Reintroduction biology of tuatara (Sphenodon

punctatus): identifying suitable founder

animals and conservation translocation sites

Scott William Jarvie

A thesis submitted for the degree of

Doctor of Philosophy

at the University of Otago,Dunedin, New Zealand

2016

Mel J. Rivera Rodríguez

Abstract

Abstract

The rate of biodiversity loss is not slowing despite global commitments. Conservationtranslocations, the intentional movement and release of organisms to restore popu-lations, are an emerging tool to help reduce species loss, but translocation successrates can be low. Recent reviews of and guidelines for conservation translocations em-phasise the use of strategic approaches to improve translocation outcomes. Tuatara(Sphenodon punctatus) are endemic reptiles from New Zealand and the last survivingmember of Rhynchocephalia. The Sphenodon genus was once widespread through-out New Zealand, but, following human arrival and the introduction of mammalianpredators to the mainland, tuatara now naturally inhabit only 32 offshore islands. Along-term recovery goal is to re-establish populations of tuatara on the mainland andoffshore islands. The aim of this thesis is to increase the likelihood that conservationtranslocations of tuatara will succeed. This was addressed through two approaches:1) I compared the post-release performance and survival of captive-reared versus wild-caught juvenile tuatara, and 2) I applied correlative- and mechanistic-models to predictclimatically suitable sites for tuatara.

Post-release performance has rarely been compared between captive-reared and wild-conspecific reptiles, and results have been mixed. In Chapters 2 and 3, I compared mor-phometric, ecophysiological and behavioural traits, resource selection and, ultimately,survival of juveniles in three groups (wild-caught from a warmer climate, captive-reared from a warmer climate and captive-reared from the local climate) released intoOrokonui Ecosanctuary in southern New Zealand. For most metrics, including sur-vival, post-release performance of captive-reared and wild-caught juveniles were similar,and captive-reared groups sometimes differed between each other as much as betweencaptive-reared and wild. These findings highlight the species-specific nature of translo-cation outcomes and the need for adaptive management in the field of reintroductionbiology.

Populations might fail to establish following translocation, even when the most suitablefounder animals are selected, unless they are released into suitable areas of habitat.A key aspect of planning is therefore the selection of suitable release sites that matchthe biotic and abiotic needs of the focal species under current and future climates. InChapter 4, I used three widely applied correlative species distribution models to predictclimatically suitable sites for future translocations. Despite discrepancies for future

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Abstract

climates, the three correlative species distribution models projected an increase in areaof climatically suitable habitat for tuatara. The thermal sensitivity of physiologicalrates is a key characteristic of organisms. In Chapter 5, I measured metabolic rate andrates of evaporative water loss in juvenile tuatara, over a range of temperatures. Thesemetabolic and water loss data provide a firm basis for estimating the thermodynamicniche for tuatara. In Chapter 6, I use a biophysical model fitted with the physiologicaldata to illustrate the use of a mechanistic model for free-roaming tuatara under currentand future climates. This mechanistic model is the first for a free-roaming animalspecies in New Zealand and provides key information for tuatara conservation.

Taken together, my thesis makes an important contribution to the conservation scienceof tuatara and reptiles generally. The findings from this research are applicable to awide range of species that are translocated for conservation benefit.

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Acknowledgements

Acknowledgements

This thesis would not have been possible without the help, advice and encouragementof many people. Foremost, I would like to thank my primary supervisor Alison Cree.Alison, I am grateful for the guidance, patience and the discussions that contributedsubstantially to this research. Thanks for the opportunity to learn from you. I alsothank my secondary supervisor Philip Seddon. Phil, I am grateful for your advice,patience and feedback. Thanks to my advisors, Michael Thompson, of the Universityof Sydney, and Michael Kearney, of the University of Melbourne. I would not havebeen able to complete this project without your help.

