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Durham E-Theses The structure of dominance hierarchies in the paridae: consequences for foraging eciency and body condition Hay, Jacqueline Marie How to cite: Hay, Jacqueline Marie (2003) The structure of dominance hierarchies in the paridae: consequences for foraging eciency and body condition, Durham theses, Durham University. Available at Durham E-Theses Online: http://etheses.dur.ac.uk/4060/ Use policy The full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that: • a full bibliographic reference is made to the original source • a link is made to the metadata record in Durham E-Theses • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders. Please consult the full Durham E-Theses policy for further details. Academic Support Oce, Durham University, University Oce, Old Elvet, Durham DH1 3HP e-mail: [email protected] Tel: +44 0191 334 6107 http://etheses.dur.ac.uk 2 The Structure of Dominance Hierarchies in the Paridae: Consequences for Foraging Efficiency and Body Condition. Jacqueline Marie Hay 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. Presented in candidature for the degree of Doctor of Philosophy University of Durham School of Biological and Biomedical Sciences 2003 This thesis is dedicated to the memory of Prof. Peter R. Evans (1937-2001) For my Mam and Dad "Good parents give their children Roots and Wings....Roots to know where home is, wings to fly away and exercise what's been taught" -Jonas Salk- The Structure of Dominance Hierarchies in the Paridae: Consequences for Foraging Efficiency and Body Condition. Jacqueline Marie Hay PhD 2003 In species that form social foraging groups during all or part of their lifecycle, individuals are subject to costs in terms of a higher frequency of aggressive interactions and having to share resources with group members, which must be outweighed by the benefits of flocking for all individuals concerned if it is to remain an Evolutionary Stable Strategy. In many group living species, dominance hierarchies exist that reduce the need for repeated agonistic interactions between the same individuals, which use valuable energy and may result in injury. Dominance hierarchies allow the most dominant individuals priority of access to contested resources whilst subdominant individuals may suffer as a consequence. In this thesis, I examined how dominance hierarchies were structured in great tits, blue tits and coal tits and how dominance status affected an individual's foraging efficiency and body condition. The food handling times or vigilance levels of great tits and blue tits were not governed by dominance status but dominant coal tits were less vigilant than subdominant conspecifics thus probably allowing more time for foraging and other social activities i.e. mate finding and territory defence. Though subdominant great tits did not suffer costs in terms of food handling times, they were affected more by interference competition when foraging in mixed tit flocks compared to dominant great tits. Using ptilochronology as a measure of nutritional condition, the results from coal tits in this study contradicted those of previous studies in that dominant coal tits had slower rates of feather growth and produced feathers that were shorter and less dense than subdominant conspecifics, possibly a cost of their high dominance. Feather growth rate in great tits and blue tits was not related to dominance status. The immunocompetence of adult great tits or blue tits was not governed by dominance status nor did great tit or blue tit nestlings show any relationship between their place in the brood competitive hierarchy and immuocompetence. Declaration I declare that the material contained in this thesis has not been previously submitted for a degree in this or any other university. All the work recorded in this thesis is entirely my own, unless otherwise stated. Statement of Copyright © The copyright of this thesis rests with the author. No quotation from it should be published without prior consent and any information derived from it should be acknowledged. 