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Maternal Care and Interactions Within and Between Families: How the Environment and Chemical Communication Shape Family Life in a Social Insect

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Maternal Care and Interactions Within and Between Families: How the Environment and Chemical Communication Shape Family Life in a Social Insect Maternal care and interactions within and between families: How the environment and chemical communication shape family life in a social insect Inauguraldissertation zur Erlangung der Würde eines Doktors der Philosophie vorgelegt der Philosophisch-Naturwissenschaftlichen Fakultät der Universität Basel von Janine Wong aus Stuttgart, Deutschland Basel, 2014 Genehmigt von der Philosophisch-Naturwissenschaftlichen Fakultät auf Antrag von Fakultätsverantwortlicher: Prof. Dr. Mathias Kölliker, Assistant Professor SNF, Universität Basel Betreuer: Prof. Dr. Mathias Kölliker, Assistant Professor SNF, Universität Basel Korreferentin: Prof. Dr. Rebecca Kilner, Universität Cambridge, UK Basel, den 15.10.2013 Prof. Dr. Jörg Schibler, Dekan ACKNOWLEDGEMENTS I would like to thank all the people who were involved with my work and the accomplishment of this thesis in one way or another. First of all, I need to thank my supervisor Mathias Kölliker. Doing a PhD in your group was among the most valuable experiences I ever made. You always knew how to encourage me to go to the next level. Thank you for sharing your knowledge and your positive attitude with me, and for always having your door open for questions and discussions. You always gave me the feeling that my work is much appreciated. Major thanks go to my friend Joël Meunier. I consider myself lucky to have had the chance to learn from you! You never lost your patience with my endless row of questions, even after moving to Mainz. A big part of this thesis is your contribution. I will always admire your passion for science! Thanks to all current and former members of the Earwig Group for discussions, great team work in the lab, and story telling in the office. Also, for conscientiously replacing me in the lab when I was away for conferences, especially Lilian Röllin, Stefan Boos, and Shirley Raveh. Thank you also for teaching me (maybe unknowingly) so much about Switzerland! Big thanks to the core of “the lunch crew” César Metzger and Lucas Marie-Orléach! We spend many years of lunch breaks together and it was always fun. We shared all the ups and downs of a PhD student’s life and it made the hard times much easier. I will see you at the finish line! Thank you to Shirley Raveh for discussions about science and career options, as well as for your support during the stressful finishing phase. I value your honesty and your understanding for knowing when to speak up and when to keep quiet. I am grateful to César Metzger, Pepijn Luijckx, Kiyono Sekii, Tobias Schär, Roberto Abore, Stefan Boos, Sandra Schneider, Lara Sargenti, and Mathias Ganter for various game and movie nights, BBQs, hikes, museum visits, brunches and dinners, Rhine and lake swims, and many other non-work related activities. Danke an Viktor Mislin, Jürgen Hottinger, and Lukas Zimmermann for always helping me out with technical challenges and computer problems. Thank you to the whole Zoological Institute for teaching me so much about Evolutionary Biology during seminars and discussions. I want to thank Christophe Lucas for fruitful collaborations on two of my manuscript. Your knowledge greatly improved our work. Thanks to Rebecca Kilner for reading and evaluating my thesis and for flying in to be the external referee of my defense. Major thanks go to the Swiss National Science Foundation, the Freiwillige Akademische Gesellschaft and the Nikolaus und Bertha Burckhardt-Bürgin-Stiftung, which have for supported my four years of research financially. I thank one anonymous person for letting me down otherwise I would have never ended up in Basel. Und ein riesiger Dank geht an meine Eltern und meine Schwester. Ich danke Euch dafür, dass Ihr immer so viel Geduld mit mir hattet und Interesse an meiner Forschung. Ohne Euren Rückhalt und Eure uneingeschränkte Unterstützung hätte ich das alles nie im Leben geschafft! TABLE OF CONTENTS SUMMARY 6 GENERAL INTRODUCTION 8 CHAPTER 1 16 The evolution of parental care in insects: The roles of ecology, life history and the social environment CHAPTER 2 45 The effect of female condition on maternal care in the European earwig CHAPTER 3 62 The more the merrier? Condition-dependent brood mixing in earwigs CHAPTER 4 78 Maternal origin, mates and shared environment shape chemical signatures or earwigs CHAPTER 5 98 Cues of maternal condition influence offspring selfishness CHAPTER 6 117 Food availability shapes juvenile development and adult life history in the European earwig CONCLUDING REMARKS 136 APPENDIX 142 APPENDIX I 143 One clutch or two clutches? Fitness correlates of coexisting alternative female life-histories in