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Sexual Conflict Over Parental Care in Penduline

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Sexual Conflict Over Parental Care in Penduline SEXUAL CONFLICT OVER PARENTAL CARE IN PENDULINE TITS René Erwin van Dijk A thesis submitted for the degree of Doctor of Philosophy University of Bath Department of Biology and Biochemistry June 2009 COPYRIGHT Attention is drawn to the fact that copyright of this thesis rests with its author. A copy of this thesis has been supplied on condition that anyone who consults it is understood to recognise that its copyright rests with the author and they must not copy it or use material from it except as permitted by law or with the consent of the author. This thesis may be made available for consultation within the University Library and may be photocopied or lent to other libraries for the purposes of consultation. Contents Acknowledgements 4 Summary 8 INTRODUCTION Chapter 1 Introduction – Sexual conflict and cooperation 9 Chapter 2 Post-fertilisation reproductive strategies 15 PROCESS OF DESERTION Chapter 3 Sexual conflict over parental care in penduline tits 28 Remiz pendulinus : the process of clutch desertion Chapter 4 Parents of the Eurasian penduline tit conceal their 37 intention to desert their mate Chapter 5 What games do penduline tits play? 51 Chapter 6 Sex differences in parental care do not explain female- 68 biased uniparental care in Eurasian penduline tits Remiz pendulinus ENVIRONMENT AND PARENTAL CARE Chapter 7 The influence of habitat quality on sexual conflict over 83 care in penduline tits Chapter 8 Parental care strategies in Eurasian penduline tit are 95 not related to breeding densities and mating opportunities THE EVOLUTIONARY RAMIFICATIONS OF SEXUAL CONFLICT Chapter 9 Sexual conflict predicts morphology and behaviour in 107 two species of penduline tits 2 CONCLUSIONS AND FUTURE DIRECTIONS Chapter 10 Conclusions and future directions – sexual conflict 123 over care in penduline tits and beyond References 138 Appendix I Offspring sex ratio in the sequentially polygamous 152 penduline tit Remiz pendulinus Appendix II Strategic differences – the battle of the sexes in 168 penduline tits Appendix III Practical field guide for investigating breeding ecology 173 of penduline tits Remiz pendulinus Appendix IV Fieldwork report expedition Kazakhstan 2008 189 Affiliations of co-authors 203 3 Acknowledgements During my undergraduate studies biology at the University of Groningen, The Netherlands, I sometimes wondered whether a PhD project would really be suitable for me. Like many, I wondered if working on one study species for some four years would perhaps not be very exciting. Once I embarked on my PhD though, it soon turned out it would not be difficult to keep myself amused. Besides being lucky enough to be able to work on one of the most exciting species within research concerning sexual conflict and breeding systems and to have the privilege of living in the world heritage city of Bath, a substantial contribution to keep me happy throughout the past four years came from the fantastic bunch of the people that surrounded me. They have all been very supportive and have made life for the past few years a great deal more enjoyable. First and foremost, I have very much enjoyed the privilege to work with two experts in the fields of sexual conflict and cooperation in breeding systems, my supervisors Tamás Székely and Jan Komdeur . I am very grateful to Tamás, first of all for inviting me to apply for a PhD at Bath after my MSc research project on penduline tits. I have learned an enormous amount from him. He has been extremely supportive throughout my PhD. His enthusiasm for the work we were carrying out has stimulated me a lot and never left me short of ideas where to go next. The opportunities for me to travel and explore new sites and penduline tit populations are also gratefully appreciated. When many attendants at one of the penduline tit meetings giggled at the idea of exploring penduline tits in Kazakhstan and China next year, Tamás was convinced it would be a worthwhile expedition and pursued the idea. It turned out to be a fantastic basis for future comparative research on penduline tits. I am also very grateful for being so accessible when he was in Bath (and not chasing plovers and students in some remote corner of the planet), despite the large number of projects he had to coordinate. The many chats we have had in his office have been enjoyable and at the same time very productive. It never took long to get back with useful and constructive comments on the work I put together, which I know is something not always easy to achieve, yet something extremely useful to get from a supervisor. We all know Tamás does have his grumpy days, but this is generously compensated for by a good and vibrant sense of humour. I