Many more people provided advice or support at some stage of my research. I amgrateful to Jane Elith, Richard Earl and Grant Humphries for advice on correlativespecies distribution models, to Barbara Joos, John Lighton, Thomas Forrester, KellyHare and Marion Preest for guidance on measuring metabolic rate, to Mandy Cald-well, Nicola Mitchell, Sophie Arnall and Anna Carter for discussions about mechanisticspecies distribution models, to Alistair Senior, Yu-shun Hsu, Luke Easton, MatthieuBruneaux, Shinichi Nakagawa and Tim Jowett for advice on statistical analyses, toWilliam Gamble for advice on GIS analyses, to Ken Miller for advice on formattingfigures, to Mariano Recio and Stephen Adolph for advice on resource selection anal-yses, and Trevor Worthy, Paul Schofield, Neville Hudson, Nicolas Rawlence and AlanTennyson for advice on New Zealand’s vertebrate fossil records. Thanks also to AlistairSenior who took notes of Lance Hill’s PhD thesis on water loss of tuatara at the RareBooks and Special Collections, University of Sydney, Australia, to Anne Besson whoprovided inferred body temperature of adult tuatara data to validate the ectothermmodel, Nick Sleeman from Otago Polytechnic who provided technical support with a3D scanner, Andrew Tait from NIWA/Taihoro Nukurangi who provided access to NewZealand’s Virtual Climate Station Network data, and Emma Burns and Ellen Simawho arranged access to preserved tuatara specimens at Otago Museum.

This project entailed a great deal of fieldwork, more than one person could achievesafely by themselves, and I was fortunate to have had the assistance of the followingpeople: Stephen Adolph, Jonathan Armitage, Dan Barrett, Leon Berard, Anne Besson,Lenke Blauss, Laura Bussolini, Rachel Buxton, Andreas Bruder, Fernanda Centeno,Matthew Clarkson, Andrew Clarkson, Alison Cree, Rebecca Cumming, Luke Easton,Bastian Egeter, Valerie Fay, Maddalena Fumagalli, William Gamble, Sophie Gibson,

vii

Acknowledgements

Kieran Godfrey, Emily Gray, Stefanie Grosser, Kelly Gough, Anna Harris, KathrynHand, Samantha Haultain, Mike Hitchcock, Benedikt Holtmann, Katherine Hope, Yo-Shun Hsu, Jay Iwasaki, Frances Jacomb, Fátima Jorge, Tsukushi Kamiya, Claire Kil-ner, Jenny Kitchin, Aaron Korpal, Devonia Kruimer, Katha Lange, Carlos EstebanLara, Sara Larcombe, Nigel McDonald, Tahu MacKenzie, Fanny Maure, Noela Mc-Gregor, Ricardo Mello, Stefan Meyer, Sarah Mockett, Jennifer Morley, Georgia Moore,Natasha Noble, Grant Oakes, Katie O’Dywer, Sophie Penniket, Charlotte Penniket,Romana Salis, Junichi Sugishita, Tiff Stephens, Verity Stone, Edward Ramirez, ScottReid, Alister Robinson, Santiago Rodríguez Jiménez, Mel J. Rivera Rodríguez, CailinRoe, Amandine Sabadel, Leida dos Santos, Marcus Simons, Elton Smith, AnthonyStumbo, Xaxier Tatin, Matthew Thomson, Merenia Wright and Tim Wright.

For approval to translocate tuatara to Orokonui Ecosanctuary, I am grateful to NgatiKoata, kaitiaki or guardians of Stephens Island (Takapourewa), and Kati HuirapaRunaka ki Puketeraki, mana whenua of Otago. Many people provided assistance withthe collection or release of tuatara. Thanks to Nicola Nelson, Susan Keall, Earl Boyd,Samara Davis, Michael Elkington, Te Hotu Elkington, Pene Gieger, Noela McGre-gor, Tokotahi Paul, and Te Rangi Walters for collecting tuatara from Stephens Island(Takapourewa), and Bruce Benseman, Susan Keall, Nicola Nelson, Valerie Fay andRhys Mills, amongst others, for collecting tuatara from Nga Manu Nature Reserve.Thanks to Valerie Fay, Elton Smith, Kelly Gough, Alister Robinson, Sophie Welvaert,Nicola Nelson, Susan Keall, Rhys Mills, Bruce Benseman, Wahanui Elkington, MichaelElkington, Noela McGregor, Sophie Penniket, Anne Besson, Stephen Adolph and Ali-son Cree for release assistance. I would like to acknowledge the support of the OtagoNatural History Trust (who manage Orokonui Ecosanctuary), particularly Neville Peatand Alyth Grant, and Orokonui Ecosanctuary staff and volunteers, especially ChrisBaillie, Valerie Fay, Alister Robinson and Elton Smith.

