To the University of Wyoming: The members of the Committee approve the dissertation of Abdullahi H. Ali presented on 4/13/2016. Jacob R. Goheen, Chairperson Jeffrey L. Beck, External Department Member Matthew J. Kauffman Anna D. Chalfoun David McDonald APPROVED: Dr. Bob Hall: Director, Program In Ecology. Paula Lutz: College Dean Ali, Abdullahi H., Range Collapse, Demography and Conservation of the Critically Endangered Hirola Antelope in Kenya., Ph.D., Program in Ecology, May, 2016. The hirola (Beatragus hunteri) is one of the rarest antelopes on Earth, with a global population size of ca. 500 individuals restricted to 1500 km2 on the Kenya-Somalia border. Hirola has exhibited ongoing declines since the 1970s while the remaining populations occur almost solely on pastoral lands with no formal protection. Because of historical and political instability in the hirola’s native range, it has been difficult to clearly identify the reasons underlying hirola declines. Like many other globally endangered species, it is likely that more than one factor underlies the hirola problem. Therefore, I investigated, 1) the role of predation and range degradation in driving hirola declines, 2) mechanisms responsible for hirola range collapse and landscape change within hirola historical range, and 3) identified socially- acceptable strategies for habitat restoration and hirola recovery. I have used a combination of GPS telemetry, analysis of long-term satellite imagery, a large-scale predator exclusion zone, and structured questionnaires to address my objectives. Understanding the impacts of these factors can help determine the next steps necessary to maximize the chances of hirola persistence in the future. My findings suggest that, a combination of top-down (predation) and bottom-up (rangeland quality) forces drive hirola declines, with populations in the historical range being impacted more by poor rangeland quality. Additionally, resource selection analysis revealed that contemporary low numbers of hirola are due to loss of forage via tree encroachment. Some factors—including mega-herbivore extirpations, fire suppression, and overgrazing—were likely to have triggered tree encroachment which may be slowing contemporary recovery efforts. Local communities supported efforts to conserve elephants, 1 seed and fertilization of grass, and removal of trees as means to restore hirola historical range. However, the locals were opposed to voluntary reduction of livestock and were ambivalent towards soil ripping and control burns. Livestock wealth (ownership) and years of residency were important predictors of locals’ perceptions toward rangeland-restoration practices. I recommend a combination of rangelands restoration efforts that have local support coupled with reintroductions to enhance the chances of recovery for this globally endangered species. 2 RANGE COLLAPSE, DEMOGRAPHY AND CONSERVATION OF THE CRITICALLY ENDANGERED HIROLA ANTELOPE IN KENYA By Abdullahi H. Ali A dissertation submitted to the Department of Zoology and Physiology and the University of Wyoming in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY in ECOLOGY Laramie, Wyoming May 2016 Copyright © 2016 by Abdullahi H. Ali ii This dissertation is solely dedicated to my late brother Muhumed H. Ali who lost his life in a surgical theatre room at the Garissa General hospital (Kenya) in October 2004. May God rest his soul in eternal peace. Amen. iii Acknowledgements The financial support of this work was made possible by the generosity of the Kenya Wildlife Service, local communities in Fafi and Ijara sub-counties, Garissa County government, Ishaqbini Community Conservancy (Kenya), Northern Rangelands Trust (Kenya), Association of Zoos and Aquariums (USA), British Ecological Society (UK), Chicago Zoological Society (USA), Denver Zoo (USA), Disney Worldwide Conservation Fund (USA), Houston Zoo (USA), Idea Wild (USA), International Foundation for Science (Sweden), IUCN/SSC Antelope Specialist Group, Mohamed Bin Zayed Species Conservation Fund (UAE), National Museums of Kenya, the People’s Trust for Endangered Species (UK), Rufford Foundation (UK), St. Louis Zoo’s Center for Conservation in the Horn of Africa (USA), the University of Wyoming’s Berry Biodiversity Conservation Center, the University of Wyoming’s Haub School, and the Zoological Society of London. I wish to thank all the individuals from these organizations for their foresight and generosity. Special thanks go to Drs, Charles Musyoki, David Mallon, and Martha Fischer for their continued enthusiasm and encouragement with this research. I also thank my field collaborators; Amos Muthee, Ian Craig, Idle Farah, Isaac Lekolol, Juliet King, Mathew Mutinda, and Osman Abdi for assisting in my field work. Further, distinguished thanks go to my academic committee members: Dr. Anna Chalfoun, Dr. David McDonald, Dr. Jacob Goheen, Dr. Jeffrey Beck, and Dr. Matthew Kauffman for their continued support throughout my study period. In particular, I would like to acknowledge and extend special thanks to my advisor: Dr. Jacob R. Goheen for his guidance and dedicated efforts throughout the study period. Special appreciations also go to Dr. Matthew Kauffman, for his dedicated support during the last phase of my PhD project. I wish to also acknowledge several mentors and collaborators who iv also contributed to this research including: Dr. Adam Ford, Dr. Jefferey Evans, Mr. Matthew Hayes, Dr. Rajan Amin, Dr. Richard Muita and Dr. Samuel Andanje for their invaluable inputs and support throughout the study period. Without forgetting the faculty and students from the Department of Zoology and Physiology and Program In Ecology have all been very helpful with ideas and encouragements throughout the study period. I wish to say Thank You. Specifically, I wish to thank Program in Ecology and the Department of Zoology and Physiology administrative staff including Brenna Marcisek, Carol Pribyl, Sheila Krupp and Shawn Sheen for their help with all the logistics during the study period. Finally, I would like to thank all of my family members: My wife Zamzam and children Mohamed, Tahmin, and Tasneem for their spiritual and emotional support during my study and stay away from home. v Table of Contents Chapter 1 ................................................................................................................................... 