am very grateful to Jan for asking me to do my second MSc research project on penduline tits in 2004. I have learned a lot in that project about the fieldwork and important concepts in behavioural ecology and it thus formed an excellent basis from 4 which to start my PhD. Throughout my PhD Jan has also always been very supportive and easily accessible. The many comments he has provided on the various drafts of the chapters in my thesis, have helped enormously to improve the manuscripts. His and Tamás’ comments were often very different and, although sometimes contrasting, they usually complemented each other very well. Furthermore, it has never taken Jan much longer than a day or so to find a suitable student from Groningen to carry out a project on penduline tits and help us to collect the necessary field data. Jan’s good mood and hassle-free approach towards any issue we encountered made it a great pleasure to work with him. I appreciated that whenever I came over to visit Groningen there has always been a desk ready for me, and I could always rest assured that I could leave again having achieved everything I came for. The very different approach between the Animal Ecology Group in Groningen and Tamás’ Biodiversity lab in Bath to tackle research questions have stimulated me to try and find the right balance. Special thanks must also go to Ákos Pogány . It has been a great pleasure to work with him in the field. The long talks we have had about various issues have definitely contributed to the success of our penduline tit project. His skills to organise projects and develop field methods have been crucial to the success of our field work in South Africa and Asia. And also the more informal chats we have had in various pubs in Hungary, South Africa and England have also always been most enjoyable and may or may not have contributed to the success of our project. I am thankful to István Szentirmai for being such a great company during my first field season on penduline tits in Fehértó in 2004 and during several meetings later on. István also deserves the credits for developing most of the field methods. Lídia Mészáros also made sure our time in the field in Hungary has always been very enjoyable. She has been a great person to collaborate with and, like Ákos and István, became a good friend. All three of them were also always kind enough to offer me the opportunity to let me stay at their place at any time. Lídia also dealt with most of the practicalities in Fehértó. The help of all the students and field assistants that have participated in our project, and of the people that have collaborated in or have facilitated to start up projects at various localities is also acknowledged with gratitude: Joris van Alphen, Sander Bot, Dušan Brinkhuizen, Natalino Cuti, Johan van der Dennen, Richard ffrench-Constant, Per Ericson, Zsofia Gergely, Rob Gordijn, Martijn Hammers, Péter Horváth, Martin Irestedt, Corinne Jeffs, Evelien Jongepier, Arno wa 5 Kangeri, Penn Lloyd, Bruno Massa, Emily Mockford, Arjen Pilon, Sergey Sklyarenko, Rafaella Turco, Machiel Valkenburg, Marco van der Velde, Vera Voronova, Arend Wassink, Franjo Weissing and Xutong Yang . Without them we would have been able to collect considerably less data, and some preliminary results, important for the further development of projects, would have been produced at a much later stage or not at all. The members of the biodiversity lab in Bath, a great, international bunch of people, also deserve a great expression of gratitude. They not only helped to create a stimulating working environment, but many of them also became good friends without whom life in academia would have been much less enjoyable: Araceli Argüelles Ticó, Monif Al-Rashidi, Fiona Burns, John Burnside, Richard James, Gabriel Garcia Peña, Freya Harrison, András Kosztolányi, Clemens Küpper, Peter Long, Mark O’Connell, Valérie Olson, Alejandro Serrano Meneses, James St Clair and Richard Young . I am grateful to Nick Davies for taking up the external examination of my thesis and to Steve Dorus for doing the same as my internal examiner. I also thank the organisations and committees that have been responsible for the funding of the various projects, in particular the University of Bath for providing me with a University Studentship. All other financial support is recognised in the acknowledgements of the separate chapters in my thesis. I am grateful to the Kiskunság National Park and Szegedfish Ltd. for providing permission to conduct fieldwork in Hungary and to ESKOM for doing likewise in South Africa. I am also greatly indebted to my parents, Peter and Tineke , who have always generously supported, but never pushed, me to work at the limits of my abilities, even if this meant that, after my brother Ronald moved to Ireland, I would leave The Netherlands too.
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