Thanks to the Department of Zoology staff. I enjoyed being part of a diverse de-partment and benefited from your support. I would like to acknowledge Ronda Keen,Esther Sibbald, Wendy Shanks and Vivienne McNaughton for administrative support,Erik Liepins and Jonathon Ung for computer support, Matthew Downes, Ken Millerand Nicky McHugh for technical support, Amy Adams, Kim Garrett and Nat Limfor fieldwork and laboratory support, and last, but not least, Murray McKenzie, forassistance with equipment for both field and laboratory studies. Murray, I am gratefulfor your help and support, particularly during the metabolic rate and field studies.

I have been fortunate to share an office or the corridors of the Department of Zoologywith people I look forward to seeing each day. In particular, I would like to acknowledge

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Acknowledgements

Jason Augspurger, Anne Besson, Andreas Bruder, Leida dos Santos, Luke Easton, Bas-tian Egeter, Maddalena Fumagalli, Emily Gray, Stefanie Grosser, Samantha Haultain,Benedikt Hoffman, Frances Jacomb, Fatima Jorge, Kathryn Hand, Tsukushi Kamiya,Jo Krawiek, Sam Macaulay, Jonathan McCallum, Rohan Menzies, Georgia Moore,Jennifer Morley, Stefan Meyer, Katie O’Dywer, Dominik Paschke, Sophie Penniket,Kalinka Rexer-Huber, Marine Richardson, Romana Salis, Anthony Stumbo, JunichiSugishita and Zachiary Tobias. To my friends outside the Department, thanks foryour support and encouragement. In particular, I would like to acknowledge MahdisArzamandi, Tom Csima, Keiran Godfrey, Emily Gray, Paula Hardie, Anna Harris,Siobhan Haynes, Brenah Hearne, Matthew Jenkins, Josh Kent, Aaron Korpal, De-vonia Kruimer, Tahu MacKenzie, Jo Monks, Harvey Penfold, Sara Rahmani, AndrewReynolds, Santiago Rodríguez Jiménez, Cailin Roe, Hayden Ryder, Amandine Sabadel,Danielle Salmon, Rose Stamm, Aviva Stein, Andrew Tuaine, Penny Wood and TimWright.

Many people or organisations loaned or purchased equipment for this research. Thanksto Mike Denham and Ray McLennan, from the School of Surveying, University ofOtago, for GNNS or GPS units, Chris Jones, from Landcare Research, for the bur-rowscope, David Howarth and Nigel McDonald, from the Department of Geography,Univeristy of Otago, for meteorological equipment, Shinichi Nakagawa and Carlos Lara,from the Department of Zoology, for the loan of the spectrometer, and John Lighton,Thomas Forrester and Barbara Joos, from Sable Systems, for metabolic rate equip-ment. Thanks also to Anne Ryan who made the respirometry chambers and AlisterRobinson who bought the first passive-transponder reader unit.

For constructive comments that improved chapters, I thank Anne Besson, Sophie Pen-niket, David Towns and Anna Carter. I am grateful to the seven anonymous reviewersand the Editorial Teams at Animal Conservation, Herpertological Review, the Journalof Zoology and the New Zealand Journal of Ecology who provided valuable feedback onearlier versions of the published chapters or appendices. Thanks also to Leon Berardand Mel J. Rivera Rodríguez who provided images that appear throughout the thesis.

Funding for this thesis research was provided by Department of Zoology at the Uni-versity of Otago, the Diane Campbell-Hunt Memorial Award, the Australian Societyof Herpetologists Award, the Company of Biologists, MJ & BJ Ecology Fund, and theSociety for Research on Amphibians and Reptiles in New Zealand. Thanks to the Uni-versity of Otago Doctoral Scholarship for financial support. The University of OtagoDivision of Sciences and Department of Zoology provided funds to attend national andinternational conferences. My research would not have been possible without the sup-

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Acknowledgements

port of tangata whenua, particularly Ngati Koata, Kati Huirapa Runaka ki Puketeraki,and the Te Runanga o Ngai Tahu Research Consultation Committee at the Universityof Otago. For providing the necessary permits and approval, I am grateful to the De-partment of Conservation/Te Papa Atawhai (permit NM-32559-FAU and 39465-RES),the University of Otago Animal Ethics Committee (AEC permit 102-11 and 62-14),and the Otago Natural History Trust who manage Orokonui Ecosanctuary.

Last but not least, I would like to thank my family for all of your support during myresearch. Without years of encouragement, none of this would have been possible.

