Cooperative breeding and anti-predator strategies of the azure-winged magpie (Cyanopica cyanus Pallas, 1776) in northern Mongolia Dissertation to acquire the doctoral degree in mathematics and natural science the degree 'Doctor of Philosophy' at the Georg-August-University Goettingen in the doctoral degree programme 'Biological Diversity and Ecology' at the Georg-August University School of Science (GAUSS) Submitted by Gantulga Bayandonoi from Dornogobi, Mongolia Goettingen, 2016 0 Thesis Committee Univ.-Prof. i.R. Dr. Michael Muehlenberg Honorary Professor Conservation Biology, Workgroup on Endangered Species, Department of Animal Ecology, Johann-Friedrich-Blumenbach Institute of Zoology & Anthropology, Georg-August University, Goettingen Prof. Dr. Peter M. Kappeler Department of Sociobiology/Anthropology, Johann-Friedrich-Blumenbach Institute of Zoology & Anthropology, Georg-August University, and Behavioural Ecology & Sociobiology Unit, German Primate Center (DPZ), Goettingen, Germany Prof. Dr. Eckhard W. Heymann Department of Sociobiology/Anthropology, Johann-Friedrich-Blumenbach Institute of Zoology & Anthropology, Georg-August University, and Behavioural Ecology & Sociobiology Unit, German Primate Center (DPZ), Goettingen, Germany Members of the examination board: Referee: Univ.-Prof. i.R. Dr. Michael Muehlenberg Co-referee: Prof. Dr. Peter M. Kappeler Second Co-referee: Prof. Dr. Eckhard W. Heymann Other members of the Examination Board: Prof. Dr. Stefan Scheu Department of Animal Ecology, Johann-Friedrich-Blumenbach-Institut für Zoologie und Anthropologie, Georg-August University, Goettingen Prof. Dr. Rainer Willmann Abteilung Morphologie, Systematik, Evolutionsbiologie Johann-Friedrich-Blumenbach-Institut für Zoologie und Anthropologie, Georg-August University, Goettingen PD. Dr. Matthias Waltert Workgroup on Endangered Species, Department of Animal Ecology, Johann-Friedrich- Blumenbach Institute of Zoology & Anthropology, Georg-August University, Goettingen Date of the oral examination: 11.07.2016 1 CONTENTS ACKNOWLEDGEMENTS ..................................................................................................... 5 ABBREVIATIONS .................................................................................................................. 6 CHAPTER 1 ............................................................................................................................. 8 General introduction ................................................................................................................ 8 Cooperative breeding ........................................................................................................... 10 Anti-predator strategies ........................................................................................................ 12 The study species ................................................................................................................. 14 Cooperative breeding in the azure-winged magpie .......................................................... 17 Nest predators of the azure-winged magpie ..................................................................... 18 Study colonies .................................................................................................................. 18 The outline of this study ....................................................................................................... 21 CHAPTER 2 ........................................................................................................................... 24 Helping behaviour affects nestling condition and nest predation in a facultative cooperative breeding bird, the azure-winged magpie ......................................................... 24 Abstract ................................................................................................................................ 25 Introduction .......................................................................................................................... 26 Methods ................................................................................................................................ 29 Study area and breeding colonies ..................................................................................... 29 Data collection .................................................................................................................. 30 Nest predators ................................................................................................................... 33 Data analyses .................................................................................................................... 35 Results .................................................................................................................................. 37 Food availability ............................................................................................................... 37 Feeding rate ...................................................................................................................... 37 Nestling body mass .......................................................................................................... 41 Number of helpers at the nest ........................................................................................... 43 Duration of the chick-rearing stage .................................................................................. 43 Nest predation .................................................................................................................. 43 2 Nest attentiveness ............................................................................................................. 44 Discussion ............................................................................................................................ 44 Food availability ............................................................................................................... 44 Contribution from helpers ................................................................................................ 45 Selective factors promoting cooperative breeding ........................................................... 47 Acknowledgements .............................................................................................................. 51 CHAPTER 3 ........................................................................................................................... 52 Responsive adaptation to anti-predator strategies of the Azure-winged magpie (Cyanopica cyanus Pallas, 1776) breeding colonies in northern Mongolia ....................... 52 Abstract ................................................................................................................................ 53 Introduction .......................................................................................................................... 54 Methods ................................................................................................................................ 57 Study area and breeding colonies ..................................................................................... 57 Nest predators ................................................................................................................... 57 Data collection .................................................................................................................. 58 Data analyses .................................................................................................................... 59 Results .................................................................................................................................. 62 Predation rate .................................................................................................................... 62 Nest distance .................................................................................................................... 63 Nest concealment ............................................................................................................. 64 Effect of breeding pair experience on anti-predator strategies......................................... 64 Avoidance from the carrion crow nest ............................................................................. 65 Synchronous breeding ...................................................................................................... 65 Impacts on nest success .................................................................................................... 65 Behavioural responses against predators .......................................................................... 68 Protector species ............................................................................................................... 69 Discussion ............................................................................................................................ 69 Nest concealment ............................................................................................................. 69 Effect of helpers ............................................................................................................... 70 Effect of breeding pair experience ................................................................................... 70 Synchronous breeding .....................................................................................................