.Thesekey

principlesare:

1) Maximizethenumberofbisoninapopulation.Larger

populationsbetterretainnaturalvariation,andaremore

resilientto‘surprises’orcatastrophicevents.Striveto

achievea‘maximumsustainable’ratherthana‘minimum

viable’populationsize.

2) Supportandpromote‘wild’conditionsandbehaviours.

Wherepossible,provideanenvironmentwherebison

areintegraltocommunityandecosystemprocesses

(Table9.1).Behavioursanddemographicprocesses

shouldreflectnaturalselection,andactivemanagement

interventionsshouldbeminimized.Wildbisonherdsuse

verylargeranges.

plate 9.1 The bison is an interactive species. Here wolves are hunting and

feeding on a plains bison they have killed and ravens are scavenging (middle

photo). Top and middle photos: Douglas Smith, lower photo: Dwight Lutsey.


3) Preservegeneticintegrityandhealth.Maintainbison

lineagesandcarefullyevaluateallmovementsofbison

betweenpopulations.Considerpotentialgenetic

consequencesofallmanagementactions,especiallyfor

smallherds.

4) Routineassessmentiscentraltoscience-based

conservationofbison.Routinemonitoringandevaluation

ofdemographicprocesses,herdcomposition,habitat,and

associatedecologicalprocessesarecentraltoevaluating

herdhealthandmanagementefficacy.Assessmentsare

necessarytoanticipateorrespondtoconservationneeds

andsounddataisthebasisforinformedmanagement.

Thescientificbasisandrationalofprinciplesforconserving

bisonisprovidedinthemoredetailedguidelinesinthischapter

andotherchaptersthatreviewbisonecology,genetics,and

ecologicalrestoration.

9.2 Guidelines for population and Genetic Management

Thegeneralgoalsforpopulationandgeneticmanagementareto

achieveandsustainapopulationwithahealthylevelofgenetic

variationandasexandagecompositiontypicalofviablewild

bisonpopulations.Managementactionsneededtoachieve

thesegoalswillvarywiththesize,history,andcircumstances

ofeachparticularpopulation.Inthissection,wearticulate

morespecificmanagementobjectives,summarisebackground

informationrelevanttoourrecommendations(seealsoChapter

6),andprovidebothgeneralandspecificguidelines.

Inbison,lossofgeneticvariationisaconcernprimarilywhenthe

numberofactivelybreedinganimalsorthefoundingpopulation

sizeissmall.Ourbestestimatesarethatbisonpopulations

cangenerallybeconsidered“notsmall”(forgeneticpurposes)

whentheyexceedabout1,000animals,thepopulationhas

approximatelyequalnumbersofbullsandcows,andthesize

ofthepopulationisstable.Forthepurposesofthisreport,

thegeneticobjectiveistoattaina90%probabilityofretaining

90%ofselectivelyneutralgeneticvariationfor200years.This

objectiveislessstringentthansomepublishedobjectives,

andthusourestimatesforsustainablepopulationsizesare

smallerthanthosethatresultfromestimatesbasedonmore

conservativecriteria(Reedet al.2003;Souleet al.1986).Inall

populations,therateoflossofgeneticdiversityisdirectlyrelated

tohowrapidlyindividualsinapopulationreplacethemselves

(generationtime)andtothesizeofthebreedingpopulation.

Mostguidelinesforgeneticmanagementinthisdocumentcan

beunderstoodinthecontextofjustthesetwofactors.

Mostpopulationsarenotuniform,buthavegeneticvariation

relatedtothespatialsubstructureofthepopulation(Manelet

al.2003).Demographicandgeneticsubstructureoccursat

alargegeographicalscaleduetotraditionaluseofparticular

partsofarange(e.g.,breedingrangefidelity,seasonalranges,

calvingareas)bysegmentsofapopulation(e.g.,bisoninYNP;

Christiansonet al.2005;Gardipee2007;Goganet al.2005;

Halbert2003;OlexaandGogan2007).Withinherds,bisonare

thoughttoformfamilygroups(i.e.,matrilinealgroups,mother

cowswiththeirpreparturientdaughters)andthesefamily

groupsconstitutefine-scalepopulationstructuring.Thesetypes

ofpopulationstructureareimportantbecausetheyincrease

thelikelihoodthatanimalremovalswithoutplanstoexplicitly

accommodatesubstructuresofcowscoulddisproportionately

impactaparticularsegmentofthepopulationandresultin

agreaterlossofgeneticdiversitythannecessary.Removal

strategiesshouldbedesignedtoaccommodatethepotential

spatialstructureofherds,andinstituteproceduresthatensure

process Description

Createpatches Grazingcanproduceadynamicmosaicofvegetationpatchesthatdifferinseralstageandthatdifferduetovariationsingrazingintensity

Enhancenutrientcyclingrates

Bisongrazingcanenhancenutrientturnoverandchangedominantsystemmodefromdetritus-decompositiontoconsumption-defecation

Enhancehabitatquality

Bisongrazingcanincreasehabitatsuitabilityforprairiedogs,pronghorn,andotherspecies

Modifyfireregimes Bisonconsumefinefuelsandcreatetrailsandtrampledareasthatreducefireintensityandextent,andmodifytheeffectoffireonvegetationheterogeneity

Createdisturbances Tramplingandwallowscreateseedbedsforsomespecies;localisedtreestandsthatarenottightlyclumpedaresusceptibletomajordamagebyrubbing,horning,andthrashingofbison.

Stimulateprimaryproduction

Bisongrazingremovessenescentmaterialfromtheswardandincreaseslightpenetration,nutrientavailability,andgrowth

Disperseplantseeds Bisontransportseedsinlegfurandgut,andmayenhanceestablishment(ofnativeandexoticplants)viaconsumption,seedcoatdigestion,anddefectioninnutrient-richmedia.

Maintainfloraldiversity

Bisongrazingcanresultingreatergrassandforbspeciesdiversity

Supportcarnivoresandscavengers

Bisonarepreytosomelargecarnivores,andbisoncarcassescancontributetosupportingscavengers.

table 9.1 Ecosystem processes that bison can strongly influence. See

Hobbs (1996); Knapp et al. (1999); Larter and Allaire (2007); and Truett

et al. (2001).


animalsareproportionatelyremovedfromdifferentpopulation

segments.Thiscouldpotentiallybeaccomplishedbyremoving

animalsfromdifferentpartsoftherange.

Avarietyoffactorscanleadtoincreasedratesofgenetic

diversityloss.Afteraccountingforpopulationsize,themost

importantfactorsarelikelytobenon-randommating(i.e.,afew

bullsareresponsibleforsiringmostcalves),skewedsexratios,

andlargevariationinpopulationsize.

9.2.1 Guidelines that apply to most conservation herds

Veryfewconservationherdswillpersistwithouttheneedfor

someformofpopulationcontrol.Manyguidelinesinthis

chapterwereincludedwiththespecificintenttosupport

developmentofinformedpopulationmanagementplans.Many

ofthefollowingguidelinesapplytomostconservationherds,

andarelikelytobeincludedincomprehensivemanagement

plansforconservationherds:

1) Maintainasexratiowithneithersexconstitutingmore

than60%ofthepopulation.Ideally,theadultsexratiowill

beslightlyfemalebiased(e.g.,55cowsper100animals),

reflectingobservationsthatmortalityratesofmalestend

tobeslightlygreaterthanthoseforfemales.Avoidinga

highratiooffemalestomaleshelpsensureparticipationin

matingandtransferofgeneticdiversitybyalargernumber

ofbulls.Inlargepopulations,matingcompetitionwilllikely

besufficientwhenthereare20ormorematurebulls(six

yearsoldandolder)per100cows.Maintainingmating

behaviour,asnotedabove,callsforamoreequalsexratio.

2) Avoidremovingasignificantproportionofthepopulation.

Forpopulationssubjectedtopopulationcontrolactions,

cullingshouldbeonayearly,oreveryotheryear,schedule,

ratherthanperiodicallyatlongerintervals.Wecannot

offeradefinitivedefinitionof‘significant’,astheeffects

ofpopulationfluctuationswillbegreateraspopulation

sizediminishesandvarieswithothercircumstances.Asa

generalguideline,wesuggestlimitingremovalsofanimals

tolessthan30%ofthepopulation;

3) Avoiddisproportionateremovalofmatrilinealfemale

groups(mothercowsandtheirpreparturientdaughters).

Morespecifically,attempttoretaintheoldercows

matrilinealgroups;

4) Removeanimalsfromallspatialsegmentsofthe

population;

5) Emulatenaturalmortalitypatterns—highermortality/

removalratesforjuvenilesandoldageclasses(morethan

15years);

6) Insmallpopulations,consideractionsthatreducevariation

inthebreedingsuccessamongindividuals.Thiscouldbe

accomplishedbyreducingtheopportunitiesforcontinued

breedingbyhighlysuccessfulbulls.

7) Avoidhumanselectionformarkettraitssuchasdocility,

carcasscomposition,bodyshape,orproductivity,assuch

interventionscontradictnaturalselectionandconservation

ofgeneticvariability;

8) Routinesupplementalfeedingtoincreaseproductivity,or

tosupportapopulationsizethatexceedsrangecarrying

capacity,isdiscouragedforconservationherds;

9) Wherepractical,thefullsuiteofnaturallimitingfactors

shouldbeallowedtoinfluencepopulations,including

winterdeprivationandpredation.Thiswillresultinvariable

ratesofreproductionandsurvival.

Theneedforactivegeneticmanagementwillvarywithherd

size,geneticcomposition,andmanagementgoals.Ingeneral,

geneticallydiverseherdswithmorethan1,000animalsare

unlikelytorequireactivemanagementtoretainmostoftheir

geneticdiversityforthenext200years(Grosset al.2006).

Hedrick(2009)suggestsaherdsizeof2,000-3,000toavoid

inbreedingdepression.Inverysmallherds(fewerthanabout

250animals),long-termgenetichealthwillrequireoccasional

supplementationwithgeneticmaterialfromotherherds.The

exactnumberofanimalsneededtosupplementaparticularherd

willvarywiththegeneticcompositionofthesourceandtarget

herds,butasupplementoffourtofivebreedinganimalsper

decadeshouldbesufficientforlong-termherdgenetichealth

(Wang2004).Inadditiontotheguidelinesbelow,managers

shouldfollowtheIUCNguidelinesfortranslocationofwild

animalsbetweenestablishedherds,beingespeciallycareful

aboutgeneticpurity(i.e.,cattlegenesandgeographically

appropriatesourcesofstock)anddiseases(http://www.kew.

org/conservation/RSGguidelines.html).

Activemanagementtoretaingeneticvariation(otherthan

translocations)maybemostimportantforintermediate-sized

populationswithabout250-750animalsbecausethisisthe

sizerangewhereactivemanagementmaypreventorgreatly

reducetheneedfortranslocatinganimalstoensurelong-term

thegenetichealthofaherd(Grosset al.2006).Forconservation

herds,theoverallobjectiveistoretainallelicdiversity,which

isthebestindicatorofthegeneticresourcesavailabletothe

population.Bycontrast,geneticheterozygositymaybeabetter

short-termindicatorofthematingstructureoftheherd.In

additiontotheguidelinesprovidedabove,removalofyoung

animals,priortotheirfirstbreeding,cansignificantlyenhance

theretentionofgeneticdiversity(Grosset al.2006).Removal

ofyounganimalstopreservegeneticdiversitymayseem

counterintuitive.Geneticmaterialislostonlywhenanimalsina

populationarereplaced.Removalofyounganimalsincreases

thelengthofthegeneration(replacement)interval,andthis

therebyprolongstheretentionofgeneticmaterial.


9.2.2 herd-level population and genetic management

Formanyconservationherds,themostfrequentandcontentious

decisionswillconcernherdlevelmanagement,especially

populationcontrol.Keydecisionsaddresshowmanyanimalsto

maintain,whichonestoremove,andhowoftentoremovethem,

whentoaddanimals,andwheretosourcethem.Thissection

providesadviceforactivepopulationmanagementattheherd

level—guidelinesforestablishinganewherd,maintainingthesize

ofanexistingherd,reducingthesizeofaherdthathasbecome

muchtoolarge,andhowtodealwithknowngeneticissues.

9.2.2.1 Soft release procedures

Bisonmayneedtobemovedtosupplementanexistingherd,

ortoestablishanewherd.Insuchcases,theuseofa“soft”

releaseprocessshouldbeconsideredinvirtuallyallcases.

Softreleasestypicallyinvolveplacinganimalsina(usually

large)holdingfacilitypriortofullrelease.Holdingbisonina

largepenmayincreasetheirtendencytoremainintheareaof

releaseandestablishsomedegreeofsitefidelity.American

PrairieFoundation,forexample,heldbisonforonemonthina

largecorralpriortoreleaseontheAmericanPrairieReservein

Montana.Anadditionalbenefitofasoftreleaseprocedureisthe

effectivequarantineandassociatedabilitytomonitorandmore

easilyre-captureanimalsifanyhealthissuesbecomeapparent.

9.2.3 Establishing a new herd

Establishingandmaintainingrelated,isolatedorsemi-isolated

herds(i.e.,parentalandoneormoresatelliteherds)iscritical

tolong-termspeciesconservationinthatmultipleherdsactto

increaseeffectivepopulationsize(Ne)andreducethetotalloss

ofgeneticvariationovertime(LandeandBarrowclough1987).

Furthermore,themaintenanceofauniquegeneticpopulation

inseveralsmallherdsreducestheprobabilityofaccidental

extinction,suchasfromanaturalcatastrophebydisease,and

increasestheopportunityforlocaladaptation(Franklin1980;

Lacy1987).Intheory,andunderexperimentalconditions,

severalsmallgroups(e.g.,Neabout50)maypreservemore

geneticdiversitythanasingleherdwithasmanyindividualsas

thesmallerherdscombined(Marganet al.1998).Geneticdrift

withineachrelatedherdcanbecounteredbytheoccasional

movementofindividualsbetweenrelatedherds(Millsand

Allendorf1996).Therefore,severalmoderatelysizedherds

(i.e.,morethan300andfewerthan1,000animals)ofthe

samegeneticstockcan,ifmanagedproperly,actasalarge

metapopulationwithaneffectivepopulationsizesufficientto

impedegeneticerosion(Lacy1987).Inthissection,wearticulate

considerationsfortheestablishmentandmaintenanceofnew

bisonherdsfromexistingresources.

1. Source

Priorityshouldbegiventoestablishingsatelliteherdsfrom

extantconservationherds,withintherespectiveoriginalranges

forwoodandplainsbison,especiallyforthoseherdswithunique

geneticcharacteristics(Halbert2003;WilsonandStrobeck

1999)andthosewhichappeartobefreeofdomesticcattle

introgression(Wardet al.1999;Halbert2003;Halbertet al.

2005b).Beyondthis,establishmentofherdsofmixedancestry

shouldbeconsideredtomaximisegeneticdiversityandthe

potentialforadaptiveresponse.

Althoughbisonarelikelytobemorereadilyavailablefromherds

subjectedtoartificialselectionandsomelevelofdomestication,

westronglyrecommendacquiringbisonfrom“wild”herdsnot

subjectedtotheseinfluences.

2. Number of animals

Littlespecificinformationisavailableregardingappropriate

foundationpopulationssizes.Ingeneral,afew(4-10)individuals

shouldbesufficienttoavoidveryshort-terminbreedingeffects

(Senner1980).However,thelossofvariationinsuchasmall

populationwillbesubstantialafterthefirstfewyears(Neiet al.

1975)andadditionalbisonshouldbeimportedoveraperiod

ofseveralyearstoincreasegeneticvariation.Ifthegoalisto

conserveorduplicatemostofthegeneticmaterialinasource

herd,manymoreanimalsarerequired.Shuryet al.(2006)

proposedabaseof200“founder”animalstopreservemostof

thegeneticvariabilityin“re-established”woodbisonherds.

3. Sex ratio

Theinitialimportedbisonshouldconsistofapproximately50%of

eachsex,andtheherdshouldbemaintainedwithabalancedsex

ratiotoreduceinbreedingandmaximiseeffectivepopulationsize.

4. Breeding strategy

Ifasmallnumberofbisonareusedtofoundaherd,and

especiallyifadditionalbisonarenotbroughtintothenewherd,

breedingstrategiestomaximisethetransferofgeneticdiversity

acrossgenerationsshouldbeconsidered(e.g.,avoidexcessive

breedingbyoneorafewmales).Appropriategenetictoolsare

availabletoaccuratelyassignparentageinbison(Schnabelet

al.2000;Wilsonet al.2002),andthesemaybeusedtoassistin

captivebreedingdecisionsbyevaluatingthebreedingsuccess

ofindividualbullsandrelatednessamongcalves.

5. Age composition and behaviour

Bisonaresocialanimalsandtheimportanceofsocialstructure

withinaherdiscriticaltooverallherdhealthandsurvival

(McHugh1958).Werecommendestablishinganewherdwith

bothadultandsub-adultindividualstopreventdisintegration

ofsocialstructureandbehaviouralanomalies(e.g.,foraging

behaviour;RalphsandProvenza1999).