1 1.1 Introduction ....................................................................................................................... 1 1.2 Materials and Methods ...................................................................................................... 3 1.2.1 Study area ................................................................................................................... 3 1.2.2 Demographic field study design ................................................................................. 4 1.2.3 Demographic data collection and analysis ................................................................. 6 1.3 Results ............................................................................................................................... 8 1.4 Discussion ......................................................................................................................... 9 1.6 References ....................................................................................................................... 11 Tables .................................................................................................................................... 16 Table 1: Mean difference in survival rates, adult fecundity and sensitivities for age classes between pairs of settings, from 2012 – 2015 study period. .................................. 16 Table 2: Elasticity estimates for calves, sub-adults and adults. ........................................ 17 Figures .................................................................................................................................. 18 Figure 1: (A) Map of Kenya and the study area in Garissa County, Kenya and (B) Estimated hirola population trends from 1977 – 2011, Data courtesy of the Kenya Department of Resource Surveys and Remote Sensing. ................................................... 18 Figure 2: Mean survival rates (± 95% CI) for hirola in the sanctuary, conservancy and Ijara rangelands for (A) adult, (B) sub-adult and (C) calf on the 2012 – 2015 study period. ................................................................................................................................ 19 Figure 3: Long term growth rates (λ; ± 95% CI) of hirola in the sanctuary, conservancy, and the Ijara rangelands based on the 2012 – 2015 study period. ..................................... 20 vi Figure 4: Results from a life table response experiment (LTRE) indicating the sensitivity and percent contributions of vital rates for 2012 – 2015 to the population growth changes in hirola for paired settings A) sanctuary vs conservancy, B) sanctuary vs outlying areas (Ijara rangelands) and C) conservancy vs outlying areas (Ijara rangelands). ................... 21 Chapter 2 ................................................................................................................................. 22 2.1 Introduction ..................................................................................................................... 22 2.2 Methods .......................................................................................................................... 24 2.2.1 Study area ................................................................................................................. 24 2.2.2 Data collection and analysis ..................................................................................... 25 2.3 Results ............................................................................................................................. 32 2.4 Discussion ....................................................................................................................... 32 2.6 References ....................................................................................................................... 35 Tables .................................................................................................................................... 40 Table 1: Summary of hirola home range sizes and the proportions of tree cover within home ranges during both dry and wet season. .................................................................. 40 Figures .................................................................................................................................. 41 Figure 1: (A) Study site in Garissa County, Kenya and the historical geographic range of hirola estimated from a minimum convex polygon based on the distribution of hirola in 1963. (B) Tree cover across the hirola’s historical range (1985). (C) Tree cover across the hirola’s current range (2012). Dark gray represents tree cover and light gray represents grasslands. The linear feature at the west of both images is the Tana River. Tree cover increased 251% between 1985 and 2012. ......................................................................... 41 vii