Behavioral ecology of the African wolf (Canis lupaster) and its implication for Ethiopian wolf (Canis simensis ) conservation in the Ethiopian Highlands Tariku Mekonnen Gutema Dissertation presented for the degree of Philosophiae Doctor (PhD) 2020 Center for Ecological and Evolutionary Synthesis Department of Biosciences Faculty of Mathematics and Natural Sciences University of Oslo © Tariku Mekonnen Gutema, 2020 Series of dissertations submitted to the Faculty of Mathematics and Natural Sciences, University of Oslo No. 2298 ISSN 1501-7710 All rights reserved. No part of this publication may be reproduced or transmitted, in any form or by any means, without permission. Cover: Hanne Baadsgaard Utigard. Print production: Reprosentralen, University of Oslo. Acknowledgments First and foremost, I would like to thank almighty God for giving me health, strength and courage to complete this thesis. This work would not have been possible without the advice and support of many people. Firstly, I would like to acknowledge my principal supervisor Prof. Nils Chr. Stenseth, University of Oslo, for accepting me as a PhD student and for his continued advice, encouragement and unlimited support during the study. He offered me a full academic freedom and flexibility besides of his scientific guidance. I am very grateful to my three co-advisors, Prof Afework Bekele and Dr. Anagaw Atikem, Addis Ababa University, and Prof Claudio Sillero-Zubiri, University of Oxford, for their advice, comments and guidance throughout the study period. I would like to thank Dr. Anagaw Atickem for introducing me to the importance of the study of the recently discovered African wolf ecology and for his support from designing of the study to the write up of the thesis. I thank all my field assistants: Getabalew Asefa, Abebe Gosem, Demeke Fantabel and Fantaw Geto. They worked so hard through all of the harsh weather in the Guassa Community Conservation Areas and Borena Saynt National Park to make this project successful. I thank all those who helped with the capture and radio-telemetry of African wolves, in particular, Alo Hussein, Kassim Kedir and Dr. Alemayehu Lema for capture, immobilization and collaring. I am deeply indebted to the Ethiopian Wolf Conservation Programme (EWCP) for providing me with the capture and immobilization equipment and expertise. I also thank the Ethiopian Wildlife Conservation Authority for providing permission for this study and to capture and collar African wolves. I would also like to acknowledge Prof. Peter J. Fashing, Dr. Vivek V. Venkataraman, Dr. Diress Tsegaye, Dr. Dessalegn Chala, Dr. Jorgelina Marino and Dr. Dietmar Zinner for their valuable comments on this thesis. My sincere appreciation is also extended to many friends including Dr. Addisu Mekonnen, Behailu Etana, Dr. Berhun Gebremedhin, Dereje Tesfaye, Ejigu Alemayehu and Misganu Tamirat. This work would not have been possible without the financial support of the Centre for Ecological and Evolutionary Synthesis (CEES) at the University of Oslo, the Norwegian Quota Scheme, Rufford Small Grants Foundation, Mohamed Bin Zayed Species Conservation Fund and Jimma University. I would like to extend my heart-felt gratitude to all of them. i Finally, I owe thanks to a very special person in my life, my wife Tejitu Shiferaw, for her continued and unfailing love, support and understanding during my pursuit of this PhD degree, which made the completion of the thesis possible. She has always been on my side being the base of my career and taking care for our three beloved children, Duran, Berit and Eban. Duran, Berit and Eban – many thanks and my heartfelt gratitude for your understanding, support and love. ii Contents Acknowledgments… ................................................................................................................................................ i Contents… ............................................................................................................................................................ iii Abstract .................................................................................................................................................................. vi List of papers ......................................................................................................................................................... vi 1. Introduction ....................................................................................................................................................... 1 1.1 History and distribution of the African wolf .................................................................................................. 1 1.2 Distribution and threats of Ethiopian wolves ................................................................................................. 2 1.3 Competition among mammalian carnivores ................................................................................................ 3 1.4 The effect of competition on carnivores ........................................................................................................ 4 1.5 Resource use and coexistence of sympatric carnivores .................................................................................. 6 1.6 Anthropogenic impacts on carnivores .......................................................................................................... 6 2. Aim of the study ............................................................................................................................................... 9 3. Main Methods ................................................................................................................................................. 10 3.1 Study area ................................................................................................................................................... 10 3.2 Material and methods .................................................................................................................................. 11 3.2.1 Capture and immobilization of African wolves ........................................................................... 11 3.2.2 Competition between African and Ethiopian wolves ................................................................... 12 3.2.3 Foraging ecology of African wolves ........................................................................................... 13 3.2.4 Ranging, habitat, and activity patterns of African wolves ............................................................ 15 3.2.5 African wolf predation on livestock and associated conflict ......................................................... 15 4. Results and discussion ..................................................................................................................................... 17 4.1 Competition between African and Ethiopian wolves ................................................................................... 17 4. 2 Foraging ecology of African wolves ........................................................................................................... 17 4.3 Ranging, habitat, and activity patterns of African wolves ............................................................................ 18 4.4 African wolf predation on livestock and ensuing conflict............................................................................. 19 4.5 Appendix: Capture and immobilization of African wolves .......................................................................... 19 5. Conclusions and future recommendations ....................................................................................................... 21 6. Project in progress ........................................................................................................................................... 22 6.1 Population status and reproduction ecology of African wolves....................................................................... 22 6.2 Promoting the Co-Existence Approaches to Carnivore Conservation .......................................................... 22 7. References ....................................................................................................................................................... 23 iii Abstract Recent rapid increases in human populations are leading to conversion of wildlife habitats into agricultural areas and human settlements. This is forcing wildlife populations to increasingly concentrate into confined protected areas, which in turn increases competition between wildlife species for limited space and food. African wolves (Canis lupaster) and Endangered Ethiopian wolves (C. simensis) are sympatric in parts of the Ethiopian Highlands. The Ethiopian wolf is an ecological specialist and only small populations remaining which makes it sensitive to both exploitative and interference competition from other carnivores. To better understand these dynamics this thesis focuses on the behavioral ecology of the African wolf and its impact on Ethiopian wolves and their conservation. Due to their perceived impact as predators on livestock, I also studied the extent of human-carnivore conflict. The study was carried out from March 2015 to June 2017 in Guassa Menz Community Conservation Area and Borena Saynt Worehimenu National Park in the central Ethiopian Highlands. Focal watches of 14 colalred African wolves were carried out to study their spatial and foraging ecology, and habitat