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Liopholis Whitii: Insights Into the Evolutionary Origins of Sociality

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Liopholis Whitii: Insights Into the Evolutionary Origins of Sociality Behaviour, Ecology and Social Organisation in Liopholis whitii: Insights into the Evolutionary Origins of Sociality Benjamin Halliwell, Bsc Hons School of Biological Sciences, University of Tasmania A thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy November 2016 Declarations Declaration of originality This thesis contains no material which has been accepted for a degree or diploma by the University of Tasmania or any other institution, and to the best of my knowledge and belief, this thesis contains no material previously published or written by another person, except where due acknowledgement is made in the text of this thesis, nor does the thesis contain any material that infringes copyright. Ben Halliwell Date: 9/11/2016 Authority of access My thesis may be made available for loan and limited copying and communication in accordance with the Copyright Act, 1968. Statement regarding published work contained in the thesis The publishers of the papers comprising Chapter 2, 3, 4 and 5 (and appendices V and VI) hold the copyright for that content, and access to the material should be sought from the respective journals. The remaining non-published content of the thesis may be made available for loan and limited copying and communication in accordance with the Copyright Act 1968. Statement of ethical conduct The research associated with this thesis abides the Australian Code of Practice for the Care and Use of Animals for Scientific Purposes, 7th edition, 2004 and the University of Tasmanian Animal Ethics Guidelines. Ben Halliwell Date: 9/11/2016 ii Statement of Co-Authorship The following people and institutions contributed to the publication of research undertaken as part of this thesis: Ben Halliwell: School of Biological Sciences, University of Tasmania Geoffrey While: School of Biological Sciences, University of Tasmania Erik Wapstra: School of Biological Sciences, University of Tasmania Tobias Uller: Department of Zoology, University of Oxford; Department of Biology, University of Lund Barbara Holland: School of Physical Sciences, University of Tasmania Dave Chapple: School of Biological Sciences, Monash University Mike Gardner: School of Biological Sciences, Flinders University Thomas Botterill-James: School of Biological Sciences, University of Tasmania Emily Cooper-Scott: School of Biological Sciences, University of Tasmania Publications produced as part of this thesis: Halliwell B, Uller T, Chapple D, Gardner M, Wapstra E and While GM (2017) Habitat saturation promotes delayed dispersal in a social reptile. Behavioural Ecology: arw181 Ben Halliwell (candidate and primary author) conceived of the study, carried out associated field and laboratory work and was responsible for manuscript presentation. David Chapple (author 3) and Michael Gardner (author 4) provided academic feedback on the study. Tobias Uller (author 2), Geoffrey M While (author 6) and Erik Wapstra (author 5) contributed to the formalization, development, and refinement of the study and provided feedback manuscript presentation. Halliwell B, Uller T, Wapstra E and While GM (2016) Resource distribution mediates social and mating behaviour in a family living lizard. Behavioural Ecology: arw134. iii Ben Halliwell (candidate and primary author) conceived of the study, carried out associated field and laboratory work and was responsible for manuscript presentation. Tobias Uller (author 2), Geoffrey M While (author 4) and Erik Wapstra (author 3) contributed to the formalization, development, and refinement of the study and provided feedback during manuscript presentation. Botterill-James T, Halliwell B, Cooper-Scott E, Uller T, Wapstra E, While GM (2016) Habitat structure influences parent-offspring association in a social lizard. Frontiers in Ecology and Evolution 4: 96. Thomas Botterill-James (primary author) and Ben Halliwell (candidate and author 2) conceived of the study and along with Emily Cooper-Scott (author 3) carried out associated field and laboratory work. Thomas Botterill-James was responsible for manuscript presentation. Tobias Uller (author 2), Geoffrey M While (author 6) and Erik Wapstra (author 5) contributed to the formalization, development, and refinement of the study and provided feedback during manuscript presentation. While GM, Halliwell B and Uller T (2014) Reproductive biology and phylogeny of lizards and tuatara. Eds. Rheubert JL, Siegel DS, Trauth SE. CRC Press. 