Table Of ContentClifford Warwick · Phillip C. Arena
Gordon M. Burghardt Editors
Health and
Welfare of
Captive Reptiles
Second Edition
Health and Welfare of Captive Reptiles
(cid:129) (cid:129)
Clifford Warwick Phillip C. Arena
Gordon M. Burghardt
Editors
Health and Welfare
of Captive Reptiles
Second Edition
Editors
CliffordWarwick PhillipC.Arena
EmergentDiseaseFoundation CollegeofScience,Health,Engineeringand
London,UK Education,AcademicOperations,
EnvironmentalandConservationSciences
MurdochUniversity
Mandurah,WA,Australia
GordonM.Burghardt
DepartmentsofPsychology
andEcology&EvolutionaryBiology
UniversityofTennessee
Knoxville,TN,USA
ISBN978-3-030-86011-0 ISBN978-3-030-86012-7 (eBook)
https://doi.org/10.1007/978-3-030-86012-7
1stedition:#SpringerScience+BusinessMediaDordrecht1995
OriginallypublishedbyChapman&Hallin1995
PrioreditiontothisworkisISBN978-0-412-55080-5,“HealthandWelfareofCaptiveReptiles”,edited
byCliffordWarwick,FredricL.FryeandJamesB.Murphy.
#SpringerNatureSwitzerlandAG1995,2023
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Design&adaptation:PhillipC.Arena,GordonM.Burghardt,CliffordWarwick
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Acknowledgements
The editors would like to express their deep appreciation to all authors for their
contributions to this volume. All who have been involved in this project have so
generouslyparticipatedintheover-archinginterestsofreptilewelfare,whichisboth
gratifying and inspiring. A scientific book is defined not only by its content and
authorsbutalsobyitsreviewers.Accordingly,theeditorsalsowishtoconveytheir
enduring gratitude to the following for their expertise, dedication, and guidance
across the breadth of chapters and issues therein: Lydia Acaralp-Rehnberg,
Stephanie Allard, Rocio Aguilar, Melissa Amarello, Kevin Arbuckle, Meredith
J. Bashaw, George Bates, Dale DeNardo, Melanie Denomme, Bob Doneley, Mat-
thewEvans,MariekeGartner,DeonGilbert,SteveGorzula,RayHuey,TimJessup,
Robert Johnson, Michael Kreger, Angelo J. L. Lambiris, La'Toya V. Latney,
Mathias Loertscher, Kevin Markwell, Georgia Mason, Robert T. Mason, Anne
McBride,MeeraKumarMusic,Ann-ElizabethNash,EmmaNicholas,KarenOver-
all, Jesús Rivas, Michael Salmon, Gordon Schuett, Kurt Schwenk, Sally Sherwen,
AndreaM.Simmons,MarisaSpain,LeviStorks,CliffSummers,SashaJ.Tetzlaff,
Tim Tristan, Richard Vogt, Adrian Walton, Paul Weldon, Geoff While, Martin
Whitehead, Alexandra Whittaker, Anna Wilkinson, and Alix Wilson. Scientific
books,especiallylargeandcomplexmulti-authorprojectssuchasthis,alsorequire
a dedicated publishing team, and with this in mind we thank in particular Annette
Klaus, Bibhuti Bhusan Sharma, Lars Koerner, and all others involved at Springer,
for their guidance and patience throughout. The editors are grateful to Dr Karen
Mancera for reading and commenting on the complete final text. The editors also
wish to express their gratitude to Michael Taber for his valuable assistance in the
preparationofthespeciesandgeneraindex.ThefirsteditionofHealthandWelfare
of Captive Reptiles was, and remains, a historic and definitive milestone in the
evolution of academic and applied scientific welfare, and we envisage that this
new edition will continue to advance its messages long into the future. To all who
arepartofthisjourney,thankyou!
v
Contents
1 Introduction. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
CliffordWarwick,PhillipC.Arena,andGordonM.Burghardt
2 PhysiologyandFunctionalAnatomy. . . . . . . . . . . . . . . . . . . . . . . . 7
HarveyB.Lillywhite
3 SensorySystems. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
JennaM.Crowe-RiddellandHarveyB.Lillywhite
4 BiologyofStress. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
EricJ.GangloffandNeilGreenberg
5 NormalBehaviour. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
JamesC.GillinghamandDavidL.Clark
6 SocialBehaviourasaChallengeforWelfare. . . . . . . . . . . . . . . . . . 189
J.SeanDoody
7 Brains,Behaviour,andCognition:MultipleMisconceptions. . . . . . 211
EnriqueFont,GordonM.Burghardt,andManuelLeal
8 PsychologicalandBehaviouralPrinciplesandProblems. . . . . . . . . 239
CliffordWarwick
9 EffectsofOntogeny,RearingConditions,andIndividual
DifferencesonBehaviour:Welfare,Conservation,andInvasive
SpeciesImplications. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287
GordonM.BurghardtandDonnaG.Layne-Colon
10 ControlledDeprivationandEnrichment. . . . . . . . . . . . . . . . . . . . . 323
RobertW.MendykandLaurenAugustine
11 EffectsofNoiseandLight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357
KarenF.ManceraandCliveJ.C.Phillips
vviiii
viii Contents
12 EthologicallyInformedDesignandDEEPEthologyinTheory
andPractice. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379
NeilGreenberg
13 SpatialandThermalFactors. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 417
PhillipC.ArenaandCliffordWarwick
14 NutritionalConsiderations. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 447
MichaelThomasMaslanka,FredricL.Frye,BarbaraAnnHenry,
andLaurenAugustine
15 NaturalisticVersusUnnaturalisticEnvironments. . . . . . . . . . . . . . 487
CliffordWarwickandCatrinaSteedman
16 EvidentialThresholdsforSpeciesSuitabilityinCaptivity. . . . . . . . 509
MikeJessop,AnthonyPilny,CliffordWarwick,
andMartinWhitehead
17 RecordKeepingasanAidtoCaptiveCare. . . . . . . . . . . . . . . . . . . 535
RobertW.MendykandJudithBlock
18 ArbitraryHusbandryPracticesandMisconceptions. . . . . . . . . . . . 561
RobertW.MendykandCliffordWarwick
19 MiscellaneousFactors. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 583
PhillipC.Arena,MeredithJ.Bashaw,RachelGrant,TiffaniHowell,
AlbertMartínez-Silvestre,andCliffordWarwick
Index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 619
AnimalIndex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 633
Editors and Contributors
About the Editors
CliffordWarwick isanindependenthumanmedicalscientistandreptilebiologist.