6. Maintenance number and growth rate

Tominimisethelossofgeneticvariationandheterozygosity,and

tomaximisetheprobabilityofpopulationsurvival,newherds

shouldbeallowedtogrowasquicklyaspossibleuntilthetarget

herdsizeisattained(Neiet al.1975).Bisonherdscangrowvery

quickly,doublinginsizeinasfewasfouryears(seeChapter

6).Herdsshouldthenbemaintainedwithinanappropriatesize

range,whichwilllikelybethemaximumsizepossiblewithin

resourcelimitsforherdswithfewerthanabout1,000animals

(Grosset al.2006;Senner1980).Forsmallherds,fluctuations

inpopulationsizecanhaveasubstantialnegativeimpacton

retentionofgeneticvariation(Neiet al.1975).Maintenanceof

populationsizeismoreimportanttopopulationsurvivalthanis

thefounderpopulationsizeandshould,therefore,begivena

highpriorityforsmallherds(Senner1980).

7. relationship between founders

Selectunrelatedindividualsasfoundersforanewherd.Use

appropriategenetictoolswhenavailabletoestablishrelatedness

betweenbison(Schnabelet al.2000;Wilsonet al.2002).

8. Genetic variation and heterozygosity

Geneticevaluationshouldbecarriedoutontheparentalherd

priortoestablishmentofasatelliteherd,andrepeatedgenetic

evaluationofthesatelliteherdshouldbeusedtoensurethatall

thegeneticvariationfromtheparentalherdareincorporatedand

maintained.

9. Disease

Ingeneral,donotusediseasedbisontoestablishanewherd.

Immunesuppressionindiseasedindividualsmayleadto

infectionandspreadofotherdiseases;furthercompromising

herdestablishmentandhealth.Onenotableexceptionis

theintentionalcreationofdisease-freesatelliteherdsfroma

diseasedparentalherd.Insuchcases,useextraprecautionsto

preventthespreadofdiseasefrombisontootherwildlifeduring

theinitialdiseaseeliminationphase.

10. Monitoring success

Becauseitisexpensiveandtime-consumingtoestablishbison

herds,resourcesshouldbewiselyinvestedtomonitorbison

herdsandbroaderecologicaleffectsofbison.Ideally,habitat

characteristicsshouldbemonitoredusingavalidstatistical

processbeforebisonareintroduced.Herdcomposition,

demographicparameters,andgeneticstructure,especiallyin

thefirstfewgenerationsfollowingherdestablishment,shouldbe

monitored,alongwithecosystemchanges.Additionalmonitoring

guidelinesareprovidedbelow.

Trans-boundarytransportationofbisontoestablishanew

herdcanintroducemanyadministrativeandregulatory

considerations(Chapter8).Afteranextendedperiodofplanning

andnegotiation,woodbisonweretransportedfromCanadato

Alaskain2008.Personnelwiththerelevantagenciesmaybe

consultedforadviceonundertakingsuchanenterprise.

9.2.4 Maintaining or manipulating existing herd size

Whenabisonherdappearsinneedofinterventiontorestore

orimprovegenetichealthandpopulationviability,thefirstand

mostimportantactivityistothoroughlyevaluatethecurrent

conditionoftheherdtoavoidpremature,unnecessary,or

evendamagingmanagementdecisions.There are no simple

cookbook instructions that can be applied to any bison herd.

Thefollowinglistofbaselineevaluationswillhelpensurethat

decisionsarewellinformed:

1.Determinethehistoryoftheherdtoprovideinsightinto

currentlevelsofgeneticvariationandpopulationstructure.Tryto

determine:

• Numberandoriginofherdfounders;

• Numberandoriginofanybisonintroducedfollowingherdfoundation(transfers);

• Historicrecordsonpopulationsize,especiallywithregardtosubstantialchangesovertime.

2.Evaluatecurrentpopulationparameterstoestablishbaseline

measurementsforfuturecomparisonandtodetectattributes

thatmayleadtochangesinsocialstructureorgeneticvariation.

Variablesofinterestinclude:

• Censuspopulationsize

• Effectivepopulationsize(Ne;willnotbepossibleinallcasesandrequiresknowledgeofbreedingstructure)

• Rateanddirectionofpopulationsizechanges(e.g.,istheherdexpandingorcontracting)

• Sexratio

• Agestructure

3.Noteanyindicationsofinbreedingwithintheherd,suchas:

• Unusualphenotypiccharacterswithintheherd,especiallyanythathaverecentlyappeared;

• Recentdecreaseinrecruitmentrates;

• Highratesofmorphologicallyabnormalornon-motilespermamongbreeding-agebulls;

• Relativelylowlevelsofheterozygosityascomparedwithpreviousmeasurementsorotherbisonherdsofsimilarsizeandhistory(e.g.,Halbert2003;WilsonandStrobeck1999).

4.Assesspotentialhealthproblemsintheherd,including:

• Presenceoftransmissiblediseases,especiallythosewhichmayinfluencepopulationdynamics(e.g.,BTB,brucellosis,MCF);

• Presenceofdiseaseagentsinlivestockspeciesonnearby(especiallyadjacent)properties(e.g.,cattlewithJD,sheepcarryingMCF).


5.Evaluatetheoverallgeneticconstitutionoftheherdby

measuring:

• Uniquevariation(rareorprivatealleles)andlevelsofheterozygosityincomparisontootherbisonherds(Halbert2003;Halbertet al.2004;WilsonandStrobeck1999);

• Within-herdchangesinheterozygosityandgeneticvariationbetweengenerations(Halbertet al.2004);

• Currentbreedingstructureoftheherd(e.g.numberofmalescontributingtocalfcropeachyear,relatednessamongcalves,presenceofgeneticsubpopulations);

• Existinglevelsofdomesticcattleintrogressioninboththemitochondrial(Polziehnet al.1995;Wardet al.1999)andnucleargenomes(Halbertet al.2005b).

Usingthedatacollectedfromtheaboveevaluations,informed

andsensiblemanagementplanscanbeimplementedtobest

fittheneedsofthetargetherd.Tofurtherassistinthisprocess,

demographicandgeneticdatacanbeusedtomodelthe

effectsofvariousmanagementalternativespriortoactually

implementingadefinitivemanagementplan(Grosset al.2006;

Halbertet al.2005a).

9.2.5 transferring bison between herds

Tomaintainlong-termherdhealth,itwillbenecessaryinsome

casestotransferbisonbetweenherds(Table9.2).Thedecision

totransferbisonbetweenherds,however,mustbemadewith

extremecautionwiththefollowingconsiderations:

1. Necessity of movement

Isthereactualevidenceoflossofgeneticdiversityorinbreeding

tonecessitatethetransfer?Inbisonandothermammalian

species,wellintendedbutuninformedmanagementdecisions

totransferindividualsamongisolatedgroupshaveresultedin

detrimentalandirreversibleeffects,especiallyrelatedtogenetic

integrityanddisease.

2. Domestic cattle introgression

AsdiscussedinChapter4,fewbisonherdsappeartobefree

fromdomesticcattleintrogression(Halbert2003;Halbertet al.

2005b;Polziehnet al.1995;Wardet al. 1999).Therefore,itis

essentialtounderstandboththehistoricandgeneticevidenceof

domesticcattleintrogressionintherecipientandpotentialdonor

herdsbeforeconsideringatransfer.Ifthetwoherdsarerelated,

andespeciallyifoneisasatelliteoftheother,thetotaleffecton

introgressionlevelsduetotransferwillbenegligible.Careshould

betakentopreventtheintroductionofbisonofunknownorigin,

orquestionablehistory,intoconservationherds.Furthermore,

givenourcurrentlevelsofunderstanding,bisonshouldnotbe

transferredintothefewexistingherdswhichappeartocontain

nodomesticcattleintrogression,withthepossibleexceptionof

transfersbetweenparentalandsatelliteherds(Hedrick2009).

3. relationship between herds

Giventheobservedgeneticdistinctionsamongextantbison

herds(Halbert2003;WilsonandStrobeck1999),dilutionof

uniquegeneticcharacters(alleles)withintherecipientherd

shouldbeconsideredwhenevaluatingpotentialdonorherds

(Halbertet al.2005a).Ideally,bisonshouldbetransferred

betweensatelliteorrelatedherdstoreducethelossofrare

variants.

Number Whenpossible,thenumberofimportedbisonshouldbebasedonpriormodellingestimatestomaximizeimprovementsinheterozygosityandgeneticdiversitywhileminimizingdilutionofthenativebisongermplasm.

Sex Importingafewnewmalesintoaherdcanhavealarge,positiveandrapidgeneticanddemographicimpact.Thesameoveralleffectscanbeobtainedwhenimportingfemales,althoughtheprocesswillbesomewhatslower.Insomecases,itmayalsobeworthwhiletoconsideranyknowngeneticuniquenessofthemitochondrialgenomeandYchromosome.Forinstance,priortoimportingbisonintotheTexasStatebisonherd,itwasnotedthatthisherdcontainedauniquebisonmitochondrialhaplotypenotknowntooccurinotherbisonherds(Ward2000;Wardetal.1999).Therefore,importingmalesintothisherdwasfavouredoverimportingfemales,inparttopreventdilutionoftheuniquenativebisonmitochondrialhaplotype(Halbertetal.2005a).

Age Themostrapidinfusionofgermplasmwillbeobtainedbyimportingbreeding-ageanimals.Itmaybedesirabletochoosebisonthathavealreadyproducedoffspringtoavoidpotentialissuesofsterilityoroffspringabnormalities.Despiteplanning,geneticincompatibilitiesbetweenextantandimportedbisonmaystillinfluencecontributionsoftheimportedbisontothecalfcrop.

Quarantine Consideraquarantineofnewlyimportedbisonpriortorelease,especiallywhentherecipientherdisatahighriskofextinction.Thisallowsforaneasieradjustmentoftheimportedbisontotheirnewenvironment,aswellasearlydetectionandtreatment/removalforlatentdiseases.

Matingregime

Decidewhetherimportedbisonshouldhaveexclusivematingprivilegesforoneormoreyearsorcompetewithotherpotentialbreedersforaccesstocows.“Exclusive”matingscanbeusedtoincreasegeneticanddemographicimpacts.Afullycompetitivematingregimenpermitsextantbisontocontributetothegenepoolandprovidessomeprotectionincaseofgeneticincompatibilitybetweenthedonorandrecipientherds.

table 9.2 Additional factors to be evaluated when considering transfers

of bison between herds.


4. health and disease

Allattemptsshouldbemadetopreventthespreadofdisease

betweenbisonherds.Eveniftherecipientanddonorherdshost

thesamedisease,transfersofbisonshouldbediscouraged

sincediseasestrainvariantsbetweenherdscanleadto

differencesindiseaseprogressionoreffects.Potentialdonor

herdsshouldbethoroughlytested(seeChapter5andsection

above)toevaluatethepresenceofpathogens.

Oncetheabovefactorshavebeenevaluated,therearevarious

otherfeaturesthatmayinfluencethedemographicandgenetic

effectsofthetransfer,includingthenumber,age,andsexof

theimportedbisonaswellasfrequency(singleormultiple

introductions)anddurationofthetransfers(permanentvs.

transienttransfers,e.g.,forshort-termbreeding).Eachsituation

willdifferandacomprehensivereviewisnotpossibleheregiven

thelargenumberofpotentialmanagementscenarios.However,

thegeneralguidelinesinTable9.3shouldbeconsidered.

6. Sex

Importingafewmalesintoaherdcanhavealargeandrapid

geneticanddemographicimpact.Thesameoveralleffectscan

beobtainedwhenimportingfemales,thoughtheprocesswill

besomewhatslower.Insomecases,itmayalsobeworthwhile

toconsideranyknowngeneticuniquenessofthemitochondrial

genomeandYchromosome.Forinstance,priortoimporting

bisonintotheTexasStatebisonherd,itwasnotedthatthisherd

containedauniquebisonmitochondrialhaplotypenotknown

tooccurinotherbisonherds(Wardetal1999;Ward2000).

Therefore,importingmalesintothisherdwasfavoredover

importingfemales,inparttopreventdilutionoftheuniquenative

bisonmitochondrialhaplotype(Halbertetal.2005a).

7. Age

Clearlythemostrapidinfusionofgermplasmandimprovement

inherdviabilitywillbeobtainedbyimportingbreeding-age

animals.Insomecases,itmayalsobedesirabletochoosebison

thathavealreadyproducedoffspringtoavoidpotentialissues

ofsterilityoroffspringabnormalities.Evengiventhemostwell

thought-outplans,however,geneticincompatibilitiesbetween

nativeandimportedbisonmaystillinfluencetheeffectivenessof

theimportedbisonincontributingtothecalfcrop.

8. Quarantine

Aquarantineofnewlyimportedbisonshouldbeconsideredprior

totheirrelease,especiallywhentherecipientherdisatahigh

riskofextinction.Isolatingthenewlyimportedbisonforsome

timewillallowforaneasieradjustmentoftheimportedbison

totheirnewenvironmentandearlydetectionandtreatment/

removaloflatentdiseases.

9. Mating regime

Shouldtheimportedbisonhaveexclusivematingprivileges

foroneormoreyearsorshouldtheybeincludedwithall

otherpotentialbreederstocompeteforbreedingrights?An

“exclusive”matingregimenallowsforlargerpotentialgenetic

anddemographicimpacts.However,a“competitive”mating

regimenpermitsnativebisontocontinuetocontributetothe

genepooleachyearandprovidessomeprotectionincaseof

geneticincompatibilitybetweenthedonorandrecipientherds.

9.2.6 recovering small or threatened herds

Smallpopulations(Nelessthan50,oracensussizeoffewer

thanabout150animals),orlargerpopulationswhichhave

undergonearecentandsignificantdecreaseinpopulation

size,areespeciallyvulnerabletoalossofgeneticvariation,

decreasedfitness,and,ultimately,extinction(GilpinandSoulé

1986).Persistentlysmallpopulationsareadditionallysusceptible

toinbreeding,whichcanleadtoanoveralllossofheterozygosity

andincreaseinrare,andoftendetrimental,genetictraits.

Disease risk Factors Disease Examples (not all-inclusive)

Historyofpathogenintheregion

Anthrax,parasites

Proximitytopotentiallyinfectedpopulations(wildlifeorlivestock)

MCF,Bovinetuberculosis,brucellosis,Johne’sdisease,bovineviraldiarrhoea,foreignanimaldiseases(e.g.,Foot-and-MouthDisease)

Weatherpatternsandenvironmentalsuitability

Anthrax,parasites

Presence/abundanceofmechanicalorbiologicalvector(s)

Anaplasmosis,bluetongue,pinkeye

Populationdensity(increasedinfectiouscontacts)

Mostinfectiousdiseases(e.g.,brucellosis,tuberculosis)

Season Diseaseswithuniquetransmissionpatterns(e.g.,brucellosis,bluetongue)

Nutritionalandotherenvironmentalstress

Infectiousdiseaseswhichcapitaliseondepressedimmunity(e.g.,respiratoryviruses)

Geographiclocation/Climate

Hardypathogenscapableofsurvivingclimateextremes

table 9.3 Risk factors for disease.

5. Number

Thenumberofimportedbisonshouldbebasedonprior

modellingestimateswhenpossible,andshouldreflectthesize

ofthepopulationsothatimprovementsinheterozygosityand

geneticdiversityaremaximizedwithaminimumdilutionofthe

nativebisongermplasm.


Ifalargepopulationhasundergonearecentreduction

(=populationbottleneck)inashortperiodoftime(e.g.,fewer

thanthreegenerations),andisallowedtosubsequently

increaseinsizerapidlyandwithoutculling,theresulting

populationwillprobablysufferonlysmallreductionsinallelic

variationandheterozygosity(Neiet al.1975).Thesameisnot

trueofthebottleneckeffectinsmallpopulations,wherethe

lossofallelicvariationandheterozygositytendstobemuch

higher;inthiscase,extrameasuresmustbetakentomaximise

thetransferofgeneticdiversityandminimisethelossof

heterozygosityacrossgenerations.

Severalstrategiescanbeusedtoalterthebreedingstrategy

ofasmallherdtomaximiserecruitmentratesandgenetic

diversityinthecalfcrops.Forinstance,attemptscanbemadeto

randomisebreeding.Bisonarenaturallypolygamousbreeders,

anditmaybenecessaryordesirabletoimplementacontrolled

matingschemetoensurethatamaximumnumberofmales

arebreedingwiththeavailablefemales,andtomaximisethe

transmissionofgeneticvariationacrossgenerations.Ifsemen

viabilityorotherreproductivebarriersareanissue,artificial

inseminationmayalsobeconsidered.

Insomecases,alteringthebreedingstrategyofaherdmay

notbesufficienttoreversetheeffectsofsmallpopulationsize

(e.g.,Halbertet al.2005a).Inthesecases,itmaybenecessary

toimportbisonfromotherherdstoimproverecruitmentrates

andincreasegeneticvariation.Astheeffectsofimportingbison

intoasmallherdcanbeirreversibleandevendetrimental,the

ultimatedecisiontoimplementthisstrategyshouldbemade

onlyaftercarefulconsideration,andasalastresort(allissues

discussedinsection9.6.2shouldbeconsidered).Furthermore,

optionstomaximisedemographicandgeneticimpact(e.g.,

importingseveralmalesvs.afewfemales)shouldbeconsidered

inthreatenedherds.

