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Ecology and Evolution of Cardiaspina Psyllids, Their Bacterial Endosymbionts and Parasitoid Wasps

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Ecology and Evolution of Cardiaspina Psyllids, Their Bacterial Endosymbionts and Parasitoid Wasps Ecology and evolution of Cardiaspina psyllids, their bacterial endosymbionts and parasitoid wasps Aidan Aaron Gordon Hall BSc (Honours) Thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy Hawkesbury Institute for the Environment Western Sydney University March 2016 Acknowledgements Firstly, and most importantly, I would like to thank my principal supervisor Markus Riegler. Without Markus’ knowledge, guidance and support none of this PhD would have been possible. Markus’ passion for science is second-to-none and he was always willing to chat about ideas. I also wish to thank my co-supervisors, James Cook and Scott Johnson, for their feedback and contributions. James’ knowledge of host – parasitoid systems and coevolution were a huge asset. Secondly, my gratitude goes to Martin Steinbauer and Gary Taylor for advice about psyllid ecology and taxonomy, and to Jennifer Morrow for sharing her experience and knowledge in the molecular lab. Special thanks go to the lab group of Markus Riegler, both past and present, whose friendship, support and feedback on early drafts of manuscripts in preparation were of great importance. In particular, I must note my office mates throughout the entire PhD experience, Andrew Gherlenda and Timothy Sutton, both of whom were invaluable for their friendship and bouncing around ideas. Thanks also to my housemates over the past four years, Andrew Gherlenda, Caroline Fromont, Courtney Campany, James Ryalls, Lisa Bromfield, Mindy Hu and Stephen Bullivant for their friendship, watching movies together and playing cards. Thanks to the HIE technical and administrative team for ensuring the smooth running of the labs and arranging all the orders and pick-ups. You were imperative to the completion of this thesis. Thanks to my family for their love, support and encouragement, and for inspiring my passion of science by bringing me up on a diet of David Attenborough and bushwalks. Finally, a very special thank you goes to Jessica Esveld whose love, encouragement and patience saw me over the finishing line. You have given me so much confidence and I cannot begin to describe how much you meant to me over the last year of completing my PhD. I 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. …………………………………….. Aidan Hall March, 2016 II Preface This thesis consists of a general introductory chapter, including literature review, scope, aims, objectives and overarching hypotheses (