Diet, spatial ecology and energetics of echidnas: the significance of habitat and seasonal variation Jennifer Anne Sprent BSc (Hons), University of Tasmania Submitted in fulfilment of the requirements for the degree of Doctor of Philosophy University of Tasmania February, 2012 Declaration of originality I hereby declare that this thesis contains no material which has been accepted for the award of any other degree or diploma in any tertiary institute, and that, to the best of my knowledge and belief, the thesis contains no material previously published or written by another person, except where due reference is made in the text of the thesis, nor does the thesis contain any material the infringes copyright. Jenny Sprent February 29th, 2012 Statement of authority of access This thesis may be made available for loan and limited copying in accordance with the Copyright Act 1968. Jenny Sprent February 29th, 2012 ii Statement regarding published work contained in thesis The publishers of the paper comprising Chapters 5a 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. Jenny Sprent February 29th, 2011 Statement of Ethical Conduct The research associated with this thesis abides by the international and Australian codes on human and animal experimentation, the guidelines by the Australian Government's Office of the Gene Technology Regulator and the rulings of the Safety, Ethics and Institutional Biosafety Committees of the University. Jenny Sprent February 29th, 2011 iii Statement of publication and co-authorship The following people and institutions contributed to the publication of the work undertaken as part of this thesis: Paper 1: Latrine use by the short-beaked echidna, Tachyglossus aculeatus. Australian Mammalogy Candidate (50%), author 1 (Niels Andersen) (25%), author 2 (Stewart Nicol)(25%) Details of the Authors roles: Authors 1 and 2 contributed to the concept and planning of the study, and to data collection. Author 2 assisted with data analysis. We the undersigned agree with the above stated “proportion of work undertaken” for the above published peer-reviewed manuscript contributing to this thesis: Signed: __________________ ______________________ Stewart Nicol Elissa Cameron Supervisor Head of School School of Zoology School of Zoology University of Tasmania University of Tasmania Date:_____________________ iv The following people contributed to the research undertaken as part of this thesis Stewart Nicol (School of Zoology, University of Tasmania) provided guidance on data collection and assistance with statistical analysis and provided constructive criticism of data analysis, interpretation of results and chapter drafts. Sue Jones (School of Zoology, University of Tasmania) supervised my radioimmunoassay work (Chapter 2) and provided constructive criticism of data analysis, interpretation of results and drafts of all chapters. Niels Andersen (School of Medicine, University of Tasmania) provided field assistance and guidance on data collection (Chapters 2, 3, 4 and 5) and constructive criticism of data analysis, interpretation of results and chapter drafts (Chapters 5a). Rob Gasperini (Menzies Research Institute) performed western blot analysis of leptin (Chapter 2) Craig Mundy (TAFI, University of Tasmania) calculated home range estimates in R and converted them into a format suitable for use in a GIS (Chapter 4). Mark Hovenden and Mike Perring (School of Plant Science, University of Tasmania) calculated pasture productivity at Lovely Banks (Chapter 6). This thesis is based on field data collected by me in the period of August 2004 to October 2010, but also included data collected as part of a long term project of echidna field biology which was started in 1997. A small number of additional data collected by other researchers has been included in the data analysis for Chapter 2 - Seasonal variation in circulating leptin concentrations and body mass in free ranging short beaked echidnas (Tachyglossus aculeatus) and Chapter 4 - The influence of habitat type on the home range size of the short beaked echidna (Tachyglossus aculeatus). v Abstract Abstract The ant- and termite-eating echidna is a solitary, monotreme mammal with a highly seasonal life history. I investigated several aspects of the seasonal energetics and ecology of free-ranging echidnas: the role of leptin in the annual mass cycle, seasonal and habitat effects on diet, the relationship between home range size and habitat quality, and the effect of habitat on the siting of echidna latrines. The echidna has a large seasonal variation in fat stores, which reach their maximum prior to hibernation. I hypothesised that the hormone leptin would have the same role in the echidna as in eutherian hibernators, i.e. that there would be a direct relationship between body mass and plasma leptin that would change to allow pre-hibernatory fattening. I found significant seasonal