9.2.7 recovering herds from germplasm introgression

Ifabisonherdhashadaninfluxofgermplasm(genetic

material)fromanoutsidesource,includinganotherbison

herdorarelatedbovidspecies,theabilitytorecoverthe

germplasmoftheoriginalherddependson:1)theabilityto

detectbisoncontainingintrogressedfragments,and2)the

numberofgenerationssincetheoriginalintrogressionevent.

Forinstance,iftwodistinctbisonherdsareaccidentallymixed,

parentagetestingwouldallowforpost-matingsegregation

ofthetwoherdsandtheiroffspringprovidedthatthebison

fromeachherdaredistinguishable(e.g.,identificationtags

orsufficientgeneticdifferences)andthatalimitednumberof

generationshavepassed(fewerthanthree).Ifmorethanafew

generationshaveelapsedsincetheinitialintrogressionevent,

theintrogressedsegmentswillbecomedispersedthroughout

thegenomeoftheherd(hybridswarm)andreconstitutionof

theoriginalgermplasmwillnotbepossible(Allendorfet al.

2001).Forexample,lowlevelsofdomesticcattleintrogression

havebeendetectedinmanyextantbisonherds(Halbert2003;

Halbertet al.2005b)andcanbetracedbacktohuman-induced

hybridisationofthetwospeciesover100yearsago;inthese

cases,multipledomesticcattlefragmentsaredispersedso

thoroughlythroughoutthegenomethatitisnotpossibleto

detect,muchlessremove,allintrogressedfragments.

9.2.8 herd size reduction

Bisonhaveahighintrinsicreproductiverateandbisonherds

generallygrowrapidly(seeChapter6).Therefore,when

resourcesarelimited,bisonherdsoftenexceedthecarrying

capacityoftheirenvironmentandbegintohavenegative

impactsonothergrazersandnativeplantspecies.Asa

result,mostbisonherdsaresubjectedtosomelevelofculling

(=periodicremovals)tomaintainasuitablepopulationsize

(Table9.4).Inextremecases,itmaybenecessarytoremove

alargeproportionofthepopulationtomeetmanagement

goals.Forexample,ifbisonhavenotbeenculledfromaherd

inseveralyears,theherdmayhavenearlydoubledinsize,and

itmaythreatenthesurvivalofotherspecies.Inthesecases,

extremecautionshouldbetakentoremovebisoninamanner

thatwillminimallyinfluenceherdandgermplasmcomposition

accordingtothefollowingguidelines.Somediscretionisneeded

inapplyingtheseguidelines.Forexample,itisimportantto

avoidsocialdisruptionwhilesimultaneouslyremovinganimals

fromallsegmentsofthepopulation.Managersmustcarefully

evaluatetheirgoalsandthespecificsituationtoachievethebest

outcome(Table9.4).

9.3 Behaviour: Mating System, Social Structure, and Movements

Bisonbehaviourisanindex,orreflection,oftheconditions

experiencedbyindividualsinapopulation,andbehaviouris

anemergentpropertyoftheseconditions.Forexample,the

intensityofcompetitionformateswillbelargelydeterminedby

populationstructureanddensity,andtheabilityoftheherdto

exploitenvironmentalheterogeneitythroughforagingbehaviours

willbelargelybedeterminedbypopulationdensityandhabitat

characteristics.Vertebratesexhibitaremarkableabilityto

modifybehaviour,includingterritorialdefence,matingsystem,

orseasonalmovementpattern,inresponsetoenvironmental

factors(Lott1984).Here,wedescribedesirablebehaviours

relatedtosocialstructure,mating,foraging,andmovements.

Unlikepopulationorgeneticcomposition,behaviourscanonly

rarelybemanipulateddirectly,andbehavioural“adjustments”

mustbeaccomplishedbymodifyingotherfactors.


9.3.1 Social structure and spacing

Bisonareinherentlygregariousandtherearemanyhistorical

observationsofhugebisonherdsroamingacrossNorthAmerica.

Despitetheenormoussizeofsomebisonaggregations,astute

observersconsistentlyreportedadefinablepopulationstructure

wherecows,calves,andimmaturemalesformedmixed-sex

groups,andwherelargebullstendedtoformseparate,much

smallergroupsthroughoutmuchoftheyear.Groupsofbullsare

typicallysmallerthancow-dominatedormixedgroups,andbison

bullshavefrequentlybeenobservedalone(Allen1876;Berger

andCunningham1994;Meagher1973;Meltonet al.1989).In

winter,thegeneralpatternisoneofsmallermixedgroups,with

groupsizeincreasingtolargeaggregationsthatpeakinsize

duringthesummerbreedingseasonandthenrapidlydiminishing

(BergerandCunningham1994;Hornaday1889).

Thefundamentalsocialgroupinbisonisthoughttoconsistof

matrilinealgroups(Greenet al. 1989),althoughthepersistence

ofthesegroupsinpopulationsthatdifferinsizeandecological

circumstancesispoorlydocumented(e.g.,McHugh1958).These

generalpatternsprovideabasisforsocialbehaviouralguidelines:

1) Bisonherdsshouldhavethecapacitytoexhibitseasonal

changesingroupsize;

2) Averageherdsizeswillusuallybesmallerinmountainsor

mixedterrainthaninopenprairie;

3) Oldbullswillbeobservedaloneorinsmallgroupsduring

muchoftheyear;

4) Persistenceofmatrilinealgroupsshouldbefacilitatedand

activitiesthatdividematrilinealgroupsshouldbeavoided;

5) Activities(roundups,harvest,visitordisruptions,andsoon)

thatdisruptsocialgroupingsshouldbeavoided.Where

unavoidable,implementcarefullytominimizedisruptions.

9.3.2 Foraging and movements

Hornaday(1889)describedahighlynomadicforagingstrategy,

whereplainsbisonseemedtowandersomewhataimlessly

untiltheylocatedfavourablegrazingconditions.Bisonthen

grazeduntilaneedforwatermotivatedfurthermovement.More

recentstudiesofbisonforaginghaveshownthattheyactively

selectmorenutritiousforages,andforageinahighlyefficient

mannerthatsatisfiestheirnutritionalneedsandthatfrequently

complimentsdietselectionbysympatricherbivores(Coppock

et al.1983;HudsonandFrank1987;LarterandGates1991;

SingerandNorland1994;Wallaceet al.1995).Spatialvariation

inforageisproducedbynaturalgradientsinsoilmoisture,soil

nutrients,fire,otherdisturbances,includingforagingbybison.

AftermassivewildfiressweptalongtheAlaskaHighwayinNE

BritishColumbiaandtheSWYukonTerritoryduringtheearly

1980s,bisoncontinuedextensiveuseofrecoveringareas15

yearslater(Larteret al.2007).Bisonserveasanecosystem

Geneticdiversity Whenremovingalargeproportionofaherd,theprimarythreattolong-termpreservationoftheherdisalossofgeneticdiversitythatcanbeverydifficult,ifnotimpossible,torestore.Therefore,thoroughgeneticevaluation(e.g.,section9.2.3),isnecessarybefore,during,andafterplannedlarge-scaleherdreductions.Theprimarygeneticconsiderationsshouldbetheoverallmaintenanceofmitochondrialandnucleardiversity,suchthatthegeneticarchitectureoftheherdismaintainedduringandafterthereductionperiod.Routineexaminationofculledanimalsduringthereductionperiodwillallowfordetection—andhopefullycorrection—of“biased”removals,suchasremovalofasibshipormultigenerationalfamilygroups.Preferentialremovalofrelatedindividualscanleadtolossesingeneticdiversityandeffectivepopulationsizeandshouldbeavoided(Frankham1995).

Herdcomposition If,priortoremovals,theherdhasthedesiredcomposition,bisonshouldberemovedproportionallyfromallageandsexclassestoavoiddisruptionofsocialbehavioursanddemographicstructure.Ifthecurrentherdstructureissubstantiallydifferentfromthatdesired(e.g.section9.2),animalsmaybepreferentiallyremovedfromcertainclasses.Inthecaseofdisproportionalremovals,particularlycareshouldbetakentoassessandmitigatethepotentialeffectsofremovalsonsocialstructureandgeneticdiversity.

Populationsubstructure

Populationsubstructureislikelyimportantinmanybisonpopulations(seesection9.2).Thepresenceofdistinctsubpopulationsshouldbecarefullyevaluatedpriortolarge-scaleherdreductionsandaccommodatedinplannedreductions.

Timescale Bisonshouldberemovedatregularintervals(ratherthanlarge,occasionalevents)tominimisepotentiallyirreversibleimpactsonsocialstructureandgeneticdiversityTheexacttimeperiodforremovalswilllikelybedifferentforeachsituationandwilldependonsuchfactorsastotalherdsize,thetotalnumberofanimalstoberemoved,andtheresourcesavailable(e.g.,facilities,manpower).

Assesseffectsofmanagementactions

Beforeandaftermanagementactionsareimplemented,thoroughgenetic,health,anddemographicmonitoringisnecessarytoevaluaterecoveryefforts,andtodetecttheneedforalternativemanagementstrategies.Smallpopulationsareespeciallysensitivetomanagementchanges,andcomprehensivemonitoringmaybenecessaryforsometimetoensuretherecoveryofsuchherds.Sections9.2,1,9.2.3,and9.5.2summariseinformationthatshouldbemonitoredtodetectchangesinatimelymanner.Especiallyforsmallherds,theoverallhealthoftheherdshouldbecontinuouslymonitoredtodetectandtreatanyheritableortransmissiblediseasesthatmayimpederecoveryefforts.

table 9.4 Important considerations for culling bison herds. See section 9.2.8 for explanation.


engineer,bothrespondingto,andcreating,heterogeneity.Bison

traditionallyexploitedbroad-andfine-scalevariationinforages,

forexample,sometimesmigratinglongdistancesinresponseto

snowfallordrought.

Guidelinestohelppreservedesirablebehaviouralpatternsareas

follows:

1) Allowbisontorespondtodifferencesandchangesinthe

distribution,quality,andquantityofforagesbymoving

within,andbetween,ecosystems;

2) Provideherdrangesthatincludeabroadvarietyofhabitats

sothatbisoncanexploitshort-term(seasonal)andlong-

term(annual,multi-year)heterogeneityinforagesfrom

patchtolandscapescales;

3) Bisonherdsshouldhavetheabilitytocreateand

respondtospatialvariationinforagequality,quantity,

anddistributionthatistheresultofunderlyingvariationin

resourcesnecessaryforplantgrowth,tovariationresulting

fromherbivoreforaging(bybison,prairiedogs,andother

species),andtovariationresultingfromenvironmental

disturbancessuchasfireandflood;

4) Balancetheadvantagesoflargerpopulationsizeagainsta

needtoavoidpermanenthabitatdamage.

Theseguidelinessuggestthatbisonshouldhaveaccesstovery

largeareasinwhichtheycanexploitnaturalheterogeneityin

forageabundanceandquality.Fencesandotherimpedimentsto

movementshouldbeminimised.

9.3.3 Mating behaviour

Differentialreproductionresultingfrommatecompetitionisan

importantevolutionaryprocessand,assuch,itiscrucialto

allowbisontoexpressnaturalmatingbehaviours.Thefollowing

guidelinesforpopulationmanagementsupportthisgoal:

1) Thesexratioofapopulationshouldbenearlyequal,andin

nocaseshouldeithersexconstitutemorethan60%ofthe

population;

2) Apopulationshouldincludeabout50matureand

reproductivelyactivemalesforevery100cows(Gates1996,

unpublisheddata;Gateset al.2005;Komerset al.1992);

3) Allowinteractionandfightingbetweenbulls.

Theratioofmaturemalestocowswillgenerallybelowerthan

theoverallsexratiobecausemales(bulls)achievesexual

maturityatagreateragethanfemales(cows)andthemortality

rateofmalesishigherthanforfemales.

9.3.4 Limiting factors and natural selection

Chapter6describedfactorsthatwerehistoricallyresponsible

forseasonalandperiodicfluctuationsinthesizeanddistribution

ofbisonpopulations.Thesefactors,andthepopulation

segmentstheytendtoaffect,areconsistentwithcontemporary

observations(Chapter6;Gaillardet al.1998).

Generalguidelinesconsistentwithourunderstandingof

“normal”demographicprocessesare:

1) Naturalmortalityratesshouldbehighestforcalvesandthe

oldestageclasses;

2) A“normal”rangeforcalfsurvivalis40-90%,andcalf

survivalshouldvarywithwinterseverity,predation

pressures,andforageavailability;

3) Naturalsurvivalratesforprime-ageadultswillnormallybe

about95%;

4) Undergoodconditions(e.g.,lowdensity,mildwinter,good

forageproduction),pregnancyratesforthree-year-old

cowswillbe70%orgreater;

5) Undergoodconditions,pregnancyratesforprime-age

cows(generallyabout4-15yearsold)willnormallybe70-

90%andsometwo-year-oldcows(probablylessthan5%)

willproducecalves;

6) Diseasewillgenerallylowerreproductiveperformance.

9.4 habitat and Biodiversity Management

Bisoncan,andusuallywill,significantlyinfluencehabitatand

biologicaldiversity,andbisonaregenerallyregardedasa

foundationspeciesandecosystemengineers.Thisisespecially

trueforecosystemswherebisonarerelativelyabundantand

rangeoverlargeareas.Modern,small-hornedbisonhavealong

historyasanintegralpartoftwomajorecosystems:theNorth

AmericanGreatPlains(plainsbison)andthesedge-meadow

ecosystemsofnorthernCanadaandAlaska(woodbison).

Bisoncanprofoundlyaffectecosystemtrophicstructures,

bio-geochemicalcycling,speciescomposition,andpatterns

ofspeciesdiversity.Somemajortypesofecologicalprocesses

thatbisoninfluencearesummarisedinTable9.1,whileamore

detailedreviewisprovidedinChapter6.

Belowwelistguidelinesforbisonmanagementthatwill

helpconservebiologicaldiversity.Decisionsonactivebison

managementrequireknowledgeofproductivity,stocking

rates,andmovementpatterns.Goodsourcesofinformation

formanagementofconfinedorsemi-confinedbisonherdsin

westernhabitatsaretheUSDA’sNaturalResourceConservation

Service(NRCS)anditsFieldOfficeTechnicalGuides(http://

www.nrcs.usda.gov/technical/efotg/).Thesedocumentsprovide

informationonprimaryproductivity,recommendedstocking

rates,animalconversionunits,andotherinformationrelevant

torangemanagement.TheNRCSguides,however,focuson

obtainingthemaximumsustainedyieldoflivestock.Thereisno

comparableresourceforbiologistsmanagingnorthernbison.For


northernbisonherds,managersshouldreviewrelevantliterature

andconsultwithbiologistsinborealregionsthatsupportwood

orplainsbisonpopulations.Toenhanceandconserveregional

biologicaldiversity,bisonmanagerswillneedtoconsiderlocal

andregionalissues,culturalandeconomicissues,andland

usepatterns.Forexample,iftheconservationofprairiedogs

andotherspeciesassociatedwithshortvegetationstructureis

desired,plainsbisonstockingratesshouldbehigherthanthose

recommendedbytheNRCSfieldguides.

Thefollowingguidelinescanhelppromoteconservationof

biodiversitytoahigherdegreethanisachievedinmostlivestock

productionsystems.

1) Promotethemovementanddistributionofbisonacross

thelandscapeinas-natural-a-fashionaspossible,

includingtheexistenceofsub-herds;

2) Manageforamosaicofseralconditionsandgrazing

intensitiesacrossalandscape.Ifparticularconditionsor

seralstagesareregionallyrare,theyshouldbefavoured

throughmanagement.Thismaycontrastwithtraditional

livestockgrazingmanagementthatattemptstoimpose

relativelyuniformgrazingpressureacrossanentire

managementunitandavoidareasof“overgrazing”;

3) Managefireusingthebestavailableinformationonnatural

firepatternsfortheregion.Leaveunburnedareasas

refugiaforinvertebratesandsmallmammals;

4) Restoreand/orconserveprairiedogsandothergrazers

thatinteractwithbison;

5) Wherepossible,restoreormaintainnativepredatorsof

bison,i.e.,wolvesandbears;

6) Ifmineral,food,orwatersupplementsarenecessary

theyshouldbeprovidedinawaythatcreateshabitat

heterogeneity(asapointattractantratherthanbeing

distributeduniformlyacrossthelandscape);

7) Managesothatbisondonotgrazenaturallyinaccessible

areas,forexampleisolatedbuttesandsteepslopes,which

increaseslandscapeheterogeneity;

8) Leavecarrionin situ.

9.5 Disease Guidelines: Considerations for Infected and Uninfected herds

Asallwildlifepopulationsarehoststoawidevarietyofnatural

pathogens,andthesepathogensformanintegralcomponentof

ecosystemhealth,welimitthefocusofthissectionto:

• Pathogensthatlimitbisonpopulationrecoverydirectlybyreducingsurvivaland/orreproduction,(demonstratingabisonpopulationimpact),and/or

• Pathogensthatindirectlypreventbisonrecoveryastheyformthreatstoexistinglivestockandwildlifepopulations(e.g.,so-calledeconomicdiseases).

Ingeneral,pathogensthatfittheabovecategoriesareexotic

(i.e.,havespilledoverfromdomesticlivestockpopulations),

suchasbovinetuberculosis,brucellosis,bovineviraldiarrhoea

(BVD),andmalignantcatarrhalfever(MCF).

