Impacts of Capture and Handling on Wild Birds
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CARDIFF UNIVERSITY SCHOOL OF BIOSCIENCES ORGANISMS AND ENVIRONMENT IMPACTS OF CAPTURE AND HANDLING ON WILD BIRDS Leila Magna Gomes DUARTE SUPERVISORS: Dr R. Thomas, Dr I. Vaughan, Dr N. Marples and Dr J. Ramos Thesis submitted to Cardiff University, UK in partial fulfillment for the degree of Doctor of Philosophy Cardiff, September 2013 I Declaration This work has not previously been accepted in substance for any degree and is not concurrently submitted in candidature for any degree. Signed ………………………………………… (candidate) Date …………………… STATEMENT 1 This thesis is being submitted in partial fulfillment of the requirements for the degree of PhD. Signed ………………………………………… (candidate) Date …………………… STATEMENT 2 This thesis is the result of my own independent work/investigation, except where otherwise stated. Other sources are acknowledged by explicit references. Signed ………………………………………… (candidate) Date …………………… STATEMENT 3 I hereby give consent for my thesis, if accepted, to be available for photocopying and for inter-library loan, and for the title and summary to be made available to outside organisations. Signed ………………………………………… (candidate) Date …………………… STATEMENT 4: PREVIOUSLY APPROVED BAR ON ACCESS I hereby give consent for my thesis, if accepted, to be available for photocopying and for inter-library loans after expiry of a bar on access previously approved by the Graduate Development Committee. Signed ………………………………………… (candidate) Date …………………… II III Acknowledgments The doctoral journey has been a challenge in many ways, and it would not have been possible to complete it without the help and support of the people around me, to only some of whom it is possible to give particular mention here. Firstly I wish to acknowledge Fundação para a Ciência e a Tecnologia (FCT) in Portugal for supporting this project through a Ph.D. Grant (reference SFRH/BD/61404/2009/J520351426XS). I want to specially express my gratitude to my main supervisor Dr. Rob Thomas, for proposing this project to me, for trusting me, for his support and the inestimable effort he has put into guiding me through this doctoral project. It would have not been possible to complete this thesis withouth his invaluable input. Additionally I wish to thank Dr. Ian Vaughan for his valuable comments and thesis corrections; to Dr. Jaime Ramos for his promptness, availability, practicality, support, important suggestions and guidance, without him a considerable part of my data collection would have been completely jeopardized; to Dr. Peter Ferns and his insight, advice and guidance; and to Dr. Nicola Marples for her knowledge and advice during the doctoral project. I wish also to acknowledge Dr Kate Buchanan for her advice and guidance relating to hormonal sampling in wild birds. The data presented in this thesis would have not been possible to collect without the collaboration of bird ringers, some of which preferred to remain anonymous. I particularly acknowledge the help and standardized data collection of the team of A Rocha Field Centre and all of their support and friendship, either during the regular bird ringing sessions or the “Stormies” season. I specially acknowledge Guillaume Rethoré, whose help and commitment to the project was precious. Thank you very much Gui! A good part of the data presented in this chapter would have not been possible to collect without the help and support of Helder Cardoso, an enthusiastic bird ringer and a beloved husband, whose unconditional love, patience and knowledge have supported me to go through this project. I also wish to acknowledge Pedro Andrade for his enthusiasm, for his interest in the project, for allowing me to collect data with him during his Master’s field project, and for disseminating the “questionnaire survey” on Facebook. Next I wish to acknowledge all those who were positively interested in the project and have punctually helped me either with data, facilities for data collection, their insights and/ or their acceptance, namely: Pedro Geraldes, José Alves, LabOR (Laboratório de IV Ornitologia da Universidade de Évora), Pedro Miguel Araújo, the “Stormies” volunteers of 2011, 2012 and 2013, Jenny Shepperson, Grace Emeny, Alice Ashbury, Xavier Schmidt, the participants in the questionnaire survey, and all of those who have in any constructive manner participated in the project. Last but not least, I wish to acknowledge my mother and my brother for all of their support and their pride in me; Renata Medeiros for all of her remarkable support, for facilitating my establishment in Cardiff and talking me into the PhD; Pierfrancesco, for believing in me, being my comrade in the stressful events and a reliable friend in the remaining ones; David Brown, Mafalda Costa