THE MANAGEMENT OF LIONS (PANTHERA LEO) IN SMALL, FENCED WILDLIFE RESERVES A thesis submitted in fulfilment of the requirements for the degree of DOCTOR OF PHILOSOPHY of RHODES UNIVERSITY by Orla McEvoy March 2019 ABSTRACT Reintroduced lion (Panthera leo) populations pose several ecological and management challenges in small (< 1,000 km2), fenced wildlife reserves. Changes in the natural social- ecological conditions of reintroduced lion populations may lead to rapid population growth and a breakdown of natural predator-prey relationships. Reduced competition with other lions also likely reduces the potential for reintroduced lions to naturally form groups. My study used a combination of questionnaire surveys with tourists, existing lion demographic data from 16 wildlife reserves across South Africa and a controlled lion social experiment to address these ecological and management issues. Tourism was the primary reason for lion reintroductions. Tourists scored lions highly in terms of preference for viewing on safari, in particular, lions in larger, natural groups and adult males. Viewing lions also enhanced a tourists’ overall safari experience. The breakdown of natural social behaviour may likely therefore reduce tourist satisfaction related to lions. The number of resident prides and male coalitions in a reserve affected lion vital rates. Lion population growth rate was highest in reserves that contained a single resident pride, and the presence of unknown adult males significantly reduced cub survival and lioness birth intervals. The ratio of male cubs born also increased in reserves with a higher density of unknown adult males. Fertility control measures (deslorelin implants and unilateral hysterectomy) were effective at limiting lion population growth. Deslorelin treatment increased the age of first reproduction or the birth interval and decreased the subsequent litter size of treated lionesses to closer reflect natural vital rates in larger (> 10, 000 km2) systems. However, there was variability in infertility response between lionesses including adverse reactions in a small proportion of treated individuals. The number of resident prides and male coalitions in a reserve affected lion social behaviour. Lionesses formed larger groups in reserves with a higher density of unknown adult female neighbours, likely driven by territory defence. Lion prides with resident cubs were generally more fragmented, likely in response to reduced competition from unknown adult males. However, in areas with a high density of unknown adult female neighbours, prides with cubs formed larger groups likely in response to heightened territory defence. Therefore, with smaller foraging group sizes, predation rate was increased in reserves with reduced competition from unknown lions. My study supports a metapopulation approach for the management of lions in small, fenced reserves, and the standardisation of lion management procedures and database management. Endorsed by the Biodiversity Management Plan for lions in South Africa, this will enhance the long-term conservation potential of isolated populations. i ACKNOWLEDGEMENTS With utmost gratitude and appreciation, I would like to acknowledge the backing and invaluable contributions of a vast network of people, who provided integral support during the execution of this research. First and foremost, I would like to extend my sincerest gratitude towards my adept supervisors, Dr Dan Parker and Dr Sam Ferreira. My utmost thanks to Dan for the unfettered, friendly encouragement, skill and patience and over the past four years, particularly during the write-up phase. My utmost thanks to Sam for his crucial involvement in establishment of this work, and for his adept guidance throughout. They have each been an indispensable support to this research. I would next like to extend my greatest gratitude to all members of the Lion Management Forum of South Africa (LiMF) and remaining wildlife reserve staff, without whom, this research would not have been possible. Dr Susan Miller, the LiMF Secretary, requires special recognition for her continuous enthusiasm and skilled professionalism throughout this research, along with invaluable support and data contributions. Similarly, Brian Courteney, another LiMF member and founder of Southern African Conservation Trust, has provided integral support to this research, through continued friendly support, funding provisions and the provision of a vital research vehicle for the field work phase. I would also like to extend my greatest gratitude to the National Research Foundation for their significant contribution as a chief sponsor of this research. I would also like acknowledge SANParks administration and staff at participating national parks for their