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Evolutionary Ecology of Fig Wasps Associated with the Port Jackson Fig

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Evolutionary Ecology of Fig Wasps Associated with the Port Jackson Fig Evolutionary ecology of fig wasps associated with the Port Jackson fig Timothy Leonard Sutton BSc (Honours) Thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy Hawkesbury Institute for the Environment Western Sydney University 2016 Acknowledgements I would first like to acknowledge my supervisor, James Cook, for his support and guidance throughout the last four years. Thanks for having the faith to take me on as a student at very short notice (and from the other side of the planet), for your willingness to chat science, sport and the world, and for teaching me not to take things too seriously. Huge thanks to my co-supervisors, Markus Riegler and Jane DeGabriel, for all of their hard work and support throughout my candidature. Markus’ enthusiasm and work ethic are second-to-none and I am grateful for the encouragement he gave me to pursue a PhD. Jane not only taught me what real fieldwork is like, but was also a calming influence during stressful periods and an invaluable resource on how to forge a career in science. The amount of work she (and, by association, Clementine) put in during the final weeks of my candidature is unbelievable, and I’m not sure how I would have got over the line without it. I am grateful for having received an Australian Postgraduate Award and top-up scholarships from Western Sydney University and Hawkesbury Institute for the Environment. I would like to acknowledge the Hawkesbury Foundation for granting me the F.G. Swain Award in 2013, which funded a large portion of the molecular work for this thesis. Thanks also to the Australian Biological Resources Study and the Australian Entomological Society for funding my attendance at numerous conferences in Australia and Europe. I would like to thank my office mates of the past five years; Aidan Hall and Andrew Gherlenda. It’s been a pleasure to share this journey with you and see the world together, and I am sure many more adventures await us. Thanks also to my close friend and confidant, Adam Frew, for sharing his thoughts and support through the good times and the bad. This rollercoaster of a PhD was made a lot easier with friends like you. I would like to acknowledge past and present members of the Cook and Riegler lab groups – Caroline Fromont, Ashley Montagu, Will Balmont, Lluvia Flores-Renteria, I Peter Kern, Jennifer Morrow, Duong Nguyen, Shannon Smith and Robert Müller – for sharing ideas, support and encouragement, and assisting with field and lab work throughout my candidature. Jennifer, in particular, was an extremely helpful during my early years in the molecular lab and was always willing to help me out or answer any questions I had (and there were a lot!). Much of the fieldwork for this thesis was undertaken in collaboration with Jane DeGabriel, who led the fieldwork component in northern Queensland. Thanks also to Caroline Fromont, Will Balmont, Eliza Langford, Aidan Hall, Adam Frew, Nina White, Jennifer Hechinger and Ashley Montagu for their assistance in the field and/or setting up experiments. Aidan Hall, Andrew Gherlenda and Ashley Montagu gave helpful advice on manuscripts submitted during my PhD candidature. I also owe a big thanks to Paul Rymer for guiding me through the statistical component of population genetics in the early days of my candidature. The research conducted at the Hawkesbury Institute for the Environment would not be possible without the wonderful work of the admin and technical staff. I would like to acknowledge David Harland, Jenny Harvey, Patricia Hellier, Lisa Davison, Gillian Wilkins, Gavin McKenzie, Kerri Sherriff, Nga Le, Markus Klein, Carolina Janitz and Jocelyn King for the hard work they put in everyday to ensure everything at the HIE runs smoothly. I owe thanks to Leanne and David – my ‘weekend parents’ – for feeding and housing me for at least two-sevenths of the past three-and-a-bit years. Thanks for your kind words of support and encouragement, and for letting me wander in at ungodly hours of the morning after running all-night experiments. To my family, particularly my parents – Diane and Glenn – thank you for your amazing patience and support (both moral and financial) throughout all of my educational endeavours. You taught me the importance and value of education and hard work, and this is the culmination of those lessons. I appreciate everything you have done for me (though it doesn’t always show), and I am forever indebted to you. Finally, my love and thanks to Eliza. I cannot begin to thank you enough for the amazing support, encouragement and love you have blessed me with. You believed in me when I didn’t believe in myself, and I could not have done this without you. II Statement of authentication The work presented in this thesis is, to the best of my knowledge and belief, original except as acknowledged in the text. I hereby declare that I have not submitted