Heholdsseveraladvancedprofessionalqualificationsspecialisinginreptilescience,
as well as a PhD in reptile welfare biology. He has produced approximately
150peer-reviewedarticles,bookchapters,andbooks,mostlyconcerningthebiology
andwelfareofreptiles,withafocusonanthropogenicimpacts.
PhillipC.Arena isanawardededucatorandfunctionalmorphologistspecialising
inreptilebiology,health,andwelfare.Hehaspublishedmaterialontheanaesthesia,
anatomy, and physiology of reptiles and contributed to key investigations on the
impact of farming and recreational practices on the lives of reptiles, including
rattlesnakes,marineturtles,andcrocodiles.
Gordon M. Burghardt is Alumni Distinguished Professor of Psychology and
EcologyandEvolutionaryBiologyattheUniversityofTennessee.Hehaspublished
widelyonreptilebehaviour,behaviouraldevelopmentandevolution,captiveanimal
issues, animal play, and conceptual and historical aspects of animal ethology and
comparativepsychology.
Contributors
Phillip C. Arena College of Science, Health, Engineering and Education, Aca-
demicOperations,EnvironmentalandConservationSciences,MurdochUniversity,
Mandurah,WA,Australia
LaurenAugustine SaintLouisZoo,SaintLouis,MO,USA
SmithsonianInstituteNationalZoologicalPark,Washington,DC,USA
Meredith J. Bashaw Life Sciences and Philosophy Building, Franklin and Mar-
shallCollege,Lancaster,PA,USA
Judith Block Registrar Emerita, Zoo Registrars Association, Washington, DC,
USA
iixx
x EditorsandContributors
Gordon M.Burghardt Departments ofPsychology andEcology &Evolutionary
Biology,UniversityofTennessee,Knoxville,TN,USA
DavidL.Clark DepartmentofBiology,AlmaCollege,Alma,MI,USA
JennaM.Crowe-Riddell DepartmentofEcologyandEvolutionaryBiology,Uni-
versityofMichigan,AnnArbor,MI,USA
J.SeanDoody UniversityofSouthFlorida,St.Petersburg,FL,USA
EnriqueFont EthologyLab,CavanillesInstituteofBiodiversityandEvolutionary
Biology,UniversityofValencia,Valencia,Spain
Fredric L. Frye Fund for Clinical Research, La Primavera Organic Farm,
Cloverdale,CA,USA
Eric J. Gangloff Department of Zoology, Ohio Wesleyan University, Delaware,
OH,USA
James C. Gillingham Department of Biology, Central Michigan University, Mt.
Pleasant,MI,USA
Rachel Grant Department of Applied Science, London South Bank University,
London,UK
NeilGreenberg DepartmentofEcologyandEvolutionaryBiology,TheUniversity
ofTennessee,Knoxville,TN,USA
BarbaraAnnHenry CincinnatiZoo&BotanicalGarden,Cincinnati,OH,USA
Tiffani Howell School of Psychology and Public Health, La Trobe University,
Bendigo,VIC,Australia
MikeJessop VeterinaryExpertServices,Swansea,UK
Donna G. Layne-Colon Texas Biomedical Research Institute, San Antonio, TX,
USA
Manuel Leal Division of Biological Sciences, University of Missouri, Columbia,
MO,USA
Harvey B. Lillywhite Department of Biology, University of Florida, Gainesville,
FL,USA
Karen F. Mancera Facultad de Medicina Veterinaria y Zootecnia, Universidad
NacionalAutónomadeMéxico,CiudaddeMéxico,Mexico
Albert Martínez-Silvestre Catalonian Reptiles and Amphibians Rescue Centre,
Barcelona,Spain
Michael Thomas Maslanka Department of Nutrition Science, Smithsonian
National Zoological Park and Conservation Biology Institute, Washington, DC,
USA
EditorsandContributors xi
RobertW.Mendyk DepartmentofHerpetology,AudubonZoo,NewOrleans,LA,
USA
Department of Herpetology, Smithsonian National Zoological Park, Washington,
DC,USA
Clive J. C. Phillips Curtin University Sustainable Policy (CUSP) Institute,
Bentley,WA,Australia
AnthonyPilny ArizonaExoticAnimalHospital,Phoenix,AZ,USA
CatrinaSteedman EmergentDiseaseFoundation,London,UK
CliffordWarwick EmergentDiseaseFoundation,London,UK
MartinWhitehead ChippingNortonVeterinaryHospital,Oxon,UK
