Resource Use and Coexistence of Sympatric Tasmanian Devils and Spotted-Tailed Quolls
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Resource use and coexistence of sympatric Tasmanian devils and spotted-tailed quolls Georgina Elizabeth Andersen (MSc, BSc) Submitted in fulfilment of the requirements for the Degree of Doctor of Philosophy School of Biological Sciences University of Tasmania May 2016 Statements by the author Declaration of originality This thesis contains no material which has been accepted for a degree or diploma by the University or any other institution, except by way of background information and duly acknowledged in the thesis, and to the best of my knowledge and belief no material previously published or written by another person except where due acknowledgement is made in the text of the thesis, nor does the thesis contain any material that infringes copyright. Authority of access This 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 was conducted in compliance with the Australian Code of Practice for the Care and Use of Animals for Scientific Purposes. The research presented in this thesis was carried out in accordance with the University of Tasmania Animal Ethics Committee Permit # A12361 and with permission from the Tasmanian Department of Primary Industries, Parks, Water and Environment (DPIPWE) under the scientific permits TFA12166 and TFA13969. Signed: _______________________________ Georgina E. Andersen 03/05/2016 2 Statement of co-authorship The following people contributed to the work undertaken as part of this thesis: Georgina Andersen (Candidate), School of Biological Sciences, University of Tasmania, Hobart, Australia Menna Jones (Primary supervisor), School of Biological Sciences, University of Tasmania, Hobart, Australia Christopher Johnson (Secondary supervisor), School of Biological Sciences, University of Tasmania, Hobart, Australia Leon Barmuta (Co-author on chapter 2 and 4), School of Biological Sciences, University of Tasmania, Hobart, Australia Chapter details and relative author contributions are listed below: Chapter 2 is in preparation for publication: Andersen, G.E., Johnson, C.N., Barmuta, L.A., and Jones, M.E. Diet composition and overlap of two sympatric carnivores, the Tasmanian devil and spotted-tailed quoll Georgina Andersen was the primary investigator and lead author. Chris Johnson and Menna Jones contributed to the idea, the study design and edited chapters. Leon Barmuta assisted with statistical advice. Chapter 3 is in preparation for Wildlife Research: Andersen, G.E., Johnson, C.N., and Jones, M.E. Spatial ecology of sympatric Tasmanian devils and spotted-tailed quolls. Georgina Andersen was the primary investigator and lead author. Chris Johnson and Menna Jones contributed to the idea, the study design and edited chapters. Chapter 4 is in review in Journal of Applied Ecology: Andersen, G.E., Johnson, C.N., Barmuta, L.A., and Jones, M.E. Anthropogenic habitat modification enhances traveling and hunting opportunities for two medium-sized carnivores. 3 Georgina Andersen was the primary investigator and lead author. Chris Johnson and Menna Jones contributed to the idea, the study design and edited chapters. Leon Barmuta assisted with statistical advice. Chapter 5 is published in Behavioural Ecology and Sociobiology: Andersen, G.E., Johnson, C.N., and Jones, M.E. Sympatric predator odour reveals a competitive relationship in size-structured mammalian carnivores. Georgina Andersen was the primary investigator and lead author. Chris Johnson and Menna Jones contributed to the idea, the study design and edited chapters. We the undersigned agree with the above stated “proportion of work undertaken” for each of the above submitted peer-reviewed manuscripts contributing to this thesis: Signed: Signed: Menna Jones Anthony Koutoulis Primary Supervisor Head of School School of Biological Sciences School of Biological Sciences University of Tasmania University of Tasmania 4th November 2016 4th November 2016 4 Acknowledgements I would first like to thank my supervisors Menna Jones and Chris Johnson for letting me embark on this great project, which has taught me so much and challenged me in many ways. Thank you Menna for your enthusiasm, support and many discussions on devils and quolls. Thank you Chris for responding to my email enquiring about a PhD and for offering me such an amazing project. You always helped me see the bigger picture and provided sound advice when I needed it. Much of my PhD was spent in the beautiful Arthur-Pieman Conservation Area (APCA), which quickly became my second Tassie home. A huge thank you to the rangers of the APCA, for all your help over the past 4 years and for always welcoming me back! A special thanks to Mel Pfieffer, who became a good friend and constant support during field work. Thanks for putting up with some major bush bashing expeditions after dropped GPS collars and for chats over morning tea. Thank you Greg Cash for providing me with the nicest field accommodation, despite the kookaburra attacks! I could not have successfully completed field work