INTRAGUILD INTERACTIONS BETWEEN NATIVE AND DOMESTIC CARNIVORES IN CENTRAL INDIA _________________________________ A Dissertation presented to the Faculty of the Graduate School at the University of Missouri __________________________________________________ In partial Fulfilment of the Requirements for the Degree of Doctor of Philosophy __________________________________________________ by ABI TAMIM VANAK Dr. Matthew E. Gompper, Dissertation Supervisor December 2008 The undersigned, appointed by the dean of the Graduate School, have examined the dissertation entitled INTRAGUILD INTERACTIONS BETWEEN NATIVE AND DOMESTIC CARNIVORES IN CENTRAL INDIA presented by Abi Tamim Vanak, a candidate for the degree of Doctor of Philosophy, and hereby certify that, in their opinion, it is worthy of acceptance. ______________________________________________________ Matthew E. Gompper ______________________________________________________ Mark R. Ryan ______________________________________________________ Joshua J. Millspaugh ______________________________________________________ Hong S. He ______________________________________________________ Mark R. Ellersieck ACKNOWLEDGEMENTS In a seminal paper destined for Nature, that Morgan Wehtje and I plan to eventually write, we propose a new system of classifying the relationship between students and faculty advisors based on their study systems. Much like the free-ranging carnivores that he studies, Matt treated us the same, letting us wander off to do our own thing at the end of a long rope, occasionally monitored by radio-telemetry, but if lost, always welcome to tug on that rope and return to the den for new direction over a cup of coffee. For all this and more, to my dissertation advisor, Matt Gompper, many thanks! My heartfelt thanks also to Matt’s wonderful family, Anne and the kids, Izzie and Alex. Theirs was the first home I knew in the US, and to me it shall most certainly remain one of the warmest. I acknowledge the contribution of my dissertation committee members, Drs. Mark Ryan, Josh Millspaugh, Hong He, and Mark Ellersieck. This dissertation has improved greatly due to their inputs, from the proposal to the final defense. Thanks especially to Dr. Millspaugh for his help with the latest in home-range and resource selection analyses. The research undertaken for this dissertation would not have gotten off the ground if not for the financial assistance provided by the following funding agencies: University of Missouri Research Board – Grant to M. Gompper, Wildlife Conservation Society Research Fellowship Program, USA, Rufford Small Grants Foundation, UK and the University of Missouri Conservation Biology Fellowship. My heartfelt thanks also to the Maharashtra State Forest Department (MFD), in particular the Chief Wildlife Warden, Mr. B. Majumdar. After struggling and failing after ii a year and a half to obtain permits from the Andhra Pradesh Forest Department, it was a pleasure to interact with the MFD. A large part of this is due to the interest in this project shown by Mr. Majumdar, who, despite his position as the head of a large government agency, is still one of the most accessible as well as astute people I have come across in the Indian government service. The field staff of the Great Indian Bustard Sanctuary were also helpful, and provided me with logistic support at Nannaj. I also received great support in the beginning of the project from Mr. D. Parihar, the Conservator of Forests, Pune. I thank Dr. Brian Cypher (ESRP, California) for all of his advice and help and for letting me “wrassle” with his kit foxes in Bakersfield. He also generously gave me the first five radio-collars for my foxes. The rest of this acknowledgement section is a world apart – one for folks in the US and the other for people in India. To folks in India: I would like to begin with thanking my family in Chennai, for always being there for me, and to my dad for also generously bearing the upfront costs for a field vehicle when I did not have sufficient funds at the start of the project. I was fortunate to find a rather fine field crew, headed by Mr. Abhijeet Kulkarni, my “field station manager”, plumber, electrician, carpenter, general handyman and good friend. Others in the team included Sarang Mhamane, Vinayak Shitole and for a thankfully brief period Salim Ansare. I thank Aditya Bhaskaran for this help with the field work and scat analysis. My most sincere thanks to Dr. Aniruddha Belsare, Wildlife Veterinarian extraordinaire – this project would have struggled to do half of what it did, if iii not for his help. I know that I can always count on him for his friendship for the rest of my life, and look forward to reading his eulogy to me in his dissertation acknowledgement five years hence. I thank Dr. Asad