Wobeser(2002)outlinedfourgeneraldiseasemanagement

philosophies:(1)prevention,(2)control,(3)eradication,and(4)

thelaissez-faireapproach(donothing).Preventativemeasures

arethosedesignedtoinhibitthespreadofdiseasetouninfected

individualsorpopulations.Forexample,theBisonControlAreain

theNorthwestTerritoriesismanagedtopreventthemovementof

diseasedbisonfromWoodBuffaloNationalPark(WBNP)tothe

MackenzieBisonSanctuary(Nishiet al.2002c).Controlmeasures

reducethefrequencyofoccurrenceortheeffectsofadisease

withinapopulationorcontainthespreadofthedisease.Under

thisregime,adiseasewillnormallypersistindefinitely,requiring

continuedmanagement.TheYellowstoneNationalPark(YNP)

cooperativebisonmanagementplanincorporatesnumerous

controlmeasuresincludingtest-and-slaughterofdiseased

bison,hazingofbisonbackintothepark,vaccination,andradio

telemetryofpregnantbison(NPS-USDOI2000).Totaleradication

ofadiseaseisdifficultand,insomecases,maynotbepossible

givencurrenttechnologyandresources.Test-and-slaughter

programmes,inconcertwithvaccination,mayeradicatea

diseasefromacaptivepopulation(Nishiet al.2002c);however,

thesetechniquesaredifficulttoapplytofree-rangingwildlife

(Wobeser2002).Inlargerpopulations,oroverlargerareas,

intensivemanagement,emphasisingtreatmentandvaccination,

maybeinappropriate,unsustainable,orsimplyimpractical

(Woodruff1999).Inthesecircumstances,managingpopulation

size,structure,areaofoccupancy,ortheriskofcontactbetween

hostspeciesoradjacentpopulations,couldofferalternatives

tomoreintensiveinterventions.Depopulation(=eradication)of

aninfectedherdisapotentialoption;however,theremaybe

considerablelogisticalchallengesandconservationandpolicy

issuesincludinggeneticconservationorsalvage,cascading

ecologicaleffects,andpublicopposition(Nishiet al.2002c;

Wobeser2002).Selectionofadiseasemanagementapproach

dependsontherationaleformanagement,whetherthedisease

isalreadypresentinapopulation,theavailabilityoffunding,

andthelikelihoodofsuccess(Wobeser2002).Managersshould

alsounderstandtheecologyandpathologyofthedisease,

thedynamicsofthepathogen-hostrelationship(Bengiset al.

2002;Wobeser2002)andtherisktoadjacentuninfectedhost

populations,includingbison.

Ourdiseaserecommendationsfocusonfourdiseasecontrol

strategies:prevention,surveillance,management,andresearch.

Werecommenddevelopmentofadiseasemanagementplan

undertheumbrellaofarestorationprogrammeplanthatis


consistentwithconservationprogrammegoalsandincorporates

theexpertcounselofwildlifeveterinarians,epidemiologists,

andotherdiseasespecialists.Diseasemanagementplans

shouldbedevelopedinalocalcontextandinvolveconsiderable

stakeholderparticipation.

9.5.1 prevention

Thorougheffortsshouldbemadetopreventtheintroduction

ofexoticdiseasesintoexistingandfuturefree-rangingbison

populations.Introductionofnovelpathogensintobison

populationscouldoccurbycontactwithfree-rangingwildlife

orthroughcontactwithcaptivewildlifeorlivestock(herein

referredtoas“potentialdiseasesources”).Asageneralstrategy,

managersshouldstrivetomaintainpopulationattributesthat

reducethelikelihoodofdiseaseestablishment,oranincreasein

diseaseprevalenceshouldapathogenbeintroduced(Table9.5).

Forexample,animaldensitymayinfluencediseasetransmission

andnutritionalstatusofanimals.Habitatconditions(e.g.,

marshyareasforbluetongueordryconditionsforanthrax)and

thepresenceorabsenceofpredatorscaninfluencedisease

establishmentorprevalence.

Adiseaseriskassessmentshouldbeconductedforexisting

andfuturefree-rangingbisonpopulations.Thisriskassessment

shouldincludecomponentsofdiseasesurveillance(inboth

thepotentialdiseasesourceandthepopulationatrisk)to

determinewhatpotentialpathogensareinvolved,contact

potential(todetermineriskofdiseasetransmission),potential

consequencesofdiseasetransmission,recommendedstrategies

tomitigatediseaserisk,andcollateralimpacts

oftheseactions.Preventiveactionsmayinclude

preventionofdispersalbetweeninfectedand

atriskpopulations,habitatmodification,and

maintainingoptimalpopulationdensity,aswellas

understandingthehistoryofpertinentdiseases

withintheregion.

Thedevelopmentofaclinicalinfectiousdisease

involvesacomplexinteractionbetweenthe

host(bison),theagent(pathogen),andthe

environment(habitat).Alterationstoanyoneof

thesefactorsmayinfluencetheabilityofadisease

tobeintroducedorestablishedwithinagiven

population.Therefore,athoroughunderstanding

ofthebiologyofthehost,agent,andenvironment

isnecessarytominimisetheriskofintroducingor

amplifyingnon-nativediseases.

9.5.2 Surveillance

Thefirststepinmanagingdiseasesinapopulation

istodetermineifapathogenispresent,andifno

infectedanimalsaredetected,theprobabilitythat

thediseaseispresent,butatanundetectablelevel.Surveillance

canalsobeusedtodeterminetheprevalenceofadiseaseknown

tooccur,andtomonitorchangesinitsprevalenceovertime.

Diseasesurveillancecanbepassiveoractive.

Passive,oropportunistic,surveillancewouldincludedisease

testingofanimalswithclinicalsignsand/orthosethatarefound

deadormoribund.Ifacauseofdeathisnotapparent,itmay

beprudentandinformativetosubmittheentirecarcass,where

possible,forafulldiagnosticnecropsytodeterminecause

ofdeath.Localmanagementstaffshouldbetrainedinbasic

necropsytechniques,andtocorrectlycollectcriticalsamples

whenitisnotfeasibletosubmitentirecarcasses.Diagnostic

evaluationisparticularlyimportantifhumancontactmayhave

ledtotransmissionofazoonoticdiseasetoanemployeeora

memberofthepublic.Ifpredatorsarepresentintheecosystem,

theymayremoveorcompromisecarcassesbeforetheycanbe

collectedforinvestigation.

Activesurveillancewouldincludecapturinganimalsand

testingfordiseases,orsolicitingsamplesfromhuntersof

huntedpopulations.Often,diseasesurveillanceisperformed

bycollectingserumfrombloodsamplesandtestingthesefor

antibodiestodiseasesofinterest.Itisimportanttoremember

thatthepresenceofantibodiesdoesnotconfirmdiseaseinan

animal,onlyexposuretothepathogenatsomepointinthepast.

However,onemightinferthatthepathogenofinterestispresent

inapopulationbasedonpositiveserologicalresultsfrom

individualanimals.Additionally,mostdiagnostictestshavebeen

developedfordomesticlivestockandtheirapplicabilityinbison

passive Control Eradication

Monitorherdforclinicalsignsofdisease

Alltechniquesunderpassivecategory

Alltechniquesinpassiveandcontrolcategories

Implementmovementrestrictionsfrompopulationsthatarediseasedorofunknowndiseasestatus

Manipulatepopulationdensitytominimisespreadofdensity-dependentdiseases

Testandcullinfectedmembersofthepopulationwherescientificallyfoundedandlogisticallyfeasible

Modifyhabitattominimisecongestion

Herdleveltreatmentiffeasible(rarelyappropriateinfree-rangingpopulations)

Combinationsofvaccination,treatmentortestandculldevelopedtorapidlyeliminatedisease

Vaccinationifavailable Depopulationofhostspeciesfollowedbyre-populationwithdisease-freeanimals

Implementtemporal/spatialseparationbetweeninfectedandsusceptiblepopulations(wildlifeorlivestock).

Eliminationofbisonfromaffectedareas

table 9.5 Potential management techniques appropriate for management objectives to

passively manage, control, or eradicate disease.


maynothavebeenvalidated.Testingfaecesforparasitesor

pathogens,suchasMycobacterium aviumpseudotuberculosis

(Mptb),mayalsobebeneficial.Activesamplingallows

estimationofthepopulation-levelprevalenceofthedisease(as

itcanhavegreaterstatisticalvaluebecauseitislikelytobemore

randomthanpassivesampling),althoughpassivesurveillance

asadiseasedetectionstrategymaybemoresuitablefor

protectedpopulations.Highprioritiesfordiseasesurveillance,

basedonhuman,wildlife,andlivestockhealthconsiderations

couldincludeanthrax,bovinetuberculosis,brucellosis,BVD,JD,

andMCF,amongothers.Finally,whileforeignanimaldiseases,

suchasfoot-and-mouth(FMD)diseaseorheartwater,arenot

highlylikelytoaffectAmericanbisonpopulations,theyshould

beonthe“watch”listofpotentialdiseases,sinceintroductionof

diseasessuchasFMDtoNorthAmericawouldhavesignificant

economicimpacts.

Non-specificsignsofdiseaseshouldbemonitoredand

investigated,eventhoughdiagnosticsarerequiredtodetermine

cause(e.g.,poorconditioncouldbeduetoageorhabitat

condition,parasitism,orJD,amongothercauses;Table9.6).

9.5.3 Management

Whenapathogenhasbeendetectedinabisonpopulation,

anevaluationshouldbemadetodetermineifadisease

managementplanshouldbedevelopedthatisconsistent

withthegoalsforthebisonpopulation.Potentialdisease

managementobjectivesare:a)apassiveapproachwhereno

actions,oratleastnoactionsthatmanipulateanimals,are

takentocontrolthedisease,b)acontrolstrategywhereactions

aretakentolimitdiseaseprevalence,spread,orrisk,orc)an

eradicationstrategywhereactionsaretakentoremovethe

diseasefromthepopulation.Allthreestrategies(Table9.5)will

likelyinvolvemonitoringdiseaseprevalence(eitheractively

orpassivelyasdefinedabove).Strategiesusedwillalsobe

influencedbytheintensityofmanagementwithintheherd.For

example,managementoptions,suchasvaccination,wouldbe

moreeasilyappliedtoaherdthatisintensivelymanagedwith

round-ups.

9.5.4 research

Furtherresearchwillbenecessarytodevelopandimplement

toolsforsuccessfuldiseaseprevention,surveillance,and

management.Forexample,manyofthediagnostictests

commonlyusedinbisondiseaseprogrammesweredeveloped

foruseinthelivestockindustryandhavenotyetbeenvalidated

inbisonpopulations.Furthermore,keyquestionsremain

aboutthepresence/absenceanddistributionofdiseasesin

populations,andtheirpotentialeffectsonbisondemography

andgenetics.

Researchshouldbedesignedtomeettheneedsoflocal

managers,sothatresultscanbeappliedinmoregeneral

contexts.Alimitedlistofsomeofthekeydiseaseresearch

themesinclude:

• Diagnostics(specifictobison,withhighsensitivityandspecificitytodetectadisease);

• Vaccination/immunology;

• Roleofgeneticsindiseaseresistance;

• Diseaseepidemiology(e.g.,transmission,demography)andriskanalysis(spreadofdiseaseamongandbetweenwildanddomestichosts);

• IdentificationofemergingdiseasethreatstobisoninNorthAmerica;

• Pathology;

• Effectofdiseaseonpopulationgrowthandviability(bothindirectanddirecteffects).

Whereresearchisneededforaparticulardiseasesurveillanceor

managementquestion,bisonmanagersareencouragedtowork

withfederal,state,university,andprivateresearcherstomeet

thisneed.Anadaptivemanagementapproachwillbenecessary,

especiallywheninformationaboutaspecificdiseaseisscarce.

9.5.5 Stakeholder involvement

Insummary,bisonpopulationsshouldbemanagedtoprevent

theintroductionandspreadofdiseasesthatdirectly,or

indirectly,impactbisonrecovery.However,bisondisease

managementstrategieshavebeen,andcontinuetobe,

controversialbecausetheapparentsolutiontothedisease

problems(or“cure”)isoftenperceivedtobeworsethanthe

diseaseitself.Extensivestakeholderinvolvementindisease

managementplansisabsolutelycriticaltosuccessfulbison

diseasemanagement;suchmanagementstrategieshave

oftenfailedwithoutit.Typicalstakeholdersinbisondisease

Loss of body condition Abnormal behaviour

Abnormalexudatesfrombodyorifices

Isolationfromtheherd

Cloudyeyes Abnormallossofhaircoat

Diarrhoea Abortion

Abnormallypoorhaircoat Lameness(multiplelimb)

Somnolence Abnormalinteractionwithhumans

Unexpected/abnormalmortalityevents

table 9.6 Non-specific clinical signs of disease.


managementincludestateandfederalagencies(animalhealth

regulators,landmanagementagencies,andwildlifeagencies),

landowners,livestockproducers,conservationorganisations,

sportsmen’sorganisations,andnativepeoplegroupsand

organisations.

9.6 Active Management: handling, herding, Infrastructure

Bisondiffersubstantiallyfromcattleandtheyoftenrespond

poorlytohandlingthatwouldberoutineforcattle.Bisonshould

betreatedaswildlifeandhandledinfrequentlyorpreferably,

notatall.Whenhandlingisabsolutelynecessary,suitable

precautionsmustbeobserved,forexample,oldbulls(andcows)

canbeverydangerousanddifficulttohandle.Handlingfacilities

designedespeciallyforbisonareneededtoensurethesafetyof

boththeanimalsandpeoplethatworkwiththem.

Theoverarchingprincipleisthattopreservethetrue,wild

natureofbison,activemanagement,throughherdingorother

interventions,shouldbeminimised.Handlingbisoncanresult

inchangestobisonbehaviourandleadtomanagement-based

selectionthat,overtime,altersgeneticcompositionoftheherd

(Lott1998).Thesechangescanbeirreversibleanddetrimental

toconservingorrestoringa“wild”stock.Thegeneral

guidelinesonpreservingnormalbisonbehaviourbelowareonly

anintroduction.Anunderstandingoftheconceptsofbison

behaviour,practicalexperience,andperhaps,specialtrainingis

requiredtohandlebisonwell.Werecommendconsultingknown

expertsforadvice.Bisonhandlingpresentsagreaterchallenge

thanhandlingdomesticstockandmanagingfor“wild”behaviour

isarelativelynewconcept.

9.6.1 handling

“Sure,youcanherdbison…anywhere

theywanttogo.”

Whenactivemanagementofbison

isnecessary,use“calmanimal”

techniquesbasedonanapproach

thatadjustshumanbehaviourtofitthe

naturalresponseoftheanimal,rather

thantheotherwayaround(Grandin

andJohnson2004;Roberts1996).This

approachsimplifieshandling“wild”

animals,anditreducesthetendency

formanagerstoinadvertentlyremove

ecologicallydesirabletraitsovertimeby

selectiveculling.

Guidelinesforhandlingbisonare

predicatedonexploitingtheirnatural

instincts(Lott1991).Bisonarestrongly

motivatedbyfood,bythreatof

predation,andbytheneedtomaintain

socialcohesion.Managerscanexploitthesetendencies:bison

canbeledwithfood,andlighterfencingisadequateifbetter

foodsarenotdetectedacrossafence.Byappearingasa

predator,managerscanprecipitateuncontrollableflightoreven

attack.Lessaggressivetechniquescanbeusedtocontrolbison

movementswhileminimisingriskandeffort.Bison’sherding

“instincts”prevailandgroupsofbisoncanbemotivatedto

movesimplybymotivatingtheleadcow.Bythesametoken,

disruptingtheestablished“peckingorder”orcow-calfbondsin

aherdstressesbisonandmakesthemhardertohandle.

Socialcohesioninbisonhasimportantimplicationsforhandling.

Inthewild,herdsofbisonfoundfoodandfendedoffpredators

betterthanloneanimals,andsocialcommunicationprovides

importantclueswhenhandlingbison.Potentialdangersignals

includeposturessuchastailsup,intensestaring,snortingand

pawing,and“growling”(bybulls)(Lee1990a).Moresubtle

signalscanadvertiseanxiety,intenttomoveaway,orwillingness

tofollow.

Itiseasiertoleadthantodrivebison(Lee1990b).Oncetrained

tocometovehiclesforfood,bisonwillreadilyfollowavehicleto

differentpartsoftheirhomerange,ortheycanbegatheredfor

processing.Fooddispensedatcorralsduringannualprocessing

canmotivatebisontomoveontheirowntowardcorralsatthe

appropriatetimethenextyear.

Predator-relatedbehavioursofbisonthathandlerscanuseto

theirbenefitinclude:

1) Atendencytointerpretadirectapproachorstaringasa

threat;

plate 9.2 Meeting of stakeholders at Vermejo Park Ranch, IUCN Bison Specialist Group. Photo:

John Gross.


2) Atendencytofleeifapproachedtooclosely,tooswiftly,or

toodirectly;

3) Atendencytodriftawayifapproachedslowlyand

tangentially;

4) Reducedintensityofresponsewithrepeatedharmless

encounters.

Implicationsofbisonbeingattractedbyfoodinclude:

1) Thedifficultyoffencingthemawayfromgood-qualityfood;

2) Atendencyforbisontoseekoutthehighest-quality

foragesintheirhomeranges;

3) Thepoweroffood,whenproperlymanaged,toamplify

desiredbehaviourandreduceundesiredbehaviour.

