This electronic thesis or dissertation has been downloaded from Explore Bristol Research, http://research-information.bristol.ac.uk Author: Mellor, Emma L Title: Does natural foraging niche influence captive animal health and welfare? General rights Access to the thesis is subject to the Creative Commons Attribution - NonCommercial-No Derivatives 4.0 International Public License. A copy of this may be found at https://creativecommons.org/licenses/by-nc-nd/4.0/legalcode This license sets out your rights and the restrictions that apply to your access to the thesis so it is important you read this before proceeding. Take down policy Some pages of this thesis may have been removed for copyright restrictions prior to having it been deposited in Explore Bristol Research. 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Emma Louise Mellor A dissertation submitted to the University of Bristol in accordance with the requirements for award of the degree of Doctor of Philosophy in the Faculty of Health Sciences Bristol Veterinary School September 2020 64,325 words Abstract Understanding the basis for differences in how species typically respond to captivity is fundamental for addressing welfare-relevant management problems created by captivity. This can be achieved by formal cross-species comparisons and testing hypotheses for how species-typical ecology and biology might affect their representatives’ typical captive response. My PhD examined potential foraging-niche related risk factors, i.e. the way a species uses its habitat and resources in relation to gaining food, for two different types of welfare-relevant management problem across three taxonomic groups. Using phylogenetic comparative methods to control for species non-independence due to shared ancestry, I examined relationships between hunting behaviour and route-tracing severity across zoo- housed Carnivora; and determined whether wild food-search or -handling behaviours predict the prevalence of feather-damaging behaviour and other stereotypic behaviour in pet Psittaciformes. Across zoo-housed Lemuriformes, foraging niche was just one of three aspects of species-typical biology assessed for potential predictive effects on a second type of management problem: susceptibility to weight gain. Taking an epidemiological approach, I also explored several individual-level potential demographic and environmental risk factors for susceptibility to weight gain across a subset of four Lemuriform species. Naturally relying on extensive food-handling was revealed as risk factor for feather-damaging behaviour across Psittaciformes, and adaptations to unpredictable wild food resources might be a risk factor for weight gain in Lemuriformes. Regarding the latter, epidemiological analyses revealed several demographic risk factors and one environmental one in male lemurs (being housed with only fixed climbing structures). Based on my results, I make practical recommendations to help address these specific management problems; describe fundamental, collection-management benefits this approach yields, by identifying types of species less-suited to captivity; and detail areas for future research, with the overall aim of improving wellbeing of thousands of animals across many species. i Acknowledgements Where to start? This might have my name on it, but this sort of thing is never done in isolation and represents input from many people to whom I am incredibly grateful. First, my thanks go to my fantastic supervisors Mike Mendl, Innes Cuthill and Georgia Mason for their endless support, questioning, patience, and invaluable advice. During a visit to another university, a researcher there summed you up perfectly: “hell’uva supervisory team” (best said in broadest South Western). I feel privileged to have you as my supervisors, and you’re fantastic role models. Thanks also for your continued support, keeping me in work doing what I adore. I also wish to thank Sarah Collins, who I am pretty sure has little idea of how her encouragement during my undergraduate years influenced me and the path I’ve taken. And I must also thank Ben Brilot, as without his idea of making the abstract I supplied for my PhD interview a mini-poster rather than plain ol’ text, I very much doubt the outcome would have been even half as successful. I also wish to thank fellow lab mates at Bristol Vet School, especially Vikki Neville, Marco Ramirez and Melissa Smith, for their company during this bonkers shared experience. From the Mason Lab past and present, my thanks particularly to Carole Fureix and Andrea Polanco, and to Miranda Bandeli for sharing with me the beast that is the carnivore database. I am grateful also to Heather McDonald Kinkaid for allowing me