Human-animal interaction in the modern zoo: Live animal encounter programs and associated effects on animal welfare. Lydia Acaralp-Rehnberg ORCID: 0000-0002-0955-2077 Doctor of Philosophy (PhD) July 2019 The Animal Welfare Science Centre, Faculty of Veterinary and Agricultural Sciences, The University of Melbourne This thesis is being submitted in total fulfilment of the degree. 1 Abstract Live animal encounter programs are an increasingly popular occurrence in the modern zoo. The welfare implications of animals participating in these programs has not been studied extensively to date. The aim of the current thesis was, therefore, to explore animal welfare effects associated with encounter programs in selected zoo-housed species, using a combined approach of survey data and empirical investigations. The survey assessment, which involved over 500 accredited zoos and aquariums in Australia and overseas, revealed that most surveyed institutions (85 %) engaged in an encounter program with one or more species, most of which were mammals. Behaviours indicative of a typical fight-or-flight response were the most commonly reported welfare concerns in relation to encounters. Behaviour indicative of positive welfare, as well as voluntary participation and interaction with visitors, were commonly mentioned in regard to positive welfare during encounters. Positive welfare experiences outweighed the number of reported concerns in birds and mammals, but the opposite was true for reptiles. The empirical studies involved zoo-housed servals, giraffes and shingleback lizards, and sought to identify a potential cause-effect relationship between behavioural and physiological welfare indices and short-term variations in encounter frequency. A similar methodology involving a repeated treatment design where the frequency of encounters was manipulated to reflect the regular frequency of encounters, a temporary withdrawal, and a temporary intensification of regime, was adopted across the three studies. Behavioural changes indicative of a positive welfare effect when participating in encounters were observed in the servals and giraffes. Servals exhibited a significant reduction in stereotypic pacing on weeks when participating in interactive presentations, or presentations and behind-the-scenes encounters combined. The giraffes engaged in amicable social interactions significantly more often when participating in visitor feeding encounters, at either regular or intensified frequency. By contrast, a potentially aversive effect of encounters was observed in the shinglebacks, who significantly increased their use of concealed locations within the enclosure when handled at either the regular or intensified frequency. Approach behaviour during encounters differed significantly between individual giraffes, and significant differences in coiling behaviour while handled was observed in the lizards. The findings of the empirical studies were in agreement with the survey data, which identified taxonomic as well as individual variation as important influences on the welfare of animals participating in encounters. The nature of the encounter the animals participate in was identified as another key factor, in which 2 encounters that maximise choice and control, and where animals are rewarded for participation, were likely to contribute to a more positive welfare experience. Declaration I, Lydia Acaralp-Rehnberg, hereby declare: i) The thesis comprises only my original work towards the PhD. ii) Due acknowledgement has been made in the text to all other materials used. iii) The thesis is fewer than the maximum word limit in length, exclusive of tables, maps, bibliographies and appendices, as approved by the Research Higher Degrees Committee. Date: 07.07.2019 3 Acknowledgements I would like to extend my warmest thanks to my supervisory panel, with principal supervisor Dr. Ian Bland, co-supervisors Prof. Grahame Coleman, Dr. Michael Magrath and Dr. Vicky Melfi, for providing much valuable academic advice, support and guidance throughout the course of my candidature. I would also like to acknowledge the contribution of my advisory committee, including Dr. Lauren Hemsworth, Dr. Jean-Loup Rault and Dr. Megan Verdon, and Prof. Paul Hemsworth and Dr. Sally Sherwen, for valuable discussions and input into the current project. Many thanks to Dr. Kerry Fanson for her much valuable contribution to the physiological analysis, and to Dr. Kym Butler for valuable input and advice in regard to experimental design and statistical analysis. Thanks also to the other students, research fellows and research assistants at The Animal Welfare Science Centre for their support and assistance with the current project. My warmest thanks to the management and keeper