support throughout. Other LiMF members and contributions otherwise that require special recognition for their invaluable supporting role in this research include the following: - John Adendorff, Roxanne Erusan, Angela Gaylard, David Zimmerman and other SANParks staff (past and present) for their support from Frontier region and the staff at Gorah Elephant Camp. - Mark and Amy Palmer, Natalie Fowler and the staff at Amakhala Game Reserve. - Tim Parker and Ouida Van de Venter for their close-to-home support at Blue Canyon Conservancy, and the management and guiding teams at Moditlo River Lodge and Vuyani Safari Lodge therein. - Jono Berry, Brendan and Stevie Strydon, the volunteer teams and the support staff at Gondwana Game Reserve. - Hanno Killian at Khamab Kalahari Reserve. ii - Sabine Trollop at Entabeni Game Reserve. - Audrey Delsink Kettles, Ross Kettles and Susan Miller for their support regarding the Greater Makalali Private Game Reserve. - Jason Turner and the Global White Lion Protection Trust. - Mike Fuller, Jason Loest, Jarrett Joubert, Chris Reynecke and the volunteer teams at Kariega Game Reserve. - Colin Rowles at the Klaserie Private Nature reserve and the guiding and management teams at Ivory Wilderness, Klaserie Drift and Africa on Foot therein. - Angus Sholto-Douglas, Ina Fölscher and the guiding team at Kwandwe Game Reserve. - Piet Nel and Carlien Esterhuizen for their assistance at Madikwe Game Reserve. - Simon Naylor, Cilla Pickering along with the support staff and volunteer teams at Munyawana Private Game Reserve. - Mike Potter and Brett Deetlefs at Nambiti Game Reserve. - Ulrich Schutte, Christine van der Walt, Reinhardt and the team at Nkomazi Game Reserve. - Liesl Eichenberger and Paul Vorster and supporting staff at Sanbona Wildlife Reserve. - Dusty and Lourette Joubert along with Elizabeth Spencer and supporting staff at Selati Game Reserve. - Tinus Nel and Jacques Blignault, and the guiding team at Shambala Private Game Reserve. - John O Brien, Anja Truter and Alexandra Budd at Shamwari Private Game Reserve, and the guiding and volunteer teams therein. - Cathariné Hanekom and the supporting staff at Tembe Elephant Park. - Warren Beats, Mariana Venter, Alison Squance and the team at Thanda Private Game Reserve. - Bryan Havemann and Almero Bosch at Timbavati Private Nature Reserve, along with the management and guiding teams at Motswari Private Game Reserve, Shindzela Tented Camp and Kings Camp therein. Many thanks also to Bryan and Almero for the loan of research equipment used during field work for the project. - Brigitte Church and Eduard Goosen at uMkhuze Game Reserve, and the supporting team therein. - Andre Burger at Welgevonden Game Reserve for his support, including with the occasional stranded researcher, and the support on staff on the reserve. - Karen Holmes and Dane Antrobus, along with the guiding and support teams at Manyoni Game Reserve. - The staff and volunteer teams with African Impact. iii - A special thanks also to Gerrie Camacho from the Mpumalanga Parks and Tourism Agency for his support. - Nerosha Govender and the iSimangaliso Wetland Park Authority and other authorities and relevant staff including the North West Parks and Tourism Board, Department of Economic Development, Environment and Tourism (Limpopo), Ezemvelo KZN Wildlife, the Department of Economic Development, Environmental Affairs and Tourism (Eastern Cape) and Cape Nature (Western Cape). - The management and support team at Hoedspruit East Gate airport, including Tersia van der Westhuizen, and all respondents to the tourist survey respondents. - Craig Packer, Natalia Borrego and the Lion Research Centre at the University of Minnesota for their support and contributions. - Rob Slotow, Henk Bertschinger and the many other affiliates who have provided experience and support in this research, and the many teams who provided integral assistance with the field work. Although the list is long, thank you sincerely again to all the above mentioned for your time and dedication, without your support this research would not have been possible. My sincerest gratitude to all the support staff and students at Rhodes University, including the Registrar, post-graduate funding and the Department of Zoology and Entomology for their continued support in making this research possible. I would also like to acknowledge the Southern African Wildlife Management Association, Endangered Wildlife Trust, the Symposium of Contemporary Conservation Practise and the African Lion Working Group for their support in the expansion of the knowledge-base and communication of this research. I would also like to extend my greatest appreciation to my extended family and friends for their support. My deepest gratitude to my parents, Margaret and