Waysinwhichsocialcohesioncanaffecthandlinginclude:

1) Thestrongtendencyforsocialgroupstofollowthelead

animal’sresponse;

2) Thedifficultyofseparatingcowsfromtheiryoungcalves

duringprocessing;

3) Thestressanddisorientationthataccompanydisruptionof

socialgroups;

4) Theeaseoftranslocatingandmovinganimalsifsocial

groupsremainintact.

9.6.2 Fencing

Motivatedbisoncaneasilycrossordestroyfencesgenerally

effectiveatconstrainingcattle.Bison-prooffencescanbe

expensive,andifnotcarefullydesigned,mayhinderpassageby

otherwildlife.Effortstoreduceabison’smotivationtobreach

fencescangreatlyreducethecostsoffencingrequiredto

containanimals,andreduceadverseeffectsonotherspecies.

Appropriatefencedesignsvarywithcircumstance,anda

detaileddiscussionisbeyondthescopeofthischapter.More

detailedrecommendationsandevaluationsshouldbeconsulted

beforeanyconstructionbegins(e.g.,Butterfield1990a;1990b;

Gates2006).Ingeneral,athree-strandbarbed-wirefencecan

holdbisonthathavebeentrainedtoavoidfencesandthatare

notstronglymotivatedtocrossthefence.High-tensilewireis

morecommonlyusedtobuildnewbisonfencesortoreinforce

existingones.Someprefernet-wirefences,butdependingon

design,theycanbeformidablebarrierstootheranimalsthat

needpassage.Electricfences,hightensileorotherwise,greatly

increasethebarriereffecttobison,andalsoconditionthemto

avoidfencesingeneral.

Theneedtoallowpassageforotherwildlifeaffectsfence

designwheredeer,pronghorn,elk,(orotherlargeungulates)are

present.Hightensilefenceswiththebottomwireatleast51cm

(20”)offthegroundandthetopwire107-132cm(42”-52”)off

thegroundwillconstrainbisonundermostcircumstances,while

stillpermittingdeerandpronghorntopassunderthefenceand

mostelktojumpoverthefence(KarhuandAnderson2003).A

three-wireelectricfencewiththebottomandtopwires56cm

(22”)and107cm(42”)offtheground,respectively,offeredbetter

passagefordeer,pronghorn,andelkthandidtwo-orfour-wire

designs(KarhuandAnderson2003).Gates(2006)provides

additionaldetailsandrecommendationsthatvaryfromthose

above(e.g.,topwire152cm(60”)aboveground).Additional

guidanceshouldbeobtainedtoensurefencingmeetstheneeds

ofanyspecificapplication.

Factorsthatcanmodifytheeffectivenessoffencinginclude:

1) Bisondensity;asdensityincreases,moresecurefencing

mayberequired;

2) Deepsnow-packmayrequirespecialdesign

considerations;

3) Damageduetofallingtrees,biggame,vandals,orbison;

4) Attractivefood,orotherobjects,ontheothersideofa

fenceincreasesbisonmotivationtobreachfences.

Factorsthatinfluencetheeffectoffencesondeer,pronghorn,or

elkinclude(Gates2006):

1) Nutritionalstress;adverseimpactsincreaseduringperiods

ofnutritionalstress;

2) Somefencedesigns(e.g.,wovenwire)havegreaterbarrier

effectsthanothers;

3) Barriereffectsthatareonlyseasonalmaynotbeevident

whenfencesarebuilt;

4) Poordesignsmayinjureorkillanimalsorseparatemothers

fromyoung;

5) Predatorsmaykillbiggamemoreeasilybychasingthem

againstfences.

9.6.3 Corrals, pens, and chutes

Corralsandassociatedfacilitiesforwildbisonneedtobemore

carefullydesignedandconstructedthansimilarfacilitiesfor

domesticlivestock.Bisonmaynotrecognisestandardfencing

asabarrier.Youngcalvesrequirespecialattentionbecause

theymayrunintosolidgatesorfences,althoughfencesthat

areabout80%solidappeartopreventthis(Lammers,personal

communication).Fencesandgates,with30-40cm(12”-16”)

planksspaced10cm(4”)apart,effectivelystopbisonand

canbeeasilyclimbedbywranglers.Openfencesnearthe

workingchutes,eventhosethatareverystrong,oftenleadto

injuryandmortality.Totallysolidfencingcanbedangerousfor

peopleworkinganimalsfromthegroundiftheyneedtoescape

crowdedorcharginganimals.

Bisonhandlingfacilitiesmustaccommodatethestrongsocial

hierarchyandaggressivebehavioursthatbisonexhibit.


Appropriatefacilitiesusuallyincludecustomsizedand

constructedchutesandalleyways,crashgates,andchute

crowdingtubes.Itisexpensivetoconstructfacilitiessafefor

bison(andthepeopleworkingwiththem),andwestrongly

recommendvisitingfacilitiesthathaveproventobesafe

andeffective.HighlycrediblefacilitiesincludethoseatYNP

(Gardiner,Montana),theBacaRanch(Colorado),Badlands

NationalPark(SouthDakota),andEINP(Alberta).

9.7 Modelling to Assess Bison populations and habitat

Computermodelsareroutinelyusedtoimproveour

understandingofbisonpopulationanddiseasedynamics,

andtoforecastprobablegeneticconsequencesresultingfrom

particularmanagementactions.Inthefuture,weshouldexpect

evenmorewidespreaduseofquantitativemodels,whichcan,

andlikelywill,beusedforabroadrangeofpurposes.Adetailed

treatiseonmodellingiswellbeyondthescopeofthisplan.The

maingoalsofthissectionare,therefore,toprovidereaders

withtheminimalbackgroundnecessarytoseriouslyconsider

theutilityofusinganexistingmodel,orofconstructinganew

management-orientedmodel,andtoprovidesufficientinsight

tothemodellingprocess,thattheycanreasonablyevaluatethe

validityandusefulnessofmodelresults,oratleastaskquestions

thatwillhelpresolvetheseissues.

Forconservationpurposes,populationviabilityanalysis(PVA)

andpopulationhabitatviabilityassessment(PHVA)have

becomecommon,andimportant,approachesforassessing

existingpopulationsandforevaluatingpotentialrestoration

orreintroductionprojects.WerestrictPVAandPHVAto

analysesthatemployquantitativemodellingtoassessthe

riskofextinction,orwhichattainaquantitativepopulation

thresholdgreaterthanextinction(“quasi-extinction”,from

Ginzburget al.1982;Burgmanet al.1992;Rallset al.2002).

Otherthresholdsforevaluationcouldincludeattaininga

specifiedlevelofinbreedingdepressionorallelicdiversity,or

estimatingthelikelihoodthataproposedintroductionplanwill

resultinestablishment.Conclusionsdrawnfromexpertpanels,

committees,andothersourceofopinions,intheabsenceofa

quantitativemodel,donotconstituteaPVA(Reedet al.2002).

PHVAisamuchbroaderprocessthanPVA,andincludes

evaluationofgeographical,social,regulatory,andecological

considerationsthatmaysignificantlyaffectaspecies.ThePHVA

processincludesabroadrangeofstakeholdersandleadsto

specificrecommendationsforconservingaspeciesinthearea

considered(http://www.cbsg.org/cbsg/phva/index.asp).Viability

analysisisimportanttobisonconservationbecausesomany

bisonpopulationsaresmallandclearlyatrisk,andbecausewe

havearichknowledgeoffactorsnecessarytoconductcredible

andinsightfulevaluations.

Thesmallsizeofmanybisonherdshasraisedconcernsabout

retentionofgeneticdiversity,andtheseconcernsmotivated

detailedsimulationstoevaluateeffectsofmanagementactions

onretentionofgeneticvariationinbisonherds(Grosset al.

2006;Halbertet al.2005;WilsonandZittlau2004).Other

modellingstudieshavefocusedonbrucellosisdynamicsandits

controlinbison(DobsonandMeagher1996;Grosset al.1998;

2002;Petersonet al.1991;Treanoret al.2007)andonillustrating

populationdynamicsofbison(Brodie2008).Allwildlifemodels

areultimatelylimitedbydataavailability,andmodelresults

canbemisleadingwhenforecastsarepresentedwithan

apparentprecisionthatisnotjustifiedbytheunderlyingmodel

assumptions,structure,ortheaccuracyofmodelparameters

(Rallset al.2002;Reedet al.2002).Ingeneral,themost

appropriateuseofsimulationmodelresultsistoevaluatethe

meritsofalternativemanagementactions,ratherthantodefine

anabsolutethresholdpopulationsize.Inparticular,minimum

criticalpopulationsizesmaybesensitivetosmallerrorsin

parameterestimates,ortothefunctionalstructureofstrong

environmentalperturbations.

9.7.1 Guidelines for using computer simulations

Thefirstcriticalstepistoclearlydefinetheobjectivesofthe

modellingexercise.Iftheintentistoevaluatemanagement

actions,thebestobjectivesarequantitative,specific,time-

bound,andconsistof“treatment”variables(e.g.,numberof

founders,numberorproportionremoved)thatcanreasonablybe

simulatedbyacomputermodel.Agoodobjectivemustinclude

thelikelihoodofachievingthedesiredresults,thequantitative

valueofathreshold,andatimehorizon.Forexample,abison

PVAusedthegeneticobjectivetoachievea90%probabilityof

retaining90%ofcurrentlyobservedselectivelyneutralgenetic

heterozygosityfor200years(Grosset al.2006).

Below,weliststepsthatwillberequiredtoconstructacomputer

modeltosupportbisonconservation.Anumberofrecent

treatisesprovidemoredetailedinformationaboutthisprocess

(weespeciallyrecommendBurgmanet al.1993;Bessinger

andWestphal1998;BessingerandMcCullough2002;Hilborn

andMangel1997).Althoughweliststepssequentially,most

modellingexercisesareiterativeandinvolvesimultaneously

workingthroughanumberofthesetasksandrevisitingthemas

moreinformationorinsightbecomesavailable.

1.Clearlyarticulatetheobjectivesofthemodellingexercise.Itis

essentialtoclearlyidentifyasmall,discretesetof“treatments”

and“responses”.

• Whatmanagementmustbeevaluated?

• Whatistherelevanttimeframe?

• Whatmodeloutputsaretobeevaluated?


2.Determinetherequiredscopeofmodel.

• Singleormultiplespecies?

• Ageorstagestructured?

• Oneormorepopulationunits?

• Spatiallyhomogeneousorwithspatialstructure?

• Whatisthegeographicalextent?

• Areanimal-habitatfeedbacksnecessary?

3.Evaluateexistingsoftwareanddecidewhethertousean

existingprogrammeortoconstructanewmodel.Considerable

timeandmoneycanbesavedbyusing“offtheshelf”software,

suchasRAMAS(http://www.ramas.com/software.htm),

Vortex(Lacy1993),ALEX(Possinghamet al.1992),oranother

modellingenvironment.

4.Collectnecessarydataandestimatemodelparameters.

Thiscanbeahugestep.Datawillbeneededtoestimatemean

vitalratesandrealisticestimatesofvariance.Ecosystemor

habitatmodelswillrequiremuchadditionalinformationto

determinecarryingcapacityandanimal-ecosystemfeedbacks.

Mostpopulation-habitatmodelsusedforPVAwillinclude

catastrophes,estimatesofvarianceinhabitatcarryingcapacity,

andspecificassumptionsontheformandprocessofdensity

dependence.

5.Construct,calibrate,andrunthemodel.Evaluatemodel

results.Considerableeffortmayberequiredtounderstandand

comprehensivelyevaluatemodelinputs,andtounderstand

modelresults.Outputfromasimulationexerciseusually

includeshugequantitiesofdatathatwillneedtobereduced,

summarised,andpresentedinanunderstandableform.

6.Packageresultsinadigestibleandunderstandableformat.

Thisisavastlyunderappreciatedproblem,anditwillbemuch

easierifthemodelobjectiveswereclearandconciselystatedat

theoutset.

7.Rallset al.(Table25.4inRallset al.2002)provideaspecific

checklistforevaluatingthequalityofaPVA,andthischecklist

appliesequallywelltomanyadditionalconservationmodelling

exercises.Theyprovide“yes-no”questionsthatfocusonmodel

objectives,modelstructure,dataandparameterestimation,

analysisofmodeloutcomes,handlingofmodeluncertainty,

interpretation,andpeerreview.Thesecriteriaprovideasound

frameworkforhelpingtoensuremodelsareconstructedand

usedinanappropriatefashion.

9.8 Conclusions

Whilemanytopicsareaddressedinthischapter,effective

managementofbisonultimatelyreliesonthejudicious

applicationofcommonsenseandgoodjudgement.When

bisonhaveaccesstosufficientspaceandforage,andareleft

relativelyundisturbed,theyaremorethanfullycapableoftaking

careofthemselves.Nonetheless,mostbisonwillnotexperience

naturalconditionsthatincludewide-openspacesandintact

predatorcommunities,sowehopetheguidelinesprovidedwill

supportscience-basedmanagementprogrammesthatlead

tomoreeffectiveconservationandrestorationofbison.These

guidelinesfocusonwidespreadcommonmanagementissues—

populationmanagement,disease,andgeneticmanagement.

Theseguidelinesandprincipleswillensurethatkeyissuesare

addressed,andcitationswillhelpmanagersfindmoredetailed

informationthatmaybenecessarytoaccommodatespecific

situations.


Chapter10 GuidelinesforEcologicalRestorationofBison

Lead Authors: C. Cormack Gates, Robert O. Stephenson, Peter J.P. Gogan, Curtis H. Freese, and Kyran Kunkel

10.1 Introduction

DuringPre-Columbiatimes,bisonhadthewidestdistribution

ofanylargeherbivoreinNorthAmerica,rangingfromthe

aridgrasslandsofnorthernMexicototheextensivemeadow

systemsofInteriorAlaska(Chapters2and7).Followingthe

arrivalofEuropeans,thespeciesexperiencedunparalleledrange

contractionandcollapseofpopulationsinthewild,primarily

duringthelate19thCentury(Isenberg2000).Wildbisonpersisted

inonlytwolocations,southofGreatSlaveLakeinwhatisnow

WoodBuffaloNationalPark(about300individuals),andinthe

remotePelicanValleyintheAbsarokaMountainsintheinterior

ofYellowstoneNationalPark(YNP)(fewerthan30individuals).

Thespecieswasextirpatedfromthewildthroughoutthe

remainderofitsoriginalrange.TheAmericanbisonhasachieved

aremarkablenumericalrecovery,fromapproximately500atthe

endofthe19thCenturytoabouthalfamillionanimalstoday,of

which93%nowexistundercaptivecommercialpropagation

(Chapter7).However,Sandersonet al. (2008)estimatethatbison

occupylessthan1%oftheiroriginalrange.

RarelydowildlifepopulationsinNorthAmericaachievethe

fullrangeofecologicalinteractionsandsocialvaluesexisting

priortoEuropeansettlement.Thebisonremainsextirpatedas

wildlifeandintheecologicalsensefrommuchofitsoriginal

continentalrange.Thisisparticularlytrueoftheplainsbison,

forwhichfewpopulationsinteractwiththefullsuiteofother

nativespeciesandenvironmentallimitingfactors(Chapters6

and7).Intheabsenceofcommittedactionbygovernments

(includingaboriginalgovernments),conservationorganisations,

andperhapsthecommercialbisonindustry,theconservationof

bisonasawildspeciesisfarfromsecure.Themainchallenges

weredescribedinearlierchaptersofthisvolumeandare

summarisedbyFreeseet al. (2007).Theyincludeanthropogenic

selectionandothertypesofintensivemanagementofcaptive

herds,smallpopulationsizeeffects,issuesrelatedtoexotic

diseases,introgressionofcattlegenes,managementunder

simplifiedagriculturalproductionsystems,andassociatedwith

this,widespreadecologicalextinctionasaninteractivespecies.

Contemporarybiologicalconservationisfoundedonthe

premiseofmaintainingthepotentialforecologicaladaptation

inviablepopulationsinthewild(IUCN2003;Secretariatof

theConventiononBiologicalDiversity1992;Soulé1987),and

maintaininginteractivespecies(Souleet al. 2003).Viability

relatestothecapacityofapopulationtomaintainitselfwithout

significantdemographicorgeneticmanipulationbyhumans

fortheforeseeablefuture(Soulé1987).Forlimitingfactors,

suchaspredationandseasonalresourcelimitation,adaptation

requiresinteractionsamongspecies,betweentrophiclevels,

withphysicalelementsofanecosystem.These,andother

interactionsamongindividualswithinapopulation(e.g.,resource

andmatecompetition),contributetomaintainingbehavioural

wildness,morphologicalandphysiologicaladaptations,fitness,

andgeneticdiversity.Thesefactorsenableaspeciestoadapt,

evolve,andpersistinanaturalsettingwithouthumansupportin

thelongterm(Knowleset al. 1998).

Viable,wildpopulationsofbison,subjecttothefullrange

ofnaturallimitingfactors,areofpre-eminentimportanceto

thelong-termconservation,globalsecurity,andcontinued

evolutionofthespeciesaswildlife.However,theavailability

ofextensiveecosystemscapableofsustaininglarge,free-

roaming,ecologicallyinteractivebisonpopulationsislimited.

