Asian Eden Large herbivore ecology in India Thesis committee Thesis supervisor Prof. dr. H.H.T. Prins Professor of Resource Ecology Wageningen University Thesis co-supervisor Dr. ir. I.M.A. Heitkönig Assistant Professor Resource Ecology Group Wageningen University Other members Prof. dr. F.J.J.M. Bongers, Wageningen University Dr. G.W.T.A. Groot-Bruinderink, Alterra Prof. dr. W.H. Hendriks, Wageningen University Prof. dr. P.J.H. Reijnders, Wageningen University This research was conducted under the auspices of the C.T. de Wit Graduate School of Production Ecology & Resource Conservation. Asian Eden Large herbivore ecology in India Farshid S. Ahrestani Thesis Submitted in partial fulfilment of the requirements for the degree of doctor at Wageningen University by the authority of the Rector Magnificus Prof. dr. M.J. Kropff, in the presence of the Thesis Committee appointed by the Doctorate Board To be defended in public On Monday 30 November 2009 At 4 PM in the Aula. Farshid S. Ahrestani Asian Eden - Large herbivore ecology in India 110 pages. Thesis, Wageningen University, Wageningen, NL (2009) With references, the summaries in Dutch and English ISBN 978-90-8585-487-5 Contents Acknowledgements……………………………………………………………………………...i Abstract…………………………………………………………………………………….…..iii 1. General Introduction……………………………………………………………….…...1 Farshid S. Ahrestani 2. Seasonal variation in herbaceous biomass and nitrogen content in South India: implications for large herbivores…………………………………………………….....9 Farshid S. Ahrestani, Ignas M.A. Heitkönig and Herbert H.T. Prins 3. The role of body mass in diet and habitat selection by large herbivores species in a tropical South Indian forest……………………………………………….…………...23 Farshid S. Ahrestani, Ignas M.A. Heitkönig and Herbert H.T. Prins 4. Why some species time their annual parturition with high plant quality availability? - The impact of body mass………………………………………………..….………..35 Farshid S. Ahrestani, Frank van Langevelde, Ignas M.A. Heitkönig and Herbert H.T. Prins 5. Age and sex determination of gaur, Bos gaurus…………………………...…………..47 Farshid S. Ahrestani and Herbert H.T. Prins 6. First detailed analysis of life history traits of the Asian megaherbivore, gaur Bos gaurus….……………………………………………………………………..55 Farshid S. Ahrestani, S. Iyer, Ignas M.A. Heitkönig and Herbert H.T. Prins 7. Determinants of large herbivore diversity in sub-Himalayan India…………………....69 Farshid S. Ahrestani, Ignas M.A. Heitkönig, Frank van Langevelde, V. Srinivas, M.D. Madhusudan and Herbert H.T. Prins 8. Body mass and large herbivore ecology in India: a synthesis………………………….85 Farshid S. Ahrestani References……………………………………………………………………………………..95 Summary………………………………………………………………………………….….111 Samenvatting………………………………………………………………………….…...…113 Affiliations of Co-authors………………………………………………………...….………115 Curriculum Vitae…………………………………………………………………….….……116 PE&RC PhD Education Certificate………………………………………………….…..…..117 Acknowledgements Acknowledgements This PhD wouldn’t have been possible but for the tireless support of my wife Savita to whom this thesis is dedicated with love, respect and gratitude. To have travelled thousands of inter- continental miles (often alone with our two young children), moved and lived in three different countries, ensured that our home always remained a warm and stimulating environment, befriending people in all three countries (many of whom have become dear friends) so that we had a semblance of a normal social life, being responsible for raising our two young children, and while all along working late and odd hours to support the family financially are all testament to the sacrifices Savita made and the strength and love she provided so that I remained sane and on track to see this through. This PhD, however, took shape and reached its logical conclusion because of the vision, encouragement, and inspiration of my advisors at Wageningen University. From the immediate positive reply to my first email, always remaining accessible, and through many intelligent and stimulating ideas, advice and encouragement I always felt that I would achieve my goal under the visionary, stimulating, and inspiring leadership of Herbert Prins. Equally important was the inexhaustible clever advice, encouragement, reviews, ideas and support from Ignas Heitkönig, who helped me secure the all-important WOTRO grant, develop robust research questions, and maintain a high quality of science throughout. In addition, Sip van Wieren and Frank van Langevelde were also invaluable resources of knowledge, ideas and inspiration; Sip for his in-depth knowledge of large herbivore physiology and ecology, and Frank for his amazing grasp and knowledge of modelling. The above list wouldn’t be complete without a mention of Fred de Boer, whose hard working ethic and personal warmth left an indelible mark on me. To all of you dank U wel. The Resource Ecology Group, however, extended beyond my advisors, and the following people shared and provided me with ideas, emotional support, stimulating debates, constructive criticism, and inspirational individual life stories over numerous cups of coffee, beer and sandwiches: Anil, Arend, Audrey, Benson, Christine (thanks for all the invaluable technical help and words of wisdom), Christiaan, Cornelius (thanks for being a dear friend and a pillar of strength), Edward, Eduardo, Edson, Emmanuel (for your soft-spoken straight talking and warm smile), Euridice, Geerten, Herman (your enthusiasm to help was uplifting), Henjo (thanks for all your advice and help that you were always prepared to share), Ivonne, Jasja, Jasper, Kyle, Martijn (for your special flavour of Dutch warmth), Milena, Nicky, Nicole (for your warmth and unselfish friendship), Xavier, Priya, Ralf, Rudy (much thanks for the translation of my Summary), Thomas, Tom, Tom, Tsewang, and Yolanda (your laugh and passion for elephants will remain with me forever). This list however would never be complete without a very special thanks for the superb consistent high quality administrative support that I got throughout from Gerda, Margreet, Maureen, Patricia, Petra and Willemien. I would also like to thank Nelleke for the warm hospitality during my one week stay at the Heitkönig residence, and Pat for her warmth and hospitality over a few lovely meals. Support in the Netherlands came from beyond the University too and with a special warmth and hospitality of delightful ‘Mum’ and other dear friends like Charlotta & Shammy; Christine & Guillaume; Gina & Dominick; Keli & William; Manuela & Giacomo; Rosario & Nicholas; Savitri & Herbie; Valentina & Camillo; and Vidhya & Prithvi. In addition to the vital support from all those in the Netherlands, many people and institutions helped to make sure that I completed my field work in India successfully. My collaboration with NCF was a very important component of my PhD and I gratefully acknowledge the important scientific input, superb administrational support, inspirational leadership, and uplifting social interactions that the following at NCF provided: Aaron, Ambika, Aparajita, Ayesha, Charu, Divya, Madhusudan, Naveen, Navinder, Praveen, i Aknowledgements Raghunath, Rakhee, Rana, Rohan, ShankarRaman, Sumanto, Vena, Vishwanathan, and Yashveer. Other people and institutions who helped with data collection and analysis and improved the scientific quality of my thesis were: Srini (dai NEP thanks a ton); Madhu Nair (NBSSLUP); DCF Manoj Kumar, Nanjappa and Shivanna (Mysore Zoo); Narayanan and Abhi (CAWA); Nageswara Rao, Atul Kolte and Manju (NIANP); J.C. Daniel and Asad Rehmani (BNHS); Sheshashayee (GKVK); Gopalkrishna (Mamta Laboratories); and Raman Sukumar, C. Suresh and Nandita (CES). During my time in India I was also helped with dollops of warm hospitality, shelter and delicious food by a score of friends that made my stay in Mysore and visits to Bangalore and Chennai that much more easier and enjoyable: Arvi, Nisha, and Mr. & Mrs. Srinivasan; Bejoy, Beena, Chechi and Nethra; Ganesh; Khunger and Rashmi; Kita and Divya; Mudaliar; Salim, Zafar and Reshma; Sanjeev and Stephanie; Sanjiv and Tina, and, Srini, Sunita and Aunty. If not for the support, care and hard work of Anthony, Gauramma, and Manju my stay in Mysore would not have been so pleasant, and friends like Cat, Hemant & Benedict, Milli, and Moosa were big help during the journey. This list however would be far from complete without acknowledging my gratitude towards the people living and working in and around my study site who helped me with my data collection. If not for Manba, who was much much more than just a field assistant, this PhD would never have seen the light of day and for which I am humbly grateful. I will always cherish the jungle craft you taught me, the magical times we spent tracking gaur and spotting wildlife together, the fun we had driving and walking all over our field site, and all the other small and big things you did for me during those two years. collecting data. I am also grateful to Murali for his hard work and dedication, and Boma, Ketha, Kirikila, Madha, Naseem (I will remember your smile always), and Pambu Boman for their assistance in data collection. I am thankful for the Tamil Nadu and Karnataka Forest Department for their tireless dedicated stewardship and hard work and for providing me with the permission to conduct my study and logistics at both Mudumalai and Bandipur. I would also like to thank all the staff at both Bandipur and Mudumalai that helped with logistical support. To name just a few: PCCF Sukhdev; DCF Dogra; Ranger Ramalingu; watchers