Rahmani for his support in the initial stages of the project and for his letter of collaboration that helped in get my project off the ground in Maharashtra. I also thank Dr. Bilal Habib for his hospitality when I first visited Nannaj, and for sharing the results of his research with me. I would like to thank Mr. Bharat Chheda, and the Kulkarni family in Solapur for their tremendous help, friendship and hospitality. Bharat – thanks also for taking care of my dog Permit for me. I also thank Amod Zambre for his help with scat analysis and reptile scale identification. Vidya and Ramana Athreya were very gracious hosts on my many visits to Pune. Kishor Mungale generously gave of his time, brainpower and workforce in outfitting my truck with a mobile telemetry antenna tower – at no charge. I am very grateful to Mr. Milind Bendale and Mr. R. Nale for their friendship and for helping me in obtaining my permits for field research. In the USA: The excellent faculty and staff of the Department of Fisheries and Wildlife Sciences at the University of Missouri deserve special mention and my gratefulness. I would especially like to acknowledge Dr. Jack Jones, the chair of the department as well as all the secretarial staff, especially Karen Decker. In the later part of my time at Mizzou, especially during the dissertation analysis and writing process, I was immensely helped by several people, including Tom Bonnot, Bill Dijak, Dave Jachowski, Damon Lesmeister, Ryan Monello, Morgan and Walter iv Wehtje, Karthik Masagounder and several other graduate students of the F&W department. I thank them all very much for their time and patience in enduring my constant pestering. I also thank Aditya Singh (U. Wisconsin) for being such a GIS guru and for coming up with solutions to my analysis problems in a jiffy. I leant heavily on him for several GIS analyses and he always came through for me. I know there are many others here who I have inadvertently forgotten, and I thank you for your help and hope that you don’t get cross with me while reading this section of the dissertation. Finally, and always, I want to thank my wife Maria Thaker – for everything. The chapters of this dissertation are either published, in review or in preparation for submission to peer-reviewed journals. Below I detail the citations or eventual destination of the chapters: Chapter 1: Gompper, M.E., and Vanak A.T. 2006. Vulpes bengalensis Mammalian Species, Mammalian Species No. 795, pp. 1–5 Chapter 2: Vanak A.T., and Gompper, M.E. Domestic dogs as carnivores: their role and function in intraguild competition. In review in Mammal Review Chapter 3: Vanak A.T., and Gompper, M.E. Spatial ecology of the Indian fox in a dry grassland ecosystem in central India. To be submitted to Journal of Animal Ecology Chapter 4: Vanak A.T., and Gompper, M.E. Dietary niche separation between sympatric free-ranging domestic dogs and Indian foxes in central India. In review in Journal of Mammalogy v Chapter 5: Vanak A.T., Thaker, M., and Gompper, M.E. Experimental examination of behavioural interactions between free-ranging wild and domestic canids. In review in Behavioural Ecology and Sociobiology Chapter 6: Vanak A.T., and Gompper, M.E. Spatial niche overlap between domestic dogs and Indian foxes in a human-dominated landscape in central India. To be submitted to Ecology letters vi TABLE OF CONTENTS ACKNOWLEDGEMENTS..............................................................................................ii LIST OF TABLES...........................................................................................................xii LIST OF FIGURES..........................................................................................................xv ABSTRACT..................................................................................................................xviii Chapter 1: Vulpes bengalensis: THE INDIAN FOX.....................................................1 ABSTRACT.......................................................................................................................1 TAXONOMIC CLASSIFICATION..................................................................................2 Context and content................................................................................................3 Diagnosis................................................................................................................3 General characters.................................................................................................3 Distribution ............................................................................................................5 Fossil record...........................................................................................................6