this material, either in full or in part, for a degree at this or any other institution. … ………. Timothy Sutton March 2016 III Preface The experimental chapters presented in this thesis were written as a series of independent papers that have been published, or are intended for publication, in peer- reviewed journals. Consequently, each of the five experimental chapters was structured in the style of the chosen journal. As each chapter represents a stand-alone paper, and is therefore self-contained, each has its own more detailed introduction and discussion and there is minor repetition between them. Each chapter was co- authored; nevertheless, I am the principal author and contributor to this work. I conceptualised the research with guidance from my supervisory panel. Furthermore, I collected and analysed the data, and wrote the manuscripts with editorial advice from my supervisory panel. Because each chapter is written as it appears, or will appear, in publication, I have retained the use of the collective pronoun. In addition to the experimental chapters, I conclude this thesis by highlighting the significance of the research, and discussing its limitations and potential directions of future research (Chapter 7). Below are details of published papers and papers in review that have arisen from the research presented in this thesis. Details are accurate at the time of submission. Chapter 2: Sutton, TL, Riegler, M and Cook, JM (2015) Characterisation of microsatellite markers for fig-pollinating wasps in the Pleistodontes imperialis species complex. Australian Journal of Zoology, 63(2), 122-126. Chapter 3: Sutton, TL, Riegler, M and Cook, JM (2016) One step ahead: a parasitoid disperses farther and forms a wider geographic population than its fig wasp host. Molecular Ecology, 25(4), 882-894. Chapter 4: Sutton, TL, DeGabriel, JL, Riegler, M and Cook, JM (in revision) Barriers from barcodes: how reliable are DNA barcodes in fig wasps? Heredity. IV Abstract Geography plays an important role in the study of evolutionary ecology. It has implications for the coevolution of species interactions, our ability to accurately and confidently delimit species boundaries, and can influence species’ adaptations that may provide resilience, or make them vulnerable, to climatic change. A geographic consideration of these issues can help to improve our understanding of the processes generating biodiversity and its fate under predicted climate change. In this thesis, I investigated the genetic structure of a pollinator-parasitoid interaction over a wide geographic range, assessed the reliability of DNA barcoding under different geographic scenarios, and examined the resilience of a temperate fig- pollinating wasp species to climate change. Figs are keystone species in many regions and the fig – pollinator system is a classic example of an obligate mutualism. In this thesis, I explored the evolutionary ecology of pollinator and non-pollinator fig wasps associated with the Port Jackson fig (Ficus rubiginosa) in eastern Australia. This system is particularly interesting due to the sheer number of species that interact within F. rubiginosa fruits. There are five recognised pollinating wasps of F. rubiginosa: Pleistodontes imperialis sp. 1, 2, 3, 4 and 5 (sensu Haine et al. 2006). All of these are parasitised by Sycoscapter non-pollinating fig wasps, of which there are three species: Sycoscapter sp. A, B and C. I begin this thesis by describing the development of microsatellite markers that I characterised for population genetic studies in Pleistodontes imperialis sp. 1 and demonstrating their utility in other species of the P. imperialis complex (Chapter 2). Next, I applied these markers to a comparative population genetic study of a fig- pollinating wasp (P. imperialis sp. 1) and its parasitoid (Sycoscapter sp. A) (Chapter 3). I found that the pollinator comprises two disjunct populations while the parasitoid exists as a single population throughout eastern Australia. These results suggest that the two pollinator populations have the potential to diverge ecologically in their responses to a common antagonist, which may facilitate speciation. Furthermore, I provide the first evidence that a fig wasp parasitoid likely disperses farther than its associated fig-pollinating wasp, challenging common perceptions about parasitoid fig V wasp dispersal that were based solely on measurements of flight heights in tropical rainforests. I then shifted my focus to test the reliability of DNA barcoding for delimiting fig wasp species in sympatry and allopatry (Chapters 4 and 5), focussing on the P. imperialis complex of closely related wasp species associated with F. rubiginosa. I found that Bayesian clustering analyses help to identify potential F1 hybrids, but reveal minimal interspecific gene flow between sympatric fig wasp species. Thus, my results validate the use of DNA barcoding for species delimitation of fig wasps pollinating F. rubiginosa. I then scaled genetic differentiation of allopatric P. imperialis sp.
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