teams at Zoos Victoria for granting us permission to carry out the current research at their facilities, and for their vital support and assistance throughout the course of my research. Finally, I would like to acknowledge the love and support of my wonderful family and friends for believing in the importance of my research. Special thanks to my beloved husband, Engin Acaralp, my two beautiful sons, Ricky and Aslan, and my loyal feline and canine companions for sharing every moment of my journey, challenges and joyful moments alike. The PhD candidature was funded by MIFRS (Melbourne International Fee Remission Scholarship) and MIRS (Melbourne International Research Scholarship), and the current research was funded by The Australian Research Council (ARC Linkage Grant, project id. LP140100373). 4 Table of contents Abstract p. 2 Author declaration p. 3 Acknowledgements p. 4 General introduction p. 13-15 Chapter 1. Literature review p. 16-51 1.1 Defining and assessing animal welfare p. 16 1.2 Human-animal interaction and associated p. 25 effects on animal welfare 1.3 Effects of unfamiliar humans on the welfare p. 29 of zoo-housed animals 1.4 Interactive experiences and their effects on p. 46 visitor learning, attitude and behaviour 1.5 Conclusion p. 50 Chapter 2. Methodology p. 52-61 2.1 Introduction p. 52 2.2 Research approach p. 54 2.3 Experimental design p. 56 2.4 Behavioural and physiological measures p. 57 2.5 Statistical analysis p. 60 Chapter 3. A multi-national survey assessment of live p. 61 - 108 animal encounter programs in accredited zoos and aquariums Abstract p. 61 3.1 Introduction p. 62 3.2 Methodology p. 64 3.2.1 Survey participants p. 64 3.2.2 Website review p. 65 3.2.3 Introductory survey p. 67 3.2.4 Follow-up survey p. 68 3.2.5 Data analysis p. 69 3.3 Results p. 70 3.3.1 Survey demographics p. 70 3.3.2 Current encounter programs p. 72 3.3.3 Welfare concerns and positive p. 79 5 welfare in relation to encounter programs 3.3.4 Management strategies p. 85 3.3.5 Attitudes and beliefs in relation p. 88 to encounter programs 3.4 Discussion p. 93 3.4.1 General trends in encounter p. 93 programs 3.4.2 Welfare concerns and positive p. 96 welfare in relation to encounter programs 3.4.3 Attitudes and beliefs in relation p. 101 to encounter programs 3.4.4 Study limitations p. 105 Conclusions p. 107 Chapter 4. The effect of behind-the-scenes encounters and p.109-142 interactive presentations on the welfare of captive servals (L. serval) at Werribee Open Range Zoo Abstract p. 109 4.1 Introduction p. 110 4.2 Methodology p. 113 4.2.1 Study animals p. 113 4.2.2. Visitor interaction program p. 115 4.2.3 Experimental design p. 117 4.2.4 Behavioural observations p. 118 4.2.5 Physiological measures p. 121 4.2.6 Statistical analysis p. 122 4.3 Results p. 123 4.3.1 Activity budget p. 123 4.3.2 Treatment effects p. 123 4.3.3 Effect of individual, session and p. 125 housing 4.3.4 Behaviour during serval p. 132 presentations 4.3.5 Physiological measures p. 132 4.4 Discussion p. 133 4.4.1 Behaviour outside of visitor p. 133 interaction 4.4.2 Behaviour during serval p. 137 presentations 4.4.3 Physiological measures p. 138 4.4.4 Other influences p. 138 4.4.5 Study limitations p. 140 6 Conclusions p. 141 Chapter 5. The effect of guest feeding encounters on the p. 143-171 welfare of captive giraffes (G. camelopardalis) at Melbourne Zoo Abstract p. 143 5.1 Introduction p. 144 5.2 Methodology p. 146 5.2.1 Study animals p. 146 5.2.2. Visitor interaction program p. 148 5.2.3 Experimental design p. 149 5.2.4 Behavioural observations p. 150 5.2.5 Physiological measures p. 153 5.2.6 Statistical analysis p. 154 5.3 Results p. 155 5.3.1 Activity budget p. 155 5.3.2 Treatment effects p. 155 5.3.3 Effect of individual and session p. 156 5.3.4 Behaviour during feeding p. 161 encounters 5.3.5 Physiological measures p. 163 5.4 Discussion p. 163 5.4.1 Behaviour outside of visitor p. 163 interaction 5.4.2 Behaviour during feeding p. 166 encounters 5.4.3 Physiological measures p. 168 5.4.4 Other influences p. 168 5.4.5 Study limitations p. 169 Conclusions p. 171 Chapter 6. Welfare in relation to interactive handling p. 172-198 sessions in the shingleback lizard (T. rugosa) Abstract p. 172 6.1 Introduction p. 173 6.2 Methodology p. 175 6.2.1 Study animals p. 175 6.2.2. Visitor interaction program p. 178 6.2.3 Experimental design p. 179 6.2.4 Behavioural observations p. 180 6.2.5 Statistical analysis p. 183 7 6.3 Results p. 184 6.3.1 Activity budget p. 184 6.3.2 Treatment effects p. 184 6.3.3 Effect of individual and session p. 185 6.3.4 Behaviour during handling p. 190 sessions 6.4 Discussion p. 191 6.4.1 Behaviour outside of handling p. 191 sessions 6.4.2 Behaviour during handling p. 194 sessions 6.4.3 Physiological measures p.