and all of those who have “carried” me in any manner throughout the PhD. V Summary of the Thesis Bird ringing is a key ecological research technique that involves the capture and handling of birds. It is used extensively to obtain information on population dynamics of wild birds, and many aspects of avian behavior, physiology and life-history, which would otherwise be unfeasible to obtain. Despite millions of birds ringed every year, little is known about the short- or long-term impacts on birds, and whether there are negative welfare, conservation and scientific consequences, which can ultimately bias the interpretation of data from wild bird studies. In this thesis I study the type of intrusion that capture and handling causes to the bird, by analyzing their interlinked physiological and behavioral responses to capture stress, including hormonal and immunological responses, energy regulation decisions (feeding behavior and thermal regulation) and breeding effort. I further study the immediate effects that capture and handling has on birds through analyzing types of injuries and the rates at which injuries and mortality occur. I have focused mainly on mist-netting, which is the most widely used capture technique, and captures of passerine birds, which is the most frequently sampled taxon. However, I also demonstrate how the impacts of capture and handling can be studied in marine birds and applied to other capture methods. These studies reveal the range of short-term impacts that capture and handling may have on wild birds, and highlight aspects of methodology that have a strong effect on these impacts. The longer term consequences for lifetime fitness and demographic change require further study. This thesis demonstrated the importance for researchers to be aware of the potential effects of their activities on their study subjects, particularly for susceptible species and situations, and to continuously reasses their methods for effective improvement. I propose several guidelines, which aim to promote the birds’ welfare in regards to data collection. VI List of Contents Declaration II Acknowledgments IV Summary of the Thesis VI List of Contents VII List of Tables X List of Figures XIII Chapter 1 Introduction 1 1.1 Relevance of the topic 1 1.2 What is stress and how do animals respond to stress? 7 1.2.1 The role of the endocrine system in the adaptive stress response 13 1.2.2 Consequences for the immune system 18 1.3 Consequences for energy regulation 19 1.3.1 Changes in body mass 19 1.3.2 Changes in thermal regulation 20 1.4 Capture myopathy 21 1.5 Demographic impacts: breeding success 23 1.6 Physical integrity 28 1.6.1 Direct mortality 29 1.6.2 Physical injury 32 1.7 Thesis outline 33 Chapter 2 A time budget for mist netting and bird ringing, and the immediate responses of birds to capture and handling 36 2.1 Abstract 36 2.2 Introduction 36 2.3 Methods 41 2.4 Results 45 Hypothesis 1: Different stages of the ringing process differ substantially in their contribution to the overall time that a bird is retained. 46 Hypothesis 2: Extraction from the net and handling times are influenced by the experience of the ringer. 47 Hypothesis 3: Extraction from the net and handling times vary among species. 49 Hypothesis 4: The body grasp technique accelerates the extraction stage. 52 Hypothesis 5: The length of the ringing process influences the weight loss of the birds. 52 Hypothesis 6: Bird ringing imparts extra energetic costs to breeding females. 53 VII Hypothesis 7: Each species has a typical coping response. 54 Hypothesis 8: Coping responses are individually constant throughout the ringing procedure. 57 Hypothesis 9: The utterance of calls is related to both species, and coping response. 57 2.5 Discussion 59 Chapter 3 Body mass regulation in response to capture and handling in European passerines 64 3.1 Abstract 64 3.2 Introduction 64 3.3 Methods 67 3.4 Results 70 3.4.1 Feeder attendance 70 3.4.2 Mass regulation 73 3.5 Discussion 75 Chapter 4 Overnight thermal responses of migrating passerines in response to capture and handling 78 4.1 Abstract 78 4.2 Introduction 79 4.2.1 Bird’s body temperature 79 4.2.2 Defining normothermia 80 4.2.3 Distinction between hypothermia and torpor 80 4.2.4 Which birds resort to hypothermia and why? 81 4.2.5 Capture and handling effects on thermal regulation 83 4.2.6 Stress hormones and temperature regulation 84 4.2.7 Heart rate alterations as an effect of capture and handling 85 4.2.8 Introduction to the work developed in this chapter 85 4.3 Methods 87 4.4 Results 93 4.4.1 Barn swallow case study 93 4.4.2 Yellow wagtail’s case study 99 4.5 Discussion 103 Chapter 5 Interruption to the incubation routines of nesting passerines 109 5.1 Abstract 109 5.2 Introduction 109 5.3 Methods 111 5.4 Results 115 5.4.1 Effects of capture and handling