The ecology of cheetahs and other large carnivores in a pastoralist-dominated buffer zone By Thomas M. Maddox Ph.D. Thesis Department of Anthropology, University College, London & Institute of Zoology, London 2003 “No wild animals are really bad as you will be alone without them. It is good just to see and be happy” (Iltiyogoni laigwanan, Loliondo). 2 Summary Due to the various limitations of core-protected areas, interest in semi-protected landscapes and the human-wildlife interactions that occur within them is rapidly gaining credence. Some of the most important issues within this field are human- carnivore relationships, with many large carnivores globally threatened on one hand but with the capability of potentially devastating impacts on humans on the other. In this thesis, the success of cheetahs, their competitors and their predators in two buffer zones (Loliondo and Ngorongoro) of the Serengeti National Park in Tanzania was examined in comparison with populations living inside the park. The potential role played in carnivore ecology by the Maasai pastoralists inhabiting the buffer zones was then examined to assess the extent to which their presence determines any of the differences. The results show that large carnivores and their prey are surviving successfully outside the core-protected area, coexisting with the pastoralist Maasai. Herbivores were shown to exist at equivalent diversity and density outside the park, with the two- 2 year average prey biomass significantly higher in Loliondo than inside the park (χ 2= 49, p<0.001). The only species consistently more abundant inside the park were kongoni, topi and warthog. However, temporal variation was large and the system was better described as a single, dynamic entity rather than three distinct and comparable sites. Study sites outside the park also held substantial populations of all large carnivore species. Densities of both common jackal species were higher outside the park, hyaena estimates were higher inside the park and there was no significant 2 difference between lion estimates (χ 2=0.4, NS). Lion density in Loliondo was estimated at 0.37 lions / km2, a density comparable with most protected areas. Cheetah data were limited but showed a substantial population outside the park. Several carnivore estimation methods were used in the study, and comparison of the results showed that the visual-based surveys commonly used elsewhere (line transects or driven indices) were highly limited outside protected areas. Little behavioural variation was shown in cheetahs between individuals inside and outside the park (effect of region on time spent relaxed: F5,65=0.09, NS). Both cheetahs and lions showed strong reactions to playbacks of Maasai cattle, however responses were mixed 3 with only lions outside the park showing a consistent increase in vigilance (T9 = - 2.72, p<0.05). The role of Maasai was investigated through questionnaires. Their answers showed large carnivores to be a major part of their environmental perceptions, with lions, hyaenas and leopards receiving the top salience scores, although cheetahs were not due to a lack of differentiation between the spotted cats. General attitudes were positive, particularly in Ngorongoro, but attitudes towards large carnivores were mostly negative. Costs of coexistence were significant for both sides; livestock predation was experienced by 89% of Loliondo respondents and 63% of Ngorongoro respondents, with predators accounting for 1% of cattle herds and 3% of sheep and goats, although disease accounted for far higher. Human injuries were also reported, although most (70% in Loliondo and 50% in Ngorongoro) occurred through lion hunts. The lion hunt is still an important part of Maasai culture, with over 75% of respondents reporting having attended at least one. Based on average estimates from respondents and age group leaders, 30-40 lions are thought to be ritually killed in the entire Maasai area (including Kenya) each year. However, although the relationship between the Maasai and carnivores is far from harmonious, coexistence is continuing thanks to semi-tolerant attitudes, restrictions on hunting impacts and preventative livestock management systems. 4 Declaration The material contained in this thesis has not previously been submitted for a degree at University College or any other university. The research reported within this thesis has been conducted by the author unless otherwise indicated. © The copyright of this thesis rests with the author. No quotation from it should be published without his prior written consent and information derived from it should be acknowledged. 5 Acknowledgements Firstly, I would like to acknowledge and thank my three supervisors for their guidance and supervision: Sarah Durant at the Institute of Zoology, London, Katherine Homewood at University College, London and Guy Cowlishaw, also of the Institute of Zoology, London. I would particularly like to acknowledge Sarah for initiating this study, guiding me in its development both in Tanzania and in the UK, teaching me all of the ecological field methods used in this thesis, and as my primary mentor for most of the analysis techniques used, as well as last minute chapter edits and pulling me out of mud holes. I would like to acknowledge Katherine for introducing me to the importance of the human dimension in ecology, for valuable guidance in the UK throughout the thesis and for helping me see the much broader picture and I would like to acknowledge and thank Guy for countless draft readings, analytical advice and support in the UK. Secondly I would like to give heartfelt thanks and acknowledgement to Dawn Scott for tireless support, discussion and analytical help throughout the thesis but especially in the final year of the study; my sincerest gratitude. Thirdly I would like to acknowledge and thank the various organisations that made the project possible; to the Natural Environment Research Council for funding the work, to the Tanzanian Wildlife Research Institute, Tanzanian National Parks, the Ngorongoro Conservation Authority the Wildlife Division and the Council for Science and Technology in Tanzania for allowing me permission to work in their beautiful country and to the Frankfurt Zoological Society, World Conservation Society and National Geographic for financial and logistical support for the Serengeti Cheetah Project. I would like also like to thank the people of Loliondo and Ngorongoro, in particular Jacob, Kaleya and their families, for welcoming and guiding me through some of the most fascinating and enjoyable days of my life I would also like to extend many thanks to Karyl Whitman for teaching me the art of call-ins and hiding me from hunters, colleagues at the Institute of Zoology, especially Marcus Rowcliffe and Richard Pettifor for valuable statistical advice, to Sarah 6 Randall and the members of HERG at UCL for advice and discussion on the social aspects of the work, to Andy Russell and Jon Barnard at the University of Cambridge for help with GLMs and Psion programming, to Jo and Judith for interspersions of sanity, to John McCabe, Laura de Luca, Craig Packer, Marion East, John Shemkunde, Simon Mduma and Tony Sinclair for their helpful discussions and to my family for their support and help spotting cheetahs and chasing baboons. 7 Contents SUMMARY.............................................................................................................................................3 DECLARATION ....................................................................................................................................5 ACKNOWLEDGEMENTS ...................................................................................................................6 CONTENTS............................................................................................................................................8 CHAPTER 1: INTRODUCTION....................................................................................................14 1.1 SUMMARY ............................................................................................................................14 1.2 THE PROTECTED AREA SYSTEM ............................................................................................14 History and definitions of protected areas....................................................................................14 Importance of core-protected areas..............................................................................................15 Limitations of core-protected areas..............................................................................................16 Importance of semi-protected areas and buffer zones ..................................................................18 Protected areas in Tanzania .........................................................................................................22 1.3 HUMAN-WILDLIFE CONFLICT WITHIN BUFFER ZONES ...........................................................23 Effects of humans on wildlife ........................................................................................................23 Hunting .................................................................................................................................................... 23 Habitat change.......................................................................................................................................... 24 Effects of wildlife on humans ........................................................................................................25