Thisisparticularlytrueintheoriginalrangeofplainsbisoninthe

southernagriculture-dominatedregionsofthecontinent,given

thehistoricalpost-Europeansettlementpatternsofindustrialand

post-industrialsociety.Socialandpoliticalsystemsthatprovide

spaceandenvironmentalconditionswherebisoncancontinue

toexistaswildlifeandevolveasaspecies,areseverelylimited.

Innovativeapproachesneedtobeinstigatedinsomelocations

toemulate,totheextentpossible,theoriginalecological

conditions,andtopreventdomesticationandsmallpopulation-

relateddeleteriouseffectssuchasthoseexperiencedbythe

Europeanbison(HartlandPucek1994;Prior2005;Puceket

al. 2004).Currently,thereisonlyonepopulationofplainsbison

(YNP)andthreepopulationsofwoodbison(GreaterWood

BuffaloNationalPark,Mackenzie,andNislingRiver)inNorth

Americathatcanbeconsideredecologicallyrestored(thousands

ofindividuals,largelandscapes,allnaturallimitingfactors

present,minimalinterference/managementbyhumans).

TheconservationofAmericanbisonaswildlifewouldbe

significantlyenhancedbyestablishingadditionallarge

populationstoachievelandscapescaleecologicalrestoration.

Thiswillrequireeffectivecollaborationamongavarietyof

stakeholders,wherebylocalactions,baseduponsocialand

scientificinformation,arecoordinatedwithwidergoalsfor

speciesandecosystemconservation.Thebisonwasan


ecologicallydominantkeystonespeciesovermuchofitsrange.

Thustheecologicalintegrityanddiversityofecosystemsin

whichtheyoccurred,whetherdefinedhistoricallyorbiologically,

willdependonlarge-scalerestorationofthebison.

10.2 Ecological restoration

Ecologicalrestorationprovidesaconceptualframeworkfor

bisonrestorationatmediumtobroadscales.Itcanbedefined

astheintentionalprocessofassistingtherecoveryofan

ecosystemthathasbeendegraded,damaged,ordestroyed

relativetoareferencestateoratrajectorythroughtime(SERI

andIUCNCommissiononEcosystemManagement2004).The

goalofecologicalrestorationisanecosystemthatisresilientto

perturbation,isself-sustainingwithrespecttostructure,species

compositionandfunction,isintegratedintolargelandscapes,

andsupportingsustainablehumanlivelihoods.Manyhealthy

ecosystemsareaproductofhumanendeavoursoververylong

timeperiods.Inmanycasesthen,ecologicalrestorationprojects

typicallyrequirestheparticipationofresource-dependanthuman

communities,andhavethepotentialtosupportecologically

sustainableeconomiesinruralcommunities.Bisonplay

importantecologicalroles(Chapter6),aswellasmeaningful

culturalandeconomicroles(Chapters2and7).Theyare

increasinglyprovidingaviablealternativetograzingexotic

domesticherbivores(Reneckeret al. 1989).

Sandersonet al. (2008)assertedthatbysharinganinclusive,

affirmativeandspecificvisionandknowledgeaboutbisonand

landscapeconservationwithawiderangeofstakeholders,

opportunitiescanbecreatedtorestorebisoninecologically

effectiveherdsroamingacrossextensivelandscapesinall

majorhabitatsoftheiroriginalrange.Herewedefinethefull,or

Ecological restoration of bison: the re-

establishment of a population of several

thousand individuals of the appropriate

sub-species, in an area of their original

range, in which bison interact in

ecologically significant ways with the

fullest possible set of other native species

and other biophysical elements of the

landscape, and connect in meaningful ways

with human communities, with minimal

management interventions (adapted from

Sanderson et al. 2008).

ideal,ecologicalrestorationofbisonasthere-establishmentof

apopulationofseveralthousandindividualsoftheappropriate

sub-species,inanareaoforiginalrange,inwhichbisoninteract

inecologicallysignificantwayswiththefullestpossiblesetof

othernativespeciesandbiophysicalelementsofthelandscape,

withminimalmanagementinterventions.Thisisnottosay

thatpopulationssmallerthanseveralthousandbisondonot

contributetobisonconservation,ortorestorationofecological

processes(e.g.,grazing,soildisturbance,decomposition,

nutrientcycling,predation,scavenging;Chapter6).However,

someprocesses,suchasmigrationandnaturalselection,

maybeabsentornotfunctionascompletelyatsmallerscales

(Chapter9).Sandersonet al. (2008)providespecificcriteriafor

rankingthecontributionofbisonherdstoecologicalrestoration.

10.2.1 Geographic potential for ecological restoration

TheWildlifeConservationSocietyhostedaworkshopinMay

2006atVermejoParkRanch,NewMexicothatinvolved28

people,includingbisonspecialists,indigenousgroups,bison

producers,conservationorganisations,andgovernment

andprivatelandmanagers,fromthroughoutNorthAmerica.

Amongotherobjectives,participantsworkedtodraftavision

forecologicalrecoveryoftheAmericanbison,todevelopa

consensushypothesisonmajorhabitattypeswithintheoriginal

rangethatwouldbeusefulforrepresentativeconservation

planning,andtomapareasforpotentialecologicalrecoveryover

thenext20,50,and100years(Sandersonet al. 2008;alsosee

Chapter7).Themethodsusedtoachievetheseobjectiveswere

similartothosepioneeredforjaguars(Sandersonet al. 2002)

andsubsequentlyappliedtootherspecies(e.g.,Thorbjarnarson

et al. 2006)underthetitleof“range-widepriority-setting”.

Avisionreferredtoas“TheVermejoStatement”wasdeveloped

fortheecologicalfutureoftheAmericanbison(Sandersonet al.

2008):

“Over the next century, the ecological recovery of the North

American bison will occur when multiple large herds move freely

across extensive landscapes within all major habitats of their

historic range, interacting in ecologically significant ways with

the fullest possible set of other native species, and inspiring,

sustaining and connecting human cultures.

This vision will be realised through a collaborative process

engaging a broad range of public, private, and indigenous

partners who contribute to bison recovery by:

• Maintaining herds that meet the criteria for ecological recovery, as well as herds that contribute in some significant way to the overall vision, regardless of size,

• Managing herds for the long-term maintenance of health, genetic diversity, and integrity of the species,

• Restoring native ecosystems, ecological interactions, and species,

“‘Ecosystem’ means

a dynamic complex

of plant, animal and

micro-organism

communities and their

non-living environment

interacting as a

functional unit” (Article

2 of the Convention on

Biological Diversity).


• Providing conservation incentives for bison producers, managers, and other stakeholders,

• Creating education, awareness and outreach programmes to public and policy-making constituencies,

• Building capacity among key stakeholder groups, and

• Working across international borders, where necessary.”

ParticipantsintheVermejoworkshopwereaskedtomapareas

where “ecological recovery might be possible”overthreetime

frames(20,50,and100years),consideringfuturetrendsinland

use,economicdevelopment,demography,andclimate.The

resultingmapsprovideasubjective,visualhypothesisofthe

mostpromisingplacesforecologicalrecovery(Sandersonet al.

2008).Themapsillustratethatpotentialforecologicalrecovery

existsthroughoutNorthAmerica.Long-termopportunities

areapparentacrossmuchoftheoriginalrangeoftheplains

bison,fromprivateagricultural,state,andnationalgrazing

landsinnorthernMexicoandsouthernNewMexico,tothe

agriculture-dominated,mixedtenurelandscapesoftheNorthern

GreatPlains.Innorthernregionsofthecontinent,woodbison

populationsexceedingathousandanimalsarealreadypresent

inthreelargelandscapesinCanada,andanewinitiativewill

restoreoneormorepopulationsininteriorAlaska.

Thekindsoflargeareasrequiredtoachieveidealecological

restorationofbisonarelikelytobemanagedbyseveral

jurisdictions,andmayalsoinvolveprivatelandowners.Achieving

agreementonrestoringbisontosuchlandscapesischallenging

prospect,requiringprincipled,long-termdevelopmentplanning,

soundlybasedoncommunity-basedconservationdevelopment

praxis(see:BoppandBopp2006,forpracticalguidelinesfor

communitydevelopment).

10.2.2 principles for ecological restoration applicable to bison

Successfulecologicalrestorationofbisonas

wildlifeonmulti-tenuredlandscapesrequires

carefulassessmentandcollaborativeplanning.

Whilesomerestorationprojectswillemergefrom

governmentandnon-profitorganisationinitiatives,

privatelandownersmayinitiateothers.Inmany

cases,assemblingasufficientlylargelandscape

(tensorhundredsofthousandsofhectares)for

ecologicalrestorationwillrequirecooperation

betweenpublicandprivatelandowners.

TheAmericanBisonSpecialistGroupconsidereddocuments

publishedbyIUCNandtheSocietyforEcologicalRestoration

ScienceandthePolicyWorkingGroup,anddrewuponthe

professionalandpracticalexperiencesofitsmembers,and

otherparticipants,todevelopthefollowingguidingprinciplesfor

agenciesandnon-profitconservationorganisationsinterestedin

ecologicalrestorationofbison:

1) Goalsconcerningthemanagementofland,water,and

livingresources,includingbisonrestoration,areamatterof

societalchoice.

2) Ecologicalrestorationofbisonisaninterdisciplinaryand

inclusiveundertakingrequiringtheinvolvementofall

relevantsectorsofsocietyandscientificdisciplines.

3) Planningandmanagementofecologicalrestoration

projectsshouldbedecentralisedtothelowestappropriate

level,ascloseaspossibletothehumancommunitywithin

alocalecosystem,andsupportedbythehighestlevelsof

governmentpolicy.

4) Allformsofrelevantinformation,includingscientific,

indigenousandlocalknowledge,andinnovations

andpractices,shouldbeconsideredinplanningand

implementingbisonrestoration.

5) Understandingandaddressingeconomicdriversis

imperativeforsuccessfulecologicalrestorationofbison,

including:

a. Reducingmarketdistortionsthatadverselyaffect

conservationofbisonaswildlife;

b. Developingincentivestopromoteconservationof

ecologicallyfunctioningbisonpopulationsandtheir

sustainableuses;and

c. Totheextentpossible,internalisingthecosts

andbenefitsofmanagingbisonaswildlifeinan

ecologicallyrestoredlandscape.

6) Ecologicalrestorationofbisonshouldbeundertakenat

appropriatespatialandtemporalscales,andshouldfocus

onrestoringecologicalstructure,processes,functions,and

interactionswithinadefinedecosystem.

“A functional conservation area maintains the focal

species, communities, and/or systems, and their

supporting ecological processes within their natural

ranges of variability (i.e., the amount of fluctuation

expected in biodiversity patterns and ecological

processes under minimal or no influence from human

activities)” (poiani and richter undated).

ideal,ecologicalrestorationofbisonasthere-establishmentof

apopulationofseveralthousandindividualsoftheappropriate

sub-species,inanareaoforiginalrange,inwhichbisoninteract

inecologicallysignificantwayswiththefullestpossiblesetof

othernativespeciesandbiophysicalelementsofthelandscape,

withminimalmanagementinterventions.Thisisnottosay

thatpopulationssmallerthanseveralthousandbisondonot

contributetobisonconservation,ortorestorationofecological

processes(e.g.,grazing,soildisturbance,decomposition,

nutrientcycling,predation,scavenging;Chapter6).However,

someprocesses,suchasmigrationandnaturalselection,

maybeabsentornotfunctionascompletelyatsmallerscales

(Chapter9).Sandersonet al. (2008)providespecificcriteriafor

rankingthecontributionofbisonherdstoecologicalrestoration.

10.2.1 Geographic potential for ecological restoration

TheWildlifeConservationSocietyhostedaworkshopinMay

2006atVermejoParkRanch,NewMexicothatinvolved28

people,includingbisonspecialists,indigenousgroups,bison

producers,conservationorganisations,andgovernment

andprivatelandmanagers,fromthroughoutNorthAmerica.

Amongotherobjectives,participantsworkedtodraftavision

forecologicalrecoveryoftheAmericanbison,todevelopa

consensushypothesisonmajorhabitattypeswithintheoriginal

rangethatwouldbeusefulforrepresentativeconservation

planning,andtomapareasforpotentialecologicalrecoveryover

thenext20,50,and100years(Sandersonet al. 2008;alsosee

Chapter7).Themethodsusedtoachievetheseobjectiveswere

similartothosepioneeredforjaguars(Sandersonet al. 2002)

andsubsequentlyappliedtootherspecies(e.g.,Thorbjarnarson

et al. 2006)underthetitleof“range-widepriority-setting”.

Avisionreferredtoas“TheVermejoStatement”wasdeveloped

fortheecologicalfutureoftheAmericanbison(Sandersonet al.

2008):

“Over the next century, the ecological recovery of the North

American bison will occur when multiple large herds move freely

across extensive landscapes within all major habitats of their

historic range, interacting in ecologically significant ways with

the fullest possible set of other native species, and inspiring,

sustaining and connecting human cultures.

This vision will be realised through a collaborative process

engaging a broad range of public, private, and indigenous

partners who contribute to bison recovery by:

• Maintaining herds that meet the criteria for ecological recovery, as well as herds that contribute in some significant way to the overall vision, regardless of size,

• Managing herds for the long-term maintenance of health, genetic diversity, and integrity of the species,

• Restoring native ecosystems, ecological interactions, and species,

“‘Ecosystem’ means

a dynamic complex

of plant, animal and

micro-organism

communities and their

non-living environment

interacting as a

functional unit” (Article

2 of the Convention on

Biological Diversity).


7) Restoredbisonpopulationsshouldbemanaged,tothe

extentpossible,asanintegralcomponentof,andwithin

theecologicallimitsof,anecosystem.

8) Conservingbisonandconservinglandscapesthrough

restorationofecologicallyfunctioningbisonpopulations

areinseparable.

9) Adoptingalong-termperspectiveonecological

restorationofbison,andaninclusiveprocess,willopenup

conversationsandfosterpartnershipsandpoliticalwillthat

mightnototherwisebepossible.

10) Ecologicalrestorationofbisonshouldserveboth

biodiversityconservationandecologicallysustainableuse,

andinvolvefairandequitablesharingofbenefitsamong

stakeholders.

11) Ecologicalrestorationofbisonshouldbefullyincorporated

intonationalandstate/provincialbiodiversityconservation

strategies.

12) Inter-sectoralandinter-jurisdictionalcommunication

atalllevels(betweennations,governmentministries,

managementagencies,organisations,communities,etc.)

improvesawarenessandmulti-partycooperation.

Thebisonhasbeenautilityspeciesformanyculturesand

communitiessincepeoplefirstarrivedontheNorthAmerican

continentabout12,000yearsago,withtheexceptionofa

100-yearperiodbetweenthegreatcontractionofthespecies

(circa1880;Flores1994)andrecentcommercialisation(circa

1980;Reneckeret al. 1989).Itsutilityisreflectedinthecurrent

predominanceofanimalsmanagedforprivatecommercial

captivepropagation(about93%),andthefactthatalllarge

(morethan1,000animals)free-roamingpopulationsarehunted.

TheIUCNPolicyonSustainableUseofWildLivingResources

(http://cmsdata.iucn.org/downloads/2000_oct_sust_use_of_

wild_living_resources.pdf)andtheprinciplesonsustainable

usedevelopedbytheIUCNSustainableUseSpecialistGroup

(IUCNSUSGTechnicalAdvisoryCommittee2001)applytothe

ecologicalrestorationofbison.TheIUCNPolicyonSustainable

Useprovidesthatconservationofbiodiversityiscentraltothe

IUCN’smission,whichistoinfluence,encourage,andassist

societiestoconservetheintegrityanddiversityofnature,

andtoensurethatanyuseofnaturalresourcesisequitable

andecologicallysustainable.ThePolicyconsidersthatboth

consumptiveuses(harvestingofanimalsandplants)andnon-

consumptiveuses(maintainingculturalandaestheticvaluesof

biologicaldiversity)areimportantcomponentsofasustainable

developmentagendasupportinghumanlivelihoods,while,at

thesametime,contributingtoconservation.Inaddition,the

IUCNRe-introductionSpecialistGroup(1998)offeredimportant

considerationstoensurelocalstakeholderandagencysupport

forwildliferestorationprojects.

principles for Sustainable Use of Living resources (IUCN Sustainable Use

Specialist Group 2001):

1. Sustainable use will most likely be achieved

with consideration of socio-political,

economic, biological and user factors at

the community, sub-national, national, and

international levels.

2. Sustainable use is enhanced by supportive

incentives, policies, laws and institutions at all

levels of governance, and by effective linkages

between them.

3. Local communities, and other parties who

have management responsibility for wild living

natural resources, must be supported by

recognised rights and the means to manage

the resources.

4. the contribution and needs of those who

manage wild living natural resources must be

appropriately reflected in the allocation of the

benefits from the use of those resources.

5. Adaptive management, relying on an iterative

process of timely and transparent feedback

from socio-economic, resource and ecological

monitoring, is essential for sustainable use.

6. Sustainability of living wild resource use is

enhanced if traditional/local knowledge is

taken into account.

7. Sustainable use of wild living resources is

enhanced if managerial jurisdictions match

ecological and socio-economic scales.

8. Subsidies that distort markets, promote

habitat alteration or destruction, and

unsustainable use of natural resources should

be eliminated.