without the help of all my volunteers. So a massive thank you to all those that helped me check traps at the crack of dawn and radio-track/bush bash come rain or shine. My field work acknowledgements would not be complete without thanking all the devils and quolls that took part in this study. I have had some truly memorable experiences during field work that I will always cherish. I also would like to thank several funding bodies who made this project possible: The Holsworth Wildlife Research Endowment, the Dr Eric Guiler Tasmanian Devil Research Fund and the Australian Research Council. I was the recipient of the Australian Postgraduate Award and the Tasmanian Devil Elite Scholarship. To the people at UTAS, a huge thanks goes out to the various office mates that have helped keep me sane, whist I was in the office: Halley Durrant, Shannon Troy, Lisa Cawthen and Paul Botwe. A special thanks to Halley Durrant, for all the laughs, the morning coffee chats and just for making being in the office a little more bearable! A 5 big thanks to Shannon Troy for helping me find such an amazing study site, for statistical help and for proof reading my general introduction and discussion. Thanks to the past and present postgrads for your support and friendship especially Anna Bruniche Olsen, Nick Fountain-Jones, Hugh McGregor, Bronwyn Fancourt, Amy Edwards, Jo McEvoy, Vince Scoleri, Sebastian Comte, David Hamilton and Rowena Hamer. I would also like to thank the various staff members in the School of Biological Sciences for help with everything administrative: Felicity Wilkinson, Adam Stephens, Jenny Sprent, Richard Holmes, Wayne Kelly, Barry Rumbold, Jodi Noble, Clancy Carver and Erik Wapstra. Thank you to Barry Wells for animal welfare advice and teaching me how to sedate feral cats. I’d also like to thank Katrine Sorensen and Amy Dunlop for helping to identify prey species from scats. I am also very grateful to Leon Barmuta and Joanna Potts for statistical advice and R code help. I would have been seriously stuck without your help! Thanks to the soccer team at uni for providing some much needed distraction during the write up and for giving me a good excuse to kick something and occasionally someone, oops To my friends back in Denmark, Mette Richner, Inge Findorf, Louise Toft Iversen and Maria Jellesmark, I cannot thank you enough for your love and support throughout the years. Thanks for always remaining my friends no matter where in the world my travels take me! I am eternally grateful to my parents and sister for your unconditional love and support. You have always encouraged me to follow my dreams and helped me along the way whenever I needed it. Finally, I would like to dedicate this thesis to the late Geoff King, the king of conservation in the North West. My first field season would have been a lot harder without your knowledge of the devils and quolls in the APCA and your help whenever I needed it. As we used to say “long live the devils of Arthur River” 6 Abstract Interactions between sympatric carnivores are among the key factors influencing community structure and function, and strongly affecting the population dynamics, distribution and behaviour of the interacting species. Interspecific competition occurs in two ways: exploitation competition occurs when a resource unit is consumed by one species so it cannot be consumed by another; interference competition involves direct aggressive encounters (e.g. fighting) or the threat of aggression, thereby excluding a competitor from a resource. The strength of competition between carnivores varies with factors such as body size, diet and population density. Interspecific competition is often asymmetrical such that the smaller carnivore is most affected. Due to the competitive effects that apex predators can have on sympatric carnivores, their removal from a system can result in a ‘mesopredator release’, which in turn can lead to increased predation on smaller prey species. To minimise competition and facilitate coexistence, the subordinate species can either avoid spatial overlap with the dominant species or modify temporal or behavioural patterns to reduce the chance of encounters and interactions, but still allow for spatial overlap. The intensity of competition can be reduced in species that have co- existed for a long time through coevolution of divergent ecomorphological and behavioural traits that influence the size and type of prey that are eaten. Current competition can be difficult to measure, as competition may only occur when resources are severely limited such as during drought, and can be subtle where there is a long coevolutionary history among the species. As interspecific competition between carnivores can strongly influence abundance and distribution, understanding how carnivores interact and the extent that competition might influence their ecology and demography is crucial in managing and conserving them. The Tasmanian devil (Sarcophilus harrisii) is Tasmania’s largest marsupial carnivore (5 - 14kg) and coexists with the smaller spotted-tailed quoll (Dasyurus maculatus) (0.9 - 5kg).