to use her data when it all went horribly wrong, and for answering my endless questions about her own work. And thanks to the wonderful human being that is María Díez-León, both for the mutual geeky interests (go P cats!) and her friendship. My thanks also to Yvonne van Zeeland for her valuable advice on the parrot side of things. For their advice on and support for my lemur project my thanks to: Christoph Schwitzer; Christie Eddie and the rest of the AZA Prosimian TAG; Amy Plowman and the rest of the BIAZA Research Committee; and Sarah Zehr. For taking time to read and feedback on various sections of this thesis: Sally Sherwen, Mark Puttick, Jamie Ahloy Dallaire, Sean Nee, Hafiz ii Maherali, Jason Watters, Olivia Mellor and John. And for their assistance with my lemur survey, without which I would probably still be coding data, I am grateful to Shelley Jackman, Georgia Dacombe and Rachel Brown. For their expertise in skills I am woefully lacking I need to thank the following: Elise Shepard, Paola Lunetta, Matteo Montagna, Alessia Maina, Stefan Pinkert, Claudia Serien, Suzanne Held, Christina Rohe, Laura Rodriguez Arco, and Rogelio Rodriguez Lopez for assisting with the Parrot Survey translations. I am grateful to Jack Hughes for his climate know-how, and to Miquel Vall-Llosera and Gustavo Paterno for generously providing some R code – along with an idiot’s guide of to how to use it – and Vikki for her assistance when I still came unstuck. Thanks also to Laszlo Talas for his help with mixed models. I am very grateful to the University of Bristol for providing me with my PhD studentship and research training fund. For further financial support and helping me attend various conferences over the years, I also need to thank ASAB, the Arthur Hosier Bequest, and the University of Bristol’s Alumni Foundation. For providing data, thanks to Agnieszka Sergiel and Ulrica Alhot for their captive carnivore data. Also thanks to all my wonderful ‘lemur’ zoos for data and photographs of their animals: Adelaide Zoo, Al Bustan Zoological Centre, United Arab Emirates, Alameda Wildlife Park, Alligator Farm Zoological Park, Aquazoo Friesland, Audubon Zoo, Barcelona Zoo, Birmingham Zoo, Cameron Park, Cango Wildlife Ranch, Central Florida Zoo, Cheyenne Mountain Zoo, Cleveland Metroparks Zoo, Duke Lemur Center, Furuvik Zoo, Givskud Zoo – Zootopia, Greater Vancouver Zoo, Hannover Zoo, Hemsley Conservation Centre, Honolulu Zoo, Kaliningrad Zoo, Kosice Zoo, Lake Superior Zoological Society, Lakeland Wildlife Oasis, Lehigh Valley Zoo, Lemur Conservation Foundation, Little Rock Zoo, Minsk Zoo, Nashville Zoo, National Zoological Park, Nikolaev Zoo, Omaha’s Henry Doorly Zoo and Aquarium, Paignton Zoo, Parc Animalier D'auvergne, Parc Merveilleux, Parco Faunistico Valcobra, Parken Zoo, Peak Wildlife Park, Perm Zoo, Point Defiance Zoo & Aquarium, Randers iii Regnskov, Rolling Hills Zoo, Rotterdam Zoo, Santa Fe College Teaching Zoo, Śląski Ogród Zoologiczny in Chorzów, The Cincinnati Zoo and Botanical Garden, Twycross Zoo and the East Midlands Zoological Society, Wellington Zoo Trust, Zoo Bratislava, Zoo de Lyon, Zoo Decin, Zoo Landau, Zoo Vienna (Tiergarten Schönbrunn), Zoo Wuppertal, Zoologischer Garten Schwerin, and all others that contributed. And lastly many, many thanks to my family for putting up with increasing forgetfulness (several birthdays, one wedding anniversary [mine]), grumpiness, general scattiness, and my working during weekends and occasional family events over these past few years. To my parents and in-laws, special thanks for all the childcare over this past year. John – you are the absolute best, and ‘thanks’ seems too little for all your love, support, cooking, and ability to present coffee, wine, chocolate and/or other snacks at just the right time. You’re amazing and I love you. Eve, thanks for forcing me to become more efficient. You may have made this even harder, but I wouldn’t change a thing. Plus, I love that you can correctly identify lemurs and parrots aged two. Carnivores are a work in progress (in fairness, they have pretty diverse morphologies. Maybe by the time she’s three?). And of course, to Hamish – aka Hambo-fandango – who almost, but not quite, saw the whole thing from start to finish, and who I miss terribly. iv Author’s declaration I declare that the work in this dissertation was carried out in accordance with the requirements of the University's Regulations and Code of Practice for Research Degree Programmes and that it has not been submitted for any other academic award. Except where indicated by specific reference in the text, the work is the candidate's own work. Work done in collaboration with, or with the assistance of, others, is indicated as such.