10.3 the “Ecosystem Approach” for Designing Ecological restoration of Bison

TheEcosystemApproach(Shepherd2004)isastrategyfor

integratedmanagementofland,water,andlivingresourcesthat

promotesconservationandsustainableuseinanequitableway.

ItistheprimaryframeworkforactionundertheConventionon

BiologicalDiversity.TheEcosystemApproachputspeople,and

theirnaturalresourceusepractices,atthecentreofdecision-

making.Becauseofthis,itcanbeusedtoseekanappropriate

balancebetweenconservationanduseofbiologicaldiversity

inareaswheretherearemanyresourceuserscombinedwith

importantnaturalvalues.

Planningandimplementingecologicalrestorationofbisonmay

involvemulti-tenuredlandscapesandisacomplexundertaking

thatrequiresassessingbiophysicalandsocialcomponents,

evaluatingandengagingstakeholders,consideringeconomic

conditions,andcultivatinglong-termpartnerships.Ecological

restorationplanningisadynamicprocess,bestachieved

incrementally,withampleopportunitiesforiterationand

feedback.Thefollowingelementsprovideguidanceforagencies,

organisationsandindividualsinterestedindesigningecological

restorationprojects.

10.3.1 Defining the biological landscape and objectives

Ecologicalrestorationofbisonconsidersthespeciesasan

interactiveelementofanecologicallyfunctioningrestoration

areathatprovidesthesizeanddistributionofhabitatsnecessary

tosupportarestoredbisonpopulation.Definingabiological

landscapeforthispurposeinvolvesdeterminingthesizeand

refiningtheboundaryofthearea,identifyingtheresource

requirementsofbisonandotherfocalelementsincludingtheir

spatialneeds,andmappingthedistributionofhabitatresources

(Louckset al. 2004).Thesetaskscanbeachievedbyavarietyof

processesincludingexpert-drivenworkshopsandlocalworking

groupsaidedbytechnicalexperts.Forexample,theIUCN/SSC

ConservationBreedingSpecialistGroup(CBSG)hasextensive

experiencemanagingconservationplanningworkshopsusing

itssignatureprocesses,theConservationActionManagement

PlanandPopulationandHabitatViabilityAssessment(PHVA),

toassistgroupsindevelopingspecieslevelactionplans(www.

cbsg.org/cbsg).ALandscapeCumulativeEffectsSimulator

(ALCES®;ForemTechnologies;www.alces.ca)isanother

softwaretoolthatisrapidlygainingacceptancebyindustry,

government,andthepublicasaneffectivesimulationtoolfor

exploringsustainableresourceandlandscapemanagement

alternatives.

Whateverthedecision

supportsystemisused,

commontoeachprocess

istheneedtohave

stakeholders(conservation

groups,wildlifebiologists,

relevantgovernment

agencies,andlocalprivate

andpubliclandmanagers)

involved.Agreementsare

typicallyrequiredonthe

sizeandboundariesof

theecosystemandthe

potentialbiologicalcapacityoftheareatomeettheneeds

ofbisonrestorationandotherconservationandcommunity

objectives.

10.3.2 Defining the social landscape, the main stakeholders, and cultivating partnerships

Large-scaleecologicalrestorationinvolvesmultiplelevels

ofsocialcomplexity,andtypicallyinvolvesmorethanone

jurisdiction.Thegeographicpotentialforecologicalrestoration

ofbisoninNorthAmericaisillustratedinageneralsenseby

Sandersonet al. (2008).Priorityareasmaybeconsideredas

havingthepotentialtobecomeconservationlandscapes(sensu

Louckset al. 2004)thathaveecologicalandsocialpotentialfor

restorationofbisonintheintermediatetolongterm.Careful

assessmentandunderstandingofsocial,economic,legal,and

politicalconditionswithincandidatelandscapesisanessential

preparatorystepforplanningandimplementingrestoration

projects(Louckset al. 2004;TheNatureConservancy2005),

particularlywherecommunitysupportandinvolvementis

required(ChildandLyman2005).

ThepriorityareasidentifiedbySandersonet al. (2008)represent,

inthecollectiveopinionofagroupofexperts,ahypothesisof

wherethemostpromisingplacesforecologicalrecoveryexist,

consideringfuturelandusetrends,economicforces,human

demography,andclimate.Understandingtheregionalsocial-

ecologicalsysteminsuchtargetareasisanimportantfeatureof

effectiveconservationplanning(Driveret al. 2003).Inadditionto

assessingthebiophysicalcapabilityofacandidatearea,detailed

assessmentsarerequiredtodefinethehumancommunitywithin

theecosystemboundaries.Sociallandscapeanalysis(Fieldet

al. 2003)providesatoolforunderstandingandmappingthe

humanlandscape.Itrequirescollecting,analysingandmapping

humandemographicandeconomicdata,andinformationon

landdevelopmentandownershippatternsandtrends.Social

landscapesconsistofthedemographicpatternsofpeople

(location,density,ageandgenderstructure,industryand

employmentpatterns,andgovernanceboundaries)inrelationto

landandresources.

“Conservation

landscape” refers

to a spatial plan for

a priority area that

meets fundamental

conservation objectives

while addressing other

socio-economic needs

(Loucks et al. 2004).


Thetypesofsocioeconomicdatarelevantforecologicalrestoration

planningwillvaryamonglocationsacrossthecontinent.However,

certaininformationisrelevantforalllandscapes.Detailedand

currentinformationonlanduse,includinglandusemaps,iscritical

forassessingtheimpactsofhabitatlossandtrends.Development

plansandtargetsforimportantresourcesectors(agriculture,

energy,andtransportation)providethebasisforevaluating

impactsofforeseeablechangeovertime.Spatialinformationon

landownershipandmanagementauthoritiescontributetothe

identificationofstakeholdersandassessmentofconservation

potential.

Louckset al. (2004)providedthefollowinglistofsocioeconomic

variablesusefulforconservationplanning.Thelistshouldbe

reviewedandcustomisedforeachprojectinconsultationwith

localmanagers:

1. Current patterns of land and resource use:

• Majorlandandresourceuses(includingforest,water,wildlifeuse,agriculture,extraction);

• Developmentplansandprojectedchangesinlandandresourceuse;

• Existingzoningregulations;

• Majorexistingandplannedinfrastructure(roads,dams,etc.);

• Existingprotectedareas.

2. Governance and land/resource ownership and

management:

• Politicalboundaries(provinces,districts);

• Landtenure(private,public,ancestral/communalareas);

• Agenciesresponsibleformanagementofland/resourceareas(e.g.,forest,agriculturedepartments).

3. population data:

• Humanpopulationdensityandgrowth;

• Populationcentres;

• Migrationpatterns(in-andout-migration);

• Socialcharacteristics:income,ethnicity,indigenousareas;

• Economicdata;

• Economicgrowthandlossareas;

• Landprices;

• Potentialvaluesandopportunitiesforecologicalservices;

• Potentialforincorporatingnaturalassetsintothelocaleconomy.

4. Additional factors that affect biodiversity and

potential for bison restoration:

• Access(e.g.roads,rivers,energycorridors,etc.);

• Trendsinhabitatconversion.

Bisonoccupyadistinct

iconicstatusaswildlife

withbothindigenous

andnon-indigenous

NorthAmericans.The

culturalandhistoric

significanceofbisonis

particularlyimportant

tomanyNativeNorth

Americans(Stephenson

et al. 2001;Wyckoff

andDalquest1997).In

recentdecades,bison

haveincreasedinvalue

asprivatepropertyinthe

formoflivestock(Chapter

7).Inthegrasslands

ofthecontinent,the

cattleranchingcultureandeconomyreplaceda10,000-year-

oldbisoneconomy,andcattleranchingnowoccupiesmore

than95%oftheGreatPlainsgrasslands.Thepotentialfor

restorationofplainsbisonatameaningfulecologicalscalein

thisregionthereforedependsonsupportbypeopleinvolvedin

thissector.Similarly,supportfromregulatoryauthorities,and

harmonisationofpoliciesandplanningprocessesisnecessary

toensureafeasiblestart,andsustainableoutcomesofbison

conservationprojects.

Toignoreorcontradictculturalorlocalinterests,ortheauthority

ofagencies,cangenerateunnecessaryon-goingresistance

toconservationinitiatives.An

exampleofthisistheconcept

ofthe“BuffaloCommons”or

“re-bisoning”oftheGreatPlains

proposedbyRutgersUniversity

geographersFrankandDeborah

Popper(PopperandPopper

1987).ThePopper’spredicted

economicandhumanpopulation

declinesintheGreatPlains,

nowborneoutbycurrenttrends

(Forrestet al. 2004).Theidea

ofreplacingthecattleranching

culturewithaBuffaloCommons

createdafirestormofprotestamongagriculture-based

communitiesintheregion,andcontinuestohauntdiscussions

aboutbisonconservationandecologicalrestoration.Thegeneral

“Current conservation

initiatives—parks, land

conservation, regulatory

programs—offer

important contributions

but provide solutions

to only 10% of the

problem. the remaining

90% exist at the

interface of human

populations and

ecological systems”

(Child and Lyman 2005).

Stakeholders are

people who will be

impacted by the

decisions; they

have the knowledge

to make the best

decisions, and the

power to implement

or block decisions.


lessonlearnedfromthiscaseisthattheecologicalrestorationof

bisonisnotpossibleanywherewithoutengagingstakeholders,

theirinterests,mandatesandaspirations,anddevelopinglocal

communityandagencycapacitytoengageinsustainable

ecologicalrestoration.

Managingsocial-ecologicalsystemsrequiresanexplicit

approachthatcanserveasavisionforstakeholders(Knight

et al. 2006).Conservationplannersshouldavoidperceiving

themselvesasempiriciststhatoperateoutside,ratherthan

within,social-ecologicalsystems(SayerandCampbell2004).

ClewellandAronson(2006)discussthemajormotivations

orrationalesfortherestorationofecosystemsandtheir

associatedspecies.Theseincludetechnocratic,biotic,heuristic,

idealistic,andpragmaticrationalesthatoftenresultinsocial

conflicts.Restorationofbisonandtheirnativeecosystemsis

noexception,asadiversityofsocioeconomicfactors,from

localtoregionaltointernationallevels,isinvolved.Organisers

wishingtoinitiatelargescaleecologicalrestorationprojectsare

encouragedtobecomefamiliarwiththetheoriesandpractices

ofcommunity-basedresourcemanagement(ChildandLyman

2005)andcommunitydevelopment(BoppandBopp2006),but

moreimportantly,toincludeanexperiencedpractitioneronthe

coredevelopmentteam.

Althoughbisonrestorationpresentsmanychallenges,itis

importanttorememberthatbisonhavehistoricallyprovided

manybenefitstohumansocietiesandcontinuetodosotoday.

Incollaborativeplanningforecologicalrestoration,itisimportant

toemphasiseeconomicandsocialbenefits,aswellasthose

relatedtobiodiversityconservationandecosystemhealth.

10.4 Guidelines for planning and Implementing Ecological restoration projects for Bison

TheIUCNRe-IntroductionSpecialistGroup(1998)definesthe

purposeofare-introductioninthefollowingmanner:

“The principle aim of a re-introduction should be to establish

a viable, free-ranging population in the wild, of a species,

subspecies or race, which has become globally or locally extinct,

or extirpated, in the wild. It should be re-introduced within the

species’ former natural habitat and range and should require

minimal long-term management.”

Ecologicalrestorationaddsadditionalvaluestospecies’

reintroductionprojects.Ithasasitsgoal,anecosystem

thatisresilientandself-sustainingwithrespecttostructure,

speciescompositionandfunction,aswellasbeingintegrated

intothelargerlandscapeandsupportingsustainablehuman

livelihoods(SERIandIUCNCommissiononEcosystem

Management2004).Thefollowingguidelinesforplanningand

implementinganecologicalrestorationprojectforbisonwere

adaptedfromtheIUCNRe-introductionGuidelines(IUCN

1998).Theyarealsoinformedbyotherkeydocumentson

conservationandrestorationplanning(Louckset al.2004;

TheNatureConservancy2005),communitybasednatural

resourcemanagement(ChildandLyman2005),andcommunity

developmentplanning(BoppandBopp2006).Theyaddress

biologicalandsocio-economicneedsforrestoringbisonasan

interactivespecieswithinarestoredecosystem:

10.4.1.1 Feasibility assessment

• Sitesforecologicalrestorationofbisonshouldbewithintheoriginalrangeoftheappropriatesub-speciesofbison;

• Forare-introduction,thereshouldbenoremnantpopulationofbisoninordertopreventdiseasepropagation,socialdisruption,introductionofaliengenes,ordisruptionstologistics;

• Insomecircumstances,are-introductionorreinforcementmayhavetobemadeintoanareathatisfencedorotherwisedelimited,butitshouldbewithinthesub-species’originalrangeandhabitat;

• Ecologicalrestorationmaytakeplacewheretheannualhabitatandlandscaperequirementsofmorethan1,000bisoncanbesatisfiednormally,withouttheneedforsupplementation,andapopulationofatleastthisnumberislikelytobesustainedfortheforeseeablefuturewithminimummanagementintervention.

• Thepossibilityofnaturalhabitatchangeshouldbeconsidered(e.g.forestsuccession,climatechange);

• Theeffectsofinteractionsofbisonwithotherspeciesintheecosystemshouldbedefinedandconsideredinplanningtherestorationproject;

re-introduction: an attempt to re-establish

bison in an area that was once part of its original

range, but from which it was extirpated.

re-enforcement/Supplementation/

Augmentation: Addition of individuals to an

existing population of conspecifics.

Substitution: the introduction of a closely related

species or sub-species, for subspecies that have

become extinct in the wild and in captivity. the

introduction occurs in suitable habitat within the

extinct species or subspecies historical range

(Seddon and Soorae 1999).

Source: IUCN re-introduction Guidelines

(re-introduction Specialist Group 1998)


• Legal,policy,political,andculturalconstraintsneedtobeevaluatedtodetermineifmitigationisneededorpossible;

• Determineifthefactorscausingdeclinecanbeeliminatedormitigated(e.g.,diseases,over-hunting,over-collection,pollution,poisoning,competitionwith,orpredationby,introducedspecies,habitatloss,adverseeffectsofearlierresearchormanagementprogrammes,competitionwithdomesticlivestock);

• Wherethereleasesitehasbeensubstantialdegradedbyhumanactivity,ahabitatrestorationprogrammeshouldbeinitiatedbeforethereintroductioniscarriedout;

• APopulationandHabitatViabilityAssessmentwillaidinidentifyingsignificantenvironmentalandpopulationvariables,andassessingtheirpotentialinteractions,whichcanguidelong-termpopulationmanagement;

• A prioriagreementisdesirableonpopulationobjectives,monitoring,andmethodsthatwillbeusedtomanagepopulationgrowthasthetargetpopulationsizeisapproached;

• Similarly,a prioriagreementonrangehealthobjectivesandrangemonitoringandmanagementmethodsisdesirable;

• Determinetheavailabilityofsuitablestock,includingsubspeciesorlocallyadaptedforms,genetics(e.g.cattlegenes),andabsenceofspecificdiseasesofconcerntoconservation;

• Afeasibilityassessmentshouldincludedeterminingifadequatefundingisavailabletosuccessfullycompletetheproject.

10.4.1.2 Suitable release stock

• Itispreferablethatsourceanimalscomefromwildpopulations,orcaptivestockthathavebeensubjectedtominimummanagement,suchasselectionfororagainstspecificmorphologicaltraits;

• Thesourcepopulationshouldideallybecloselyrelatedgeneticallytotheoriginalnativestockandshowsimilarecologicalcharacteristics(morphology,physiology,behaviour,habitatpreferences)totheoriginalsub-population;

• Usestockfromasourcepopulation(s)thathastestednegativeforthepresenceofcattlegenemarkers,basedonthebestavailabletechnology;

• Stockmustbeguaranteedavailableonaregularandpredictablebasis,meetingspecificationsoftheprojectprotocol;

• Individualsshouldonlyberemovedfromawildpopulationaftertheeffectsoftranslocationonthedonorpopulationhavebeenassessedandafteritiscertainthattheseeffectswillnotbenegative;

• Ifcaptiveorartificiallypropagatedstockistobeused,itmustbefromapopulationthathasbeensoundlymanagedbothdemographicallyandgenetically,accordingtotheprinciplesofcontemporaryconservationbiology;

• Re-introductionsshouldnotbecarriedoutmerelybecausecaptivestocksexist,norsolelyasameansofdisposingofsurplusstock;

• Prospectivereleasestock,includingstockthatisagiftbetweengovernments,mustbesubjectedtoathoroughveterinaryscreeningprocessforpathogensandexposuretopathogensbeforeshipmentfromoriginalsource;

• Ifevidenceofinfectionwithanynotablepathogenisfound,thetranslocationshouldbestoppedandariskassessmentconductedtodeterminethewisestaction;

• Assessthepresenceofpathogensinwildanddomesticspeciespresentinthere-introductionarea;

• Minimisetheriskofinfectionduringtransportbymanagingpotentialexposuretopathogens;

• Stockmustmeetallhealthregulationsprescribedbytheveterinaryauthoritiesoftherecipientjurisdictionandadequateprovisionsmustbemadeforquarantineifnecessary;

• Ifvaccinationisdeemedappropriatepriortoreleasethismustbecarriedoutallowingsufficienttimefortherequiredimmunitytodevelopbeforethetranslocation.