590-619. Geoffrey M While (primary author) was invited to write this review by the editors. All authors contributed to the collection and synthesis of information during the literature review, as well as preparation of the manuscript. Halliwell B, Uller T, Holland B and While GM. (submitted) Correlated evolution of viviparity and social grouping in squamate reptiles. Current Biology. Ben Halliwell (candidate and primary author) conceived of the study, conducted associated literature reviews, and was responsible for manuscript presentation. Barbara Holland (author 3) provided consultation on statistical methods. Tobias Uller (author 2) and Geoffrey M While (author 4) contributed to the formalization, development, and refinement of the study and provided feedback during manuscript presentation. iv Additional publications produced during my candidature (Appendices V and VI): While GM, Williamson J, Prescott G, Horváthová T, Fresnillo B, Beeton NJ, Halliwell B, Michaelides S, Uller T (2015) Adaptive responses to cool climate promotes persistence of a non-native lizard. Proceedings of the Royal Society of London B: Biological Sciences 282: 20142638. Bordogna G, Cunningham G, Fitzpatrick LJ, Halliwell B, MacGregor HEA, Munch KL, Wapstra E and While GM (2016) An experimental test of relatedness-based mate discrimination in a social lizard. Behavioral Ecology and Sociobiology, doi: 10.1007/s00265-016-2217-9. We the undersigned agree with the above stated proportion of work undertaken for each of the above published (or submitted) peer-reviewed manuscripts contributing to this thesis: Erik Wapstra Geoffrey While Anthony Koutoulis Candidate’s supervisor Candidate’s supervisor Head of School School of Biological School of Biological School of Biological Sciences Sciences Sciences University of Tasmania University of Tasmania University of Tasmania Date: 9/11/2016 Date: 9/11/2016 Date: 9/11/2016 v Acknowledgements I am grateful to many people for making this journey such an enjoyable one. In particular, I acknowledge and thank the following: All members of the BEER Group! Special thanks go to my partners in work and, so often, crime: Kirke Munch, Tom Botterill-James, George Cunningham, Hannah MacGregor, Rachel Lewandowsky, and Luisa Fitzpatrick. I am truly grateful to have had the privilege of your company and the opportunity to make so many good memories in it. I also thank the many Postgrads and Undergrads that I have met and interacted with during my candidature for making this experience all the richer! My supervisory team: Erik Wapstra for your ongoing guidance and support. Thank you for welcoming me into the research group and providing such an academic yet convivial atmosphere for us to work; it has been a great pleasure! Tobias Uller for what became an unofficial supervisory role. I am very grateful to have had the opportunity to work in your lab and feel that your feedback and perspective made me strive for a higher standard in many aspects of my work. Special thanks go to Geoff While. I greatly appreciate how much time and effort you invested in my development and the many opportunities you supported me in pursuing. I really can look back on the work I present here with a sense of pride and satisfaction and you have had a huge part in realising that. All in all, I feel very lucky indeed to have had you as a supervisor and to now call you a friend. All other Co-authors who contributed to publications resulting from this thesis, including: Emily Cooper-Scott, Barbara Holland, Mike Gardner and Dave Chapple. All Administrative and Technical Staff who have helped me get this project off the ground. In particular, Adam Smolenski, Richard Holmes, Adam Stephens, Jodi Noble, Morgan Green and Clancy Carver. Akash Samuel for the cover photo used for this thesis. vi All the Eggs; I truly am a man blessed to have fallen in with such a glorious carton of double-yolkers. I love and thank you all for the many adventures we have shared over the past few years and look forward to many more! My family: Mark, Shauna, Eli, Anthea, Josh, Bek, Shum and Bron. Many thanks for your ongoing support and patience while I got this thing done! Mi querida Anna - para todo. ¡Lo hicimos! En este capítulo, finalmente termina y otra espera nuestra imaginación. No puedo esperar para explorar los años con ti como nos encontremos nuestro camino juntos. Te amo, mi fantástica Sra. Zorra. vii Abstract Sociality is one of the most captivating properties of animal life, encompassing a remarkable diversity of behaviour. As well as varying greatly in form and function, social organisation is taxonomically widespread, indicating convergent benefits of group living. The centrality of social interactions in many ecological processes also makes social organization crucial to our understanding of a range of evolutionary principles. Despite this, both the mechanisms by which natural selection has led to diversity in social systems, and the conditions promoting the initial emergence of group living, remain unclear. If we are to understand how sociality emerges and diversifies over
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