The Comparative Population Ecology of Two Semi-Aquatic Varanids
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The comparative population ecology of two semi-aquatic varanids James G. Smith 2007 The comparative population ecology of two semi- aquatic varanids James Gordon Smith B. Sc. (Hons) Melbourne A thesis submitted to satisfy the requirements for the award of the degree of Doctor of Philosophy in the Institute of Advanced Studies, School for Environmental Research, Charles Darwin University, Darwin, Australia January 2007 Declaration: I hereby declare that the work herein, now submitted as a thesis for the degree of Doctor of Philosophy of the Charles Darwin University, is the result of my own investigations, and all references to ideas and work of other researchers have been specifically acknowledged. I herby certify that the work embodied in this thesis has not already been accepted in substance for any degree, and is not being currently submitted in candidature for any other degree. Signed: Date: Acknowledgments I would like to thank the innumerable volunteers (particularly my friend “features”) I have had over the years for their assistance in the capture and recapture of goannas, most of the time in less than ideal circumstances. I am indebted to Keith Newgrain for analysis of all isotopic samples, the staff at Crocodylus Park, in particular Charlie Manolis and Yaakov Bar-Lev who generously allowed me to use their crocodiles, and Eric Cox, Jason Stevens and Greg Brown at Beatrice Hill Research Farm who provided logistical support. Peter Brocklehurst from the Northern Territory Government, Department of Natural Resources, Environment and the Arts provided invaluable GIS base layers for use in many analyses. I would also like to thank Thomas Madsen for the generous use of his field laboratory, Paul Horner for assistance and access to museum specimens, Corey Bradshaw for assistance in R programming, Jeremy Freeman for all things GIS, Gavin Bedford for advice and Julian Gorman for all sorts of help, on and off the touch rugby field. After much deliberation, Bob Richards let me continue to enter his barramundi farm to carry on catching V. indicus on his land, a decision that enabled us all to learn a little more about this amazing and reclusive creature. In addition, Rod and Lorraine kept me company and kept me laughing throughout my time in the mangroves. For this, and for all the after work beers, I thank the three of them. Of my three supervisors, I thank Keith Christian for his unquestioning assistance, Barry Brook for his immediate, concise and honest advice and most of all Tony Griffiths, for this wonderful opportunity and his enduring patience. There are so many other people in my life who deserve much kudos for their help in getting me through this, whether it was by distracting me with a frisbee, a guitar or a holiday, or by staving off my insanity by providing comprehensible discussions about statistics. Accordingly, all the people in this paragraph deserve equal ranking; Lorrae McArthur, Wendy Telfer, Jennifer Koenig, Ben (I concur) Phillips, Ron Firth, Chris Brady, Taegan (lil’ t) Calnan, Christopher (features) Schaefer, Brett Murphy, James (Lyndley) Mckay, Cameron Fraser, Steve Comber, Steve (big hair) Sinclair, Jeffrey Foucault, Tchavolo Schmitt, Florin Niculescu and Birelli Lagrene. To my parents, Jennifer and Gordon, my sisters Marinda and Samantha and to my beautiful Miranda, I extend my deepest gratitude and love. Jacinda Brown spent a joyous afternoon in the mangroves with me and I am grateful to her for some of the photos on the cover pages. Whoever reads this should head to Mindil beach markets in Darwin and buy some of her pictures, there might even be a mertens or an indicus in there! Albert Koomen and the Australian Broadcasting Commission also allowed me to use some stills that I had captured from the documentary “Goannas and Rubbish Frogs” that was made during this project. To the “other” indicus boys, Ron and Chris (and Geoffrey) I thank you mostly for keeping me laughing but also for keeping me cashed up. And finally, I can’t finish without thanking the inimitable yet humble mosquito. My constant companion in the mangroves, my joints still ache just at the thought of you… This work was funded by Charles Darwin University and a grant from the Natural Heritage Trust (NHT). All research was conducted under Animal Ethics Permit No. A00015 and Northern Territory Parks and Wildlife Permit No. 15254. Table of contents 1. Introduction........................................................................................................1 Varanid phylogeny and taxonomy........................................................................1 Varanid habits and morphology...........................................................................2 Varanid physiology...............................................................................................3 Significance of varanids in Australia ...................................................................4 Current threats to Australian varanid populations..............................................5 Previous studies of varanid ecology.....................................................................6 Seasonal responses by varanids in northern Australia........................................6 Varanids used in this study...................................................................................8 Why use a comparative approach ? ...................................................................10 This study............................................................................................................10 2. The potential impact of cane toads on Australian reptiles ...............................13 Abstract ................................................................................................................13 Introduction..........................................................................................................14 Methods ................................................................................................................19 Numbers of reptile species potentially at risk ....................................................19 Reptiles’ tolerance to toad toxins.......................................................................20 Results...................................................................................................................27 Species potentially at risk from toads.................................................................27 Tolerance to toad toxins.....................................................................................27 Discussion .............................................................................................................33 Crocodiles ..........................................................................................................34 Varanids .............................................................................................................35 Agamids..............................................................................................................35 Freshwater turtles ..............................................................................................36 3. Study sites and general methods .........................................................................39 Abstract ................................................................................................................39 Study Sites ............................................................................................................41 Manton Dam (Varanus mertensi).......................................................................41 Adelaide River (Varanus indicus) ......................................................................41 Animal capture.....................................................................................................44 Previous capture methods ..................................................................................44 Capture methods used ........................................................................................45 New pipe trap .....................................................................................................45 Effectiveness of design........................................................................................46 Animal processing ..............................................................................................50 Radio transmitter implantation..........................................................................50 Analytical framework..........................................................................................52 4. Using morphometrics to predict gender in varanids ........................................58 Abstract ................................................................................................................58 Introduction..........................................................................................................59 Materials and methods........................................................................................61 Model development.............................................................................................64 Model fitting .......................................................................................................65 Model selection...................................................................................................65 Characteristic body size .....................................................................................68