10.4.1.3 preparation and release

• Constructamultidisciplinaryplanningandmanagementteam(s)withaccesstoexperttechnicaladviceforallphasesoftheprogramme;

• Establishshort-andlong-termgoalsandspecificobjectives,bothforthebisonpopulationandforthehabitatandbiodiversitymanagement,includingsuccessindicatorsandtargets;

• Definemonitoringprogrammesforevaluatinghowwellgoalsandobjectivesarebeingmet,andtheadjustmentsthatmayberequired.Eachre-introductionshouldbeacarefullydesignedexperiment,withthecapabilitytotestmethodologywithscientificallycollecteddata;

• Secureadequatefundingforallphasesofpreparationandrelease;

• Monitorthehealthandsurvivalofindividuals;

• Secureappropriateveterinaryexpertisetoensurethehealthofreleasedstock,includingadequatequarantinearrangements,especiallywherestockistransportedoverlongdistancesorcrossesjurisdictionalboundaries;

• Developtransportplansfordeliveryofstocktothesiteofreintroduction,withspecialemphasisonwaystominimisestressontheindividualsduringtransport;

• Determineappropriatereleasestrategies,includinghabituationofreleasestocktotheprojectarea,behaviouraltraining,releasetechniques,andtiming;

• Establishpoliciesoninterventionstomanageparasitesandpathogens;

• Establish,wherenecessary,adetailedcontainmentprogrammethatincludesfencedesignandmonitoring


andprotocolsfordealingwithescapedanimals;

• Interventions(e.g.,supplementalfeeding,veterinaryaid,horticulturalaid)shouldonlybeundertakenifnecessarytopreventcatastrophiclossesthatriskextirpation,orasignificantreductioningeneticdiversity,particularlywhenthepopulationissmall;

• Iffencingisrequired,usedesignsthatallowformovementofotherwildlifespecies(seeChapter9forspecifications);

• Developaconservationawarenessprogrammeforsecuringlong-termsupport:professionaltrainingofindividualsinvolvedinthelong-termprogramme,publicrelationsthroughthemassmediaandinlocalcommunity,andinvolvement,wherepossible,oflocalpeopleintheprogramme.

10.4.1.4 Socio-economic and legal requirements

TheIUCNGuidelinesforRe-Introductions(IUCN1998)also

providemeasuresforaddressingsocio-economicandlegal

requirementsofre-introductionprogrammes.Theyhavebeen

adaptedhereforecologicalrestorationprojectsinvolvingbison.

Consideringthatecologicalrestorationprojectsrequirelong-

termcommitmentsoffinancialandpoliticalsupport:

• Socio-economicstudiesareneededtoassessimpacts,costsandbenefitsoftherestorationprogrammetolocalhumanpopulationsandgovernments;

• Athoroughassessmentofattitudesoflocalpeopletowardstheproposedprojectisnecessarytodevelopandsecurelong-termconservationoftherestoredpopulation;

• Therestorationprogrammeshouldbefullyunderstood,accepted,andsupportedbylocalcommunitiesandaffectedgovernmentagencies;

• Wherethesecurityofthere-introducedpopulationisatriskfromhumanactivities,measuresshouldbetakentominimisetheseintheprogrammearea;

• Thepoliciesofaffectedgovernmentagencies(atalllevels)onrestorationandbisonmanagementshouldbeassessed.Thiswillincludeevaluatingexistingmunicipal,provincial,national,andinternationallegislationandregulations,andifnecessarynegotiatingnewmeasures;

• Restorationprojectsmusttakeplacewiththefullpermissionandinvolvementofallrelevantgovernmentagencies.Thisisparticularlyimportantinrestorationprogrammesinvolvingmulti-tenurelandscapes,suchasinborderareas,inareasinvolvingmorethanonestate,orwhereare-introducedpopulationcanexpandintootherjurisdictionsorontoadjacentprivatelands;

• Aswithotherspeciesoflargeherbivore(e.g.mooseandelk),bisonposesmall,butmanageable,risksofpersonalinjuryandpropertydamage.Theserisksshouldbeminimisedandadequateprovisionmadeforawarenessand,ifnecessary,compensation;

• Ifprojectsaresituatedadjacenttointernationalorstate

boundaries,provisionsshouldbemadeformonitoringormanagingbisoncrossingtheboundaries;

• Measuresformanagingescapedoremigratingbisonshouldbeagreedtoa prioriwithownersofadjacentlands;

• Approvalbyrelevantgovernmentagenciesandlandowners,andcoordinationwithnationalandinternationalconservationorganisationsarenecessary.

10.4.1.5 Monitoring, evaluation, and adaptation

Theimplementationofanecologicalrestorationprojectdoesnot

guaranteeitsobjectiveswillbeattainedoritsgoalsachieved.

Outcomesofrestorationprojectsinvolvingcomplexsystems

canbeunpredictable.Restoredecosystemsaredynamic

andrequireevaluationovermanyyears.Inlargelandscapes,

abisonpopulationmaynotmaturedemographicallyfor30

yearsormorefollowingreleasefrommanagementcontrolor

followingreintroduction(Gateset al. 2005;Larteret al. 2000).

Environmentalfactors,suchassporadicdrought,severewinters

orpredationeffects,contributetouncertaintyofoutcomes.

Maintainingsupportforanecologicalrestorationprojectin

thelongtermrequirescontinuousevaluationofperformance

measures(indicators)thatrepresenttheecologicalinfrastructure

andfunctioningoftheecosystem,andothersthatrepresent

humancommunityneedsaboutsustainingcultureandeconomy.

Respectforbothlocalandscience-basedknowledge,coupled

withparticipatoryprocesses,ensuresthefullandequitable

engagementofthecommunities,andthattheindicators

selected,datacollected,anddecisionsmade,meettheneedsof

agenciesandlocalcommunities.

Thefollowingguidelinesformonitoring,evaluationand

adaptationareoffered:

• Post-releasemonitoringofasignificantsampleofindividualbisonisnecessarytoevaluateindividualsurvival,health,reproduction,andmovements,andtoassessthecausesandsignificanceofunanticipatedlosses(e.g.,copperorseleniumtoxicity,behaviouralnaivetytopredators)duringtheinitialyearsofaproject;

• Demographic,ecologicalandbehaviouralstudiesofthepopulationshouldbeundertakenoverthelongtermtomonitorchangesinpopulationanddistributionpatterns;

• Habitatprotectionorrestorationmaybenecessarytosupportpopulationandbiodiversityrestorationgoals;

• Publicityanddocumentationshouldbeincorporatedintoeveryrestorationprojectbecausepublishedaccountsareimportantformaintaininglong-termsupportofaproject.Regularpublicinformationreleasesandpublicationsinscientificandpopularliteratureareusefulinstruments;

• Monitoringallthecostsandafullrangeofbenefits(monetaryandnon-monetary)toprovidedocumentation


thatshowstheimpactsoftheprojectandthatfundingsupportisjustified;

• Implementadaptivemanagementproceduresasneeded.Adaptivemanagement,asarestorationstrategy,isessentialbecausewhathappensatonestageinrestorationinformsordictateswhatneedstohappennext;

• Capacitybuildingshouldbeinformedbyresultsofthemonitoringprogrammeandtargetedtowardthehighestprioritiesandweakestaspectsofmanagement.

10.5 Summary

Thenext10-20yearspresentopportunitiesforconserving

Americanbisonasawildspeciesandrestoringitasanimportant

ecologicalpresenceinmanyNorthAmericanecosystems.Taking

anecosystemapproach,whichputspeopleandtheirnatural

resourceusepracticesatthecentreofdecision-making,offers

aparadigmforbalancingthesometimescompetingdemands

ofbisonconservation,theuseofbisonandbiologicaldiversity

bypeople,andsustaininghumancommunitiesinareaswhere

therearemanyresourceuserscombinedwithimportantnatural

values.Toachieveecologicalrestorationatbroadscales(large

herdsroamingacrossvastlandscapes,atnumerouslocations)

willrequireflexibleapproachesthatcanbeadaptedtoavariety

oflegalandsocio-economicconditions.Assemblinglarge

landscapesforconservationherdswilltypicallyinvolveseveral

landtenureholders,potentiallyincludingpublicagencies,tribal

governments,non-profitprivateorganisations,andfor-profit

corporationsorindividualentrepreneurs.Diversemandates,

interests,andincentiveswillinfluencehowstakeholderschoose

tomanagelandandwildlife,includingbison.Creativenew

approachesareneededforforgingenduringpartnershipsamong

landtenureholdersforcooperativeundertakings.Strategiesmay

rangefromtop-downgovernmentprogrammestobottom-up

market-basedorcultural-basedinitiatives.Progresstowards

large-scalerestorationwillrequireamuchmoresupportive

frameworkofgovernmentpoliciesandsignificantinvestment

bybothpublicandprivatesectors.Awarenessandsubstantial

publicsupportarenecessaryatboththelocallevelwhere

restorationoccurs,andamongnationalconstituenciesfor

whomthebisonisaniconiccomponentofNorthAmerica’s

naturalandculturalheritage.Forecologicalrestorationofbison

tobesuccessful,carefulassessmentandunderstandingof

biophysical,social,economic,legal,andpoliticalconditionsare

requiredforplanningandimplementation.Thisisparticularlytrue

wherebothcommunityandagencysupportandinvolvement

arerequired.Thischapterprovidedguidelinesforplanning

andimplementinganecologicalrestorationprojectforbison,

includingfeasibilityassessment,selectionofstock,preparation

andreleasemethods,assessingsocio-economicandlegal

requirements,monitoring,evaluation,andadaptation.


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Appendix A NorthAmericanconservationherdsofbisonandtheirmanagingauthorities

State/province Site Jurisdiction Managing Authority

AB(Alberta) CanadianForcesBaseWainwright Federal-Canada DepartmentofNational

Defence

AB ElkIslandNationalPark Federal-Canada ParksCanadaAgency

AB/SK(Saskatchewan) PrimroseLakeAirWeaponsRange FederalandProvincial

DepartmentofNationalDefence;SaskatchewanEnvironment,FishandWildlifeBranch

AB WatertonLakesNationalPark Federal-Canada ParksCanadaAgency

AK(Alaska)Wrangell-St.EliasNationalParkandPreserve-ChitinaRIver

State AlaskaDepartmentofFishandGame

AKWrangell-St.EliasNationalParkandPreserve-CopperRiver

State AlaskaDepartmentofFishandGame

AK DeltaJunctionStateBisonRange State AlaskaDepartmentofFishand

Game

AK FarewellLake State AlaskaDepartmentofFishandGame

AK PopofIsland State AlaskaDepartmentofFishandGame;ShumaginCorporation

AZ(Arizona) HouseRockStateWildlifeArea State ArizonaFishandGameDepartment

AZ RaymondStateWildlifeArea State ArizonaFishandGameDepartment

BC(BritishColumbia) PinkMountainProvincialPark Provincial BritishColumbiaMinistryofWater,LandandAirProtection

CA(California) U.S.MarineCorpsBaseCampPendleton U.S.Military U.S.MarineCorps

CA SantaCatalinaIsland NGO CatalinaIslandConservancy

CI(Chihuahua) RanchoElUnoEcologicalReserve Federal-Mexico ComisiónNacionaldeÁreas

NaturalesProtegidas

CO(Colorado) DanielsPark Municipal DenverParksandRecreation

CO GeneseePark Municipal DenverParksandRecreation

CO Medano-ZapataRanch NGO TheNatureConservancy

CO RockyMountainArsenal Federal-US U.S.FishandWildlifeService

IA(Iowa) BrokenKettleGrasslands NGO TheNatureConservancy

IA NealSmithNationalWildlifeRefuge Federal-US U.S.FishandWildlifeService

IL(Illinois) FermiNationalAcceleratorLaboratory Federal-US U.S.DepartmentofEnergy

plains bison



KS(Kansas) KonzaPrairieBiologicalStation State/NGOKansasStateUniversity,DivisionofBiology;TheNatureConservancy

KS MaxwellWildlifeRefuge State KansasDepartmentofWildlifeandParks

KS SandsageBisonRange&WildlifeArea State KansasDepartmentofWildlife

andParks

KS SmokyValleyRanch NGO TheNatureConservancy

KY(Kentucky) LandBetweentheLakesNationalRecreationArea Federal-US USDAForestService

MB(Manitoba) RidingMountainNationalPark Federal-Canada ParksCanadaAgency

MN(Minnesota) BlueMoundsStatePark StateMinnesotaDepartmentofNaturalResources,DivisionofParksandRecreation

MO(Missouri) PrairieStatePark State MissouriDepartmentofNaturalResources

MT(Montana) AmericanPrairieReserve NGO AmericanPrairieFoundation

MT NationalBisonRange Federal-US U.S.FishandWildlifeService

ND(NorthDakota) CrossRanchNaturePreserve NGO TheNatureConservancy

ND Sully’sHillNationalGamePreserve(newherd) Federal-US U.S.FishandWildlifeService

ND TheodoreRooseveltNationalPark Federal-US U.S.NationalParksService

NE(Nebraska) FortNiobraraNationalWildlifeRefuge Federal-US U.S.FishandWildlifeService

NE FortRobinsonStatePark State NebraskaGameandParks

NE NiobraraValleyPreserve NGO TheNatureConservancy

NE Sully’sHillherdatFt.Niobrara(originalherd) Federal-US U.S.FishandWildlifeService

NE WildcatHillsStateRecreationArea State NebraskaGameandParks

OK(Oklahoma) TallgrassPrairiePreserve NGO TheNatureConservancy

OK WichitaMountainsNationalWildlifeRefuge Federal-US U.S.FishandWildlifeService

SD(SouthDakota) BadlandsNationalPark Federal-US U.S.NationalParkService

SD BearButteStatePark State SouthDakotaGameFishandParksDept.

SD CusterStatePark State SouthDakotaGameFishandParksDept.

SD OrdwayPrairiePreserve NGO TheNatureConservancy

SD LameJohnnyCreekRanch NGO TheNatureConservancy

SD WindCaveNationalPark Federal-US U.S.NationalParkService

SK(Saskatchewan) BuffaloPoundProvincialPark Provincial SaskatchewanEnvironment,ParksBranch

SK GrasslandsNationalPark Federal-Canada ParksCanadaAgency


plains bison (continued)


SK OldManonHisBackConservationArea NGO NatureConservancyofCanada

SK PrinceAlbertNationalPark Federal-Canada ParksCanadaAgency

TX(Texas) CaprockCanyonsStatePark/TexasStateBisonHerd State TexasParksandWildlife

Department

TX ClymerMeadowPreserve NGO/Private TheNatureConservancy;Privaterancher

UT(Utah) AntelopeIslandStatePark StateUtahDivisionofWildlifeResources,DivisionofParksandRecreation

UT BookCliffsRecreationArea State UtahDivisionofWildlifeResources

UT HenryMountains State UtahDivisionofWildlifeResources

WI(Wisconsin) SandhillWildlifeArea State WisconsinDepartmentofNaturalResources

WY(Wyoming) BearRiverStatePark State WyomingStateParksandHistoricSites

WY GrandTetonNationalPark/Nat.ElkRefuge Federal/State

U.S.NationalParkService;U.S.Fish&WildlifeService;WyomingFishandGameDepartment

WY HotSpringsStatePark State WyomingStateParksandHistoricSites

WY/MT YellowstoneNationalPark Federal/State

U.S.NationalParkService;U.S.ForestService,MontanaFish,WildlifeandParks;MontanaDepartmentofLivestock


plains bison (continued)



AB(Alberta) ElkIslandNationalPark Federal-Canada ParksCanadaAgency

AB Hay-ZamaLakesComplex Provincial GovernmentofAlberta,FishandWildlifeDivision

AB/NWT(NorthwestTerritories) WoodBuffaloNationalPark Federal-Canada ParksCanadaAgency

AK(Alaska) PortageGlacier ENGO1 AlaskanWildlifeConservationCenter

BC(BritishColumbia) EtthithunLake Provincial BritishColumbiaDepartmentofWater,LandsandAirProtection

BC NordquistFlats Provincial BritishColumbiaDepartmentofWater,LandsandAirProtection

MB(Manitoba) ChitekLake Provincial

GovernmentofManitoba,DepartmentofNaturalResources;WaterhenFirstNation

NWT MackenzieBisonSanctuary TerritorialGovernmentofNWTerritories,Resources,WildlifeandEconomicDevelopment

NWT Nahanni TerritorialGovernmentofNWTerritories,Resources,WildlifeandEconomicDevelopment

NWT SlaveRiverLowlands TerritorialGovernmentofNWTerritories,Resources,WildlifeandEconomicDevelopment

YT(YukonTerritories) Aishihik Territorial GovernmentofYukon

herds in the progress of establishment:


AK(Alaska) MintoFlats StateAlaskaDepartmentofFishandGame-currentlyheldatAlaskaWildlifeConservationCenter

Wood bison


InternatIonal UnIon for ConservatIon of natUre

WORLD HEADQUARTERSRue Mauverney 28 1196 Gland, Switzerland [email protected] Tel +41 22 999 0000Fax +41 22 999 0002www.iucn.org


IUCn/ssCamerican BisonspecialistGroup

Am

erican Bison S

tatus Survey and

Conservation G

uidelines 2010

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