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Table of Contents

Table of Contents

Abstract v

Acknowledgements vii

Table of Contents xi

List of Tables xv

List of Figures xvii

List of Abbreviations xix

1 General Introduction 11.1 Conservation translocations . . . . . . . . . . . . . . . . . . . . . . . . 21.2 Identifying suitable founder animals for conservation translocation . . . 3

1.2.1 Captive- versus wild-caught individuals . . . . . . . . . . . . . 31.3 Identifying conservation translocation sites with suitable climates . . . 4

1.3.1 Approaches used to predict conservation translocation sites . . 51.4 Tuatara . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

1.4.1 Conservation translocations of tuatara . . . . . . . . . . . . . . 121.5 Objectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

1.5.1 Thesis outline and style . . . . . . . . . . . . . . . . . . . . . . 131.5.2 Contributions to research . . . . . . . . . . . . . . . . . . . . . 14

2 Captive-rearing affects growth but not survival in translocated ju-venile tuatara 172.1 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182.2 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192.3 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

2.3.1 Head-start facilities and release site . . . . . . . . . . . . . . . . 202.3.2 Release . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212.3.3 Summer growth rates, body condition, emergence, inferred body

temperatures and tick load . . . . . . . . . . . . . . . . . . . . . 232.3.4 Post-release dispersal and home range over summer . . . . . . . 252.3.5 Survival . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 252.3.6 Statistical analyses . . . . . . . . . . . . . . . . . . . . . . . . . 25

2.4 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 272.4.1 Summer growth rates, body condition, emergence, inferred body

temperatures and tick load . . . . . . . . . . . . . . . . . . . . . 27

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2.4.2 Post-release dispersal and home range over summer . . . . . . . 312.4.3 Survival . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

2.5 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 342.5.1 Summer growth rates, body condition, emergence, inferred body

temperatures and ticks . . . . . . . . . . . . . . . . . . . . . . . 342.5.2 Post-release dispersal and home range over summer . . . . . . . 362.5.3 Survival . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 372.5.4 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

3 Resource selection by tuatara following translocation: a comparisonof wild-caught and captive-reared juveniles 393.1 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403.2 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 413.3 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

3.3.1 Reintroduced juvenile tuatara and release site . . . . . . . . . . 423.3.2 Post-release monitoring . . . . . . . . . . . . . . . . . . . . . . . 423.3.3 Resource selection analyses . . . . . . . . . . . . . . . . . . . . . 43

3.4 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 463.4.1 Landscape scale . . . . . . . . . . . . . . . . . . . . . . . . . . . 463.4.2 Local scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 463.4.3 Retreat-site selection . . . . . . . . . . . . . . . . . . . . . . . . 49

3.5 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 493.5.1 Landscape scale . . . . . . . . . . . . . . . . . . . . . . . . . . 493.5.2 Local scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 503.5.3 Retreat-site selection . . . . . . . . . . . . . . . . . . . . . . . . 513.5.4 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

4 Identifying climatically suitable habitat for tuatara: a correlativeapproach to species distribution modelling 534.1 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 544.2 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 554.3 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . 57

4.3.1 Distribution data . . . . . . . . . . . . . . . . . . . . . . . . . . 574.3.2 Climate data . . . . . . . . . . . . . . . . . . . . . . . . . . . . 614.3.3 Correlative species distribution models . . . . . . . . . . . . . . 62

4.4 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 644.4.1 Current climate . . . . . . . . . . . . . . . . . . . . . . . . . . . 644.4.2 Climate change scenarios . . . . . . . . . . . . . . . . . . . . . . 70

4.5 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 814.5.1 Predictions of climatically suitable habitat for tuatara under cur-

rent climate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 814.5.2 Projections of climatically suitable habitat for tuatara under cli-

mate change . . . . . . . . . . . . . . . . . . . . . . . . . . . . 824.5.3 Comparison of correlative and (previous) mechanistic species dis-

tribution models for remnant populations of tuatara . . . . . . 844.5.4 Reintroductions and other conservation translocations of tuatara

under current and future climates . . . . . . . . . . . . . . . . 884.5.5 Variation in the projections of correlative species distribution

models . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90

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4.5.6 Future directions to identify climatically suitable sites for thetranslocation of tuatara . . . . . . . . . . . . . . . . . . . . . . 91

4.5.7 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91

5 Thermal effects on metabolic rate and water loss in juvenile tuatara 935.1 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 945.2 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 955.3 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . 96