11 Acknowledgements There are many people whom I have to thank for their help whilst I was in Durham, probably more than the average PhD student since I have had five supervisors and moved offices three times. Firstly, I wish to thank the late Professor Peter Evans who picked me up when things were falling apart and offered me the PhD on tits at West Hollinside. Peter was a constant source of ideas and enthusiasm for my work and was a great support in designing the study. Though Peter was fighting a terrible illness, he was always there to offer advice but remained in the background when necessary, allowing me complete control of my PhD. Next I would like to thank Dr Keith Hamer who "inherited" me following Peter's death and then five weeks later took up the offer of a job at Leeds University. Keith however, was true to his word and supported me during my writing up process even though he had many commitments at Leeds and was putting together a research team there. Since my project was funded by Durham University, Keith had little to gain by providing such help but has stuck with me throughout, for which I am indebted. Professor Brian Huntley became my official supervisor after Dr Tom Sherratt (who took over after Keith left Durham) departed for a new life in Canada. Brian was unlucky enough to take over as my supervisor just as I was planning on spending a vast amount of money on lab work and so, as the department funded me, that money came out of Brian's budget, for which I am very grateful to him. My many lab mates and friends over the years have been a tremendous support during my supervisor and lab swapping activities. Thank you to: Ruth Cox, Stuart Bearhop, Sue Lewis, Kathy Fletcher, Steve Oswald, Trish Ranner, Judy Allen, Catherine Gray, Rich Fuller, Claire Flockhart and Hannah Drewitt. I would like to extend special thanks to Steve Willis who was always willing to iii drop everything and lend a statistical ear. For reading chapter drafts and providing helpful comments, I am grateful to Matt Davey, Claire McSorely and Veronica Mands. I am also grateful to Dr Bob Baxter for allowing me use of his wet lab during my parasitology work. During my plunge into molecular biology, Debs Dawson and Andy Krupa at the Sheffield Molecular Genetics Facility and Becky, Matt and Alex in lab 10 at Durham provided me with enormous support. Thank you also to Dr Stefan Przyborski for allowing me to use his facilities at Durham. Michael Bone deserves a special mention for building my bird tables, nest boxes and roosting box. Thank you also goes to Michael for spending a summer either climbing my ladders for me or having to cope with the embarrassment of supporting my trembling bottom as I tried to battle with vertigo when only a few steps up the ladder. Thank you also to Kim Leyland and Catherine Bennison who helped collect seed handling and vigilance data. My PhD would not have been nearly as easy or enjoyable if it had not been for my soon-to-be husband, Robin Ward. Robin faithfully gave up most of his weekends and many weekday mornings during my fieldwork to help me mist net before I gained my C permit (though he did steal all the good birds). He also spent much time checking nest boxes, feeding the birds and keeping me company during observations. Since Robin knows almost everyone who works on birds, he was a constant source of contacts and information during writing up and though he had left Durham by then, was always on the end of the phone day and night, providing encouragement and support. Finally, I would like to thank my family for their love, support and encouragement over the years. My parents' belief in me has been unfailing throughout my education and they have always encouraged me to put in 100% effort. My Dad slaved away building potter traps for my work and Mam had to sacrifice her bathroom renovations to allow him the time to do it. I cannot thank them enough for all they've done and hope that I can repay them one day. iv TABLE OF CONTENTS Abstract i Declaration ii Acknowledgements Hi Table of Contents v List of Tables xiii List of Figures and Plates xvi CHAPTER 1: GENERAL INTRODUCTION 1 1.1 General introduction 1 1.2 Study species 2 1.2.1 The Paridae 2 1.2.2 Great tit Parus major 3 1.2.3 Blue tit Parus caeruleus 4 1.2.4 Coal tit Parus ater 5 1.3 The formation of foraging flocks 6 1.3.1 Benefits of joining foraging flocks 7 1.3.2 Costs of joining foraging flocks 8 1.4 Dominance hierarchies within flocks 9 1.4.1 Benefits of high dominance status 9 1.4.2 Apparent costs of high dominance status 11 1.5 Aims and objectives 12 CHAPTER 2: STUDY SITE AND METHODOLOGY 14 2.1 Study site 14 2.1.1 General description 14 2.1.2 West area 15 2.1.3 East area 15 2.1.4 Use of study site 19 2.2 Nest boxes 19 v 2.3 Feeding regime 19 2.4 Bird capture 20 2.4.1 Capture of feeding birds 21 2.4.2 Capture of roosting birds 22 2.4.3 Capture of breeding adults at nest boxes 22 2.4.4.