variations in plasma leptin, with the highest levels occurring in hibernation and in females during mating. The lowest levels were found in males after the reproductive period. Rather than the expected strong positive relationship between adiposity and plasma leptin I found a weak negative relationship, similar to that in reptiles and birds. To determine if there was any significant seasonal variation in diet associated with pre- hibernatory fattening I investigated diet using scat and stable isotope analysis. Echidna scats consisted largely of ants and the larvae of pasture cockchafer beetles. There was significant seasonal variation in percent occurrence of larvae, but not in the ant species found in scats or in the stable isotopic composition of echidna plasma. Although there was no difference in the prey items that contributed most to scat contents of animals living in different habitats, stable isotope analysis of blood showed a highly significant effect of habitat type on δ15N. Female echidnas showed a significant negative relationship between the proportion of woodland habitat and home range size, whereas there was no such relationship for males, which had significantly larger home ranges. My data suggest that female home range is scaled to available resources, while male home range is probably scaled to maximise access to females. The role of latrines and their relationship to habitat was examined by a detailed survey of part of the field site, where many echidna home ranges were known to overlap. Latrines were located more frequently in scrub than in pasture or thick bush and the highest frequency of latrines occurred where there were two home ranges that overlapped. Latrines may be important in the spatial organisation of echidnas. viiv Acknowledgments Acknowledgements Firstly my heartfelt thanks must go to my supervisor Stewart Nicol. Your comments, suggestions and guidance have greatly improved the quality of my work over the years, and your enthusiasm, encouragement and assistance were a major driving force in getting this thesis completed, especially in the last few painful months. Thanks to my co- supervisor Sue Jones for taking an active interest in my research, for your constructive comments and suggestions. Special thanks from pregnant me, and unborn Hamish, for running one of my radioimmunoassays for me when I was pregnant. Peter McQullian provided an enormous amount of information about pasture grubs, ants and how to identify them. Peter you are a mine of information, and whenever I got to opportunity to discuss my work with you I always came away with some great ideas. A large number of people assisted me throughout my project. I am most grateful to Niels Andersen for all his help in the field. Thanks for your company when driving around on the hunt for echidnas, as well as for your patience whilst I mastered the art of radio tracking and anesthetising echidnas. I won’t miss your stinky sandwiches though! Thanks to office mates and fellow postgraduates Gemma Morrow and Rachel Harris for help and company during field work as well as providing support, encouragement, advice and cupcakes. I am indebted to Rob Gasperini for advice about, and running the western blot for me, as well as attempting to explain what on earth it was all about. The GIS part of this work would have not been possible without help and advice from Arko Lucieer, Pip Bricher and Stephen Harwin, and I am most grateful to Craig Mundy for plugging my data into his R script, and thus allowing me to escape the tyranny of R. Much needed, and appreciated, statistical advice was provided by David Ratkowsky. Thank you to owners of Lovely Banks, the McShane family, for allowing access to our field site and the echidnas that inhabit it. A generous donation of two GPS logger units from NewBehavior in Switzerland made trialling the attaching GPS loggers to echidnas possible. Dick Bashford at Forestry Tasmania was kind enough to allow me to look at the Lowery ant collection. vii Acknowledgments Thank you to Dad, Chris and Mum for your continued support and interest in my work. Dad thank you for all that time you spent designing and machining the various bits and pieces that held GPS loggers onto echidnas, and Chris and Mum – thank you for being the best Granny/Gramma a little boy could wish for. It certainly made it easier going to uni knowing that Hamish loves spending his days with you. Finally thanks to Chris and Hamish for your love, support and above all, tolerance. I dedicate this work to you because without you it never would have been possible. Thanks you for your unwavering confidence in my ability to complete this, especially in those dark moments when I was awash with self-doubt. Now it is all over I can a) stop stressing, b) stop working most of the night and be part of the family again and c) get a job and start paying some bills (hopefully).