the European earwig APPENDIX II 162 Sibling cooperation through food sharing in an insect with facultative parental care SUMMARY The evolution of parental care represents an important step in the evolution of sociality and is widespread across different species and different taxa. Parental care is a trait that shows a wide diversity regarding duration and forms of care within and between species and is an important field of research both in evolutionary biology and behavioral ecology. However, its importance in these fields has only been recognized relatively recently. Environmental effects influence condition and individuals will choose distinct behavioral strategies to maximize their fitness. Condition can be communicated through chemicals and such condition-dependent cues can be used by conspecifics to adjust their own behavior. The focus of my dissertation was the investigation of environmental effects and chemical signaling on maternal care and within and between family interactions. I used the European earwig (Forficula auricularia) as a model system for my experimental work. As an introductory chapter, I have written a review article about the evolution of parental care in insects (Chapter 1). Here, I summarized present hypotheses about the roles of ecology and life history from the literature and combined them with new suggestions regarding the influence of social interactions on parental care. In my first experiment (Chapter 2), I investigated the influence of maternal nutritional condition on mother-offspring interactions. Female condition was manipulated through a high-food and a low-food treatment. I could show that the period and amount of maternal food provisioning was dependent on the condition of the female. Females in poor condition provided food to fewer nymphs and for a shorter period of time compared to females in good condition. Offspring attendance remained at a constantly high level independent of female condition and was maintained by both the female and the nymphs, suggesting strong benefits of living in a (family) group. In my second experiment (Chapter 3), I investigated effects of nymph condition and food availability on brood mixing in F. auricularia. Females provide care for foreign nymphs that join their brood. This however increases brood size and thus competition between the offspring. Previous work showed that cannibalism is directed primarily against unrelated nymphs under conditions of low food availability. My results showed that the brood mixing dynamics are influenced by the condition of the nymphs, but are independent of the food availability in the environment. The overall degree of brood mixing was high, suggesting again benefits of living in groups. Furthermore, my third experiment (Chapter 4) tested the presence of family specific cuticular hydrocarbon profiles. Insects mainly use cuticular hydrocarbons as means of communication and 6 individual recognition. Previous results from our group show that there are significant negative effects of inbreeding, which makes kin recognition important in this species. I could show that cuticular hydrocarbon profiles are indeed family specific and that the earwigs have the potential to use them to discriminate kin from non-kin. This allows to direct social behaviors to the appropriate individuals and to avoid inbreeding and the associated fitness losses. In a fourth experiment (Chapter 5), I tested the effect of maternal condition-dependent cues on nymph selfishness and survival. In species where parental care is provided, offspring could use cues of parental condition to adjust their begging behavior and their selfishness by varying the degree of sibling competition. My results show that maternal condition cues influence offspring survival depending on the time of breeding, and further suggest that offspring use these maternal condition-dependent cues to adjust their degree of selfishness, which changes during the breeding season. Together with former evidence on maternal sensitivities to condition-dependent nymph chemical cues, my study shows context-dependent reciprocal information exchange about condition between earwig mothers and their offspring, mediated by cuticular hydrocarbons. My last experiment (Chapter 6) investigated the effects of high and low food availability during the juvenile and early adult development and its influences on development, maternal care and egg production. Females of the European earwig represent two distinct phenotypes. One type produces only one clutch, the other type produces two clutches during its life. Previous experiments have shown that the phenotype is not purely inherited genetically, but likely to be condition-dependent. My results revealed that especially restriction in the late juvenile development has negative effects on development and the probability of second clutch production. Environmental conditions experienced early in an individual’s lifetime
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