5.3.1 Respirometry equipment set-up . . . . . . . . . . . . . . . . . . 975.3.2 Diel cycle in metabolic rate . . . . . . . . . . . . . . . . . . . . 985.3.3 Thermal profile for metabolic rate . . . . . . . . . . . . . . . . . 985.3.4 Effect of temperature on water loss . . . . . . . . . . . . . . . . 995.3.5 Calculations and statistical analysis . . . . . . . . . . . . . . . . 100

5.4 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1025.4.1 Diel cycle in metabolic rate . . . . . . . . . . . . . . . . . . . . 1025.4.2 Thermal profile for metabolic rate . . . . . . . . . . . . . . . . . 1055.4.3 Effect of temperature on rate of total evaporative water loss . . 107

5.5 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1105.5.1 Diel cycle in metabolic rate . . . . . . . . . . . . . . . . . . . . 1115.5.2 Thermal profile for metabolic rate . . . . . . . . . . . . . . . . 1125.5.3 Respiratory exchange ratio . . . . . . . . . . . . . . . . . . . . 1155.5.4 Effect of temperature on rate of total evaporative water loss . . 1165.5.5 Incubation conditions . . . . . . . . . . . . . . . . . . . . . . . 1185.5.6 Conclusion and implications for conservation translocations of

tuatara . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118

6 Identifying climatically suitable habitat for tuatara: a mechanisticapproach to a species distribution model 1216.1 Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1226.2 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1236.3 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . 127

6.3.1 Study species and field site . . . . . . . . . . . . . . . . . . . . . 1276.3.2 Monitoring inferred body temperature, microclimate and opera-

tive temperature . . . . . . . . . . . . . . . . . . . . . . . . . . 1276.3.3 Biophysical model . . . . . . . . . . . . . . . . . . . . . . . . . 1286.3.4 Data analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133

6.4 Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1346.4.1 Field body temperatures, copper models and weather data . . . 1346.4.2 Validation of microclimate model . . . . . . . . . . . . . . . . . 1356.4.3 Validation of the ectotherm model . . . . . . . . . . . . . . . . 1356.4.4 Projections from biophysical model under current and future cli-

mates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1356.5 Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141

6.5.1 Predictions of the microclimate and ectotherm models . . . . . 1416.5.2 Projections of the ectotherm model under current and future

climate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1426.5.3 Reintroductions and other conservation translocations of tuatara 1446.5.4 Future steps to help identify climatically suitable sites for the

translocation of tuatara . . . . . . . . . . . . . . . . . . . . . . 145

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6.5.5 Caveats of mechanistic species distribution models for tuatara . 1476.5.6 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148

7 General Discussion 1497.1 Review of the main findings . . . . . . . . . . . . . . . . . . . . . . . . 150

7.1.1 Identifying suitable founder animals for translocation: a researchsynthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150

7.1.2 Identifying conservation translocation sites with suitable climates:a research synthesis . . . . . . . . . . . . . . . . . . . . . . . . 151

7.2 Recommendations for future research . . . . . . . . . . . . . . . . . . . 1527.2.1 Identifying suitable founder animals for translocation: future re-

search opportunities . . . . . . . . . . . . . . . . . . . . . . . . 1527.2.2 Identifying conservation translocation sites with suitable climates:

future research opportunities . . . . . . . . . . . . . . . . . . . 1537.2.3 Metabolic rate and rates of total evaporative water loss in tu-

atara: future research opportunities . . . . . . . . . . . . . . . 1567.3 Management recommendations . . . . . . . . . . . . . . . . . . . . . . 157

7.3.1 Recommendation 1: wild-caught and head-started individualsare suitable as founder animals. . . . . . . . . . . . . . . . . . . 157

7.3.2 Recommendation 2: monitoring of remnant and translocatedpopulations is needed. . . . . . . . . . . . . . . . . . . . . . . . 157

7.3.3 Recommendation 3: climatically suitable sites should be identi-fied with species distribution models. . . . . . . . . . . . . . . 158

7.4 Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158

References 159

Appendices 199

A Assessing the thermal suitability of translocation release sites foregg-laying reptiles with temperature-dependent sex determination:a case study with tuatara (Sphenodon punctatus) 199

B Attaching radio transmitters does not affect mass, growth or dis-persal of translocated juvenile tuatara (Sphenodon punctatus) 219

C A non-invasive method to measure surface area in reptiles: a casestudy using tuatara 231

D Supporting materials for Chapter 2 243

E Supporting materials for Chapter 3 251

F Supporting materials for Chapter 4 257

G Supporting materials for Chapter 5 273

H Supporting materials for Chapter 6 279

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