Demography and ranging behaviour of lions (Panthera leo) within a human-occupied landscape in northern Kenya A thesis submitted to the University of Oxford for the degree of Doctor of Philosophy in Zoology Shivani Bhalla Lady Margaret Hall 2017 DECLARATION The thesis presented here represents a project that I developed as a D.Phil. student of the Zoology Department, University of Oxford. I implemented all aspects of this project, analysed the results and wrote all of the material in the thesis. While others assisted with the project at various stages including fieldwork, data entry and analysis, it represents a work entirely of my own doing and I assume full responsibility for the work presented here. The text, excluding figure legends, tables, references and appendices, does not exceed 50,000 words. All photographs included in the thesis are the copyright of the Ewaso Lions Project, except where stated. Signed Date: 18th April 2017 i ABSTRACT Lion populations across Kenya are threatened, primarily as a result of habitat loss and human persecution in response to livestock depredation. This study provides the first population insights into lions within the Samburu-Isiolo ecosystem; considering both the protected area network and the surrounding human-occupied landscape. These results are particularly pertinent given Kenya’s low lion population. The demography of the lion population in the Samburu-Isiolo ecosystem was studied to provide data on the area’s basic lion population structure. Due to the small size of the protected areas within the study area, it was expected that lions would frequently move outside the Reserves into the human-occupied landscapes that surround the protected areas and, therefore, their ranging behaviour was also assessed. Community Conservancies exist around the protected areas, where wildlife conservation is encouraged. However, the increasing human and livestock populations within these areas conflicts intensely with the spatial requirement of lions. This has numerous implications; from direct mortality to reducing the opportunities for immigration of new lions into the protected areas, or safe dispersal from them. Therefore, the presence of suitable habitat in one community area adjacent to the protected areas was examined using a Habitat Suitability Model (HSM) in order to explore these issues. The demography of the lions within the study area was found to be comparable to other populations across Africa, although displayed higher sex ratios, and lower cub dispersal figures. Whilst the ranging behaviour of the prides displayed core ranges along the rivers, it was noted that they each had their distinct areas of intensive use. Male ranges were larger than females and shrunk during the drought in 2009 and expanded again in 2010. Despite displaying ranges within the limits of the protected areas, it is known that between 2008 and 2010, 10 lions disappeared and moved outside the protected areas. The HSM showed that highly suitable habitat did exist within the Community Conservancy, highlighting the presence of safe refuges for carnivores, with more suitable habitat found to be available at night compared to during the day. Despite the presence of highly suitable habitat, human-lion conflict was a common occurrence in these areas. Anthropogenic factors will have an impact on the demography of lion populations, whether they exist inside or outside protected areas. If suitable habitat did not exist within the human-occupied landscape, it is expected that there would be reduced immigration of new males, longer pride tenures within protected areas and the potential risk of inbreeding. Conflict mitigation measures are important in reducing human-lion conflict, however, this study also recommends the presence of safe refuges in the form of Conservation Areas within the Community Conservancy network in northern Kenya. The model of Conservation Areas acting as safe refuges is essential for the conservation of lions outside protected areas throughout remaining parts of their range in northern Kenya. iii ACKNOWLEDGEMENTS I would like to thank the following people, organisations and entities for their contribution to the production of this thesis. The Government of Kenya, the National Council of Science and Technology, the Kenya Wildlife Service, the Samburu and Isiolo County Councils and Westgate Community Conservancy for giving me permission to work in northern Kenya. Professor Fritz Vollrath, my supervisor, and all the wonderful staff at the University of Oxford – including Heather Green, Lynne Bradley, Chris Holland, Jelle Ferweda who welcomed me in the Department of Zoology; Ketlin Stroo, Anke Rehm, Rev. Allan Doig who created a fantastic support system for me at Lady Margaret Hall; my committee in Oxford - Marian Dawkins, Nigel Winser, Mark Stanley Price whose guidance and academic and moral support has enabled this study to happen. My many generous sponsors - the Wildlife Conservation Network and Eve Schaeffer (Sidney Byers Scholarship); the African Wildlife Foundation and Daudi Sumba (Charlotte Fellowship); Panthera Foundation (Kaplan Graduate Award); the Rufford Small Grants Foundation (funds for fieldwork); Lady Margaret Hall (Warr-Goodman Scholarship); University of Oxford (Oxford Hardship Fund). The Douglas-Hamilton family - Oria “Mama 1” who first introduced me to Samburu and gave me shelter and much support; Saba for her invaluable knowledge and who generously taught me everything she knew about lions; Iain who always provided support and assistance – without them all, this study would never have taken place. Save the Elephants – my second home in Samburu during my early years. Lembara – for his warm welcome at all times; Jerenimo Lepirei, Chris Leadismo, Gilbert Sabinga for valuable additional lion sighting information; David Daballen – a friend and huge support; Jake Wall, George Wittemyer, Festus Ihwagi, Barnerd Lesowapir who provided important advice on data; Wainaina Kimani for his friendship and support; Njoroge Nganga for administrative assistance; Lucy King whose words of encouragement and advice meant a lot. To the team of people for the time and effort they took with data entry - Christina Tsantes, Taylor Tully, Joseph Basele, Aimee Guha Roy, Mat Beckwith, Benjamin Loloju, Lauren Ross; and for their support in the crucial, final year - Patrick Kabatha, Anu Khoda and Suzanne Offord. Paul Lolchuragi, Ricila Lelenkeju, Raphael Lekuraiyo who assisted me in the initial stages of the research; John Doherty, Pascal Fournie, Caroline Mullins, who provided additional lion information; and numerous guides who sent me a constant stream of “lion texts”- Bernard Lesirin, Serenoy Letoiye, Julius Lesooli, Sumoro Lecharkole, John, Peter (Joys Camp), Daniel Kasoo, Onesmas Lesiata; and the hospitable, helpful lodge managers of the area. The rangers and wardens of Samburu, Buffalo Springs and Shaba have been fantastic at providing me with information, assistance and support through the years – Mike v Lesiil, Boniface Lekartiwa, Ernest Ekeno, Abdi Sukuna, Jirmo, Rashid, Salat, Ibrahim (Rangers); James Lesuyai, Abdi Boru, Simon Leyarana, Moses Lolmodooni, Gabriel Lepariyo, Joy Letoiya (Wardens). The Samburu community have made me feel very welcome, at home and safe in Westgate; Stephen Lenantoiye, Daniel Letoiye - great friends who provide a fantastic support system together with all the community scouts. Tony and Ali Allport who set me up in Westgate, helped with the logistics and provided moral and emotional support when things got tough. Subsequent managers Doug and Tanya who have been continuously hospitable, helpful and supportive. Lion conservationists who have helped with so much - Alayne Cotterill, Siva Sundaresan, Zeke Davidson, Paul Funston, Leela Hazzah, Stephanie Dolrenry and Colleen Begg. Thank you for your support and friendship. My friends whose words of encouragement saw me through some dark times - Nas Merali, Pranj Acharya, Arzina Bhanjee, Shane Dallas, Deborah Rosario, Nilanga Jayasinghe and Chip Owen for his continued support and friendship. The Ewaso Lions Team - Laikos Letupukwa, Moses Letitiya, Francis Lendorop, Jeremiah Letoole, Yesalai Lemachokoti, Leshula Lenakae, Ngila Ltenesi, Ltalesi Lekuraiyo, Letoiye - you have worked hard and stayed by my side through the easy and hard times; all the warriors from “Warrior Watch” – a very special group of young men; Paul Thomson – my colleague; Jeneria Lekilelei – without whom none of this would be possible - battling cattle raids, droughts, escaping from poachers, digging out of floods, long days in the field, being robbed – always with a smile and positive outlook. He is my right-hand man, my strength. Kura – always with me. You are all my Samburu Family – and I respect and thank you for everything. Ashe Oleng. A special thanks to Irene Amoke, Heather Gurd, Charles Ngero, Mixie Wandera, Tobias Otieno and Nancy Khadioli for assistance during the re-writing of the thesis, and to my examiners Adrian and Stewart. My family in England - without your support, encouragement and endless patience, Ewaso Lions, this study and all that I strive for would not be possible. Kalpana and Shirish Jani – I would not have got through the final stages without you. My sister Shalini who provided precious practical as well as emotional support on my numerous trips to England – I couldn’t have done it without you. My parents Ashok and Sharmi Bhalla – who first introduced me to safaris in my childhood and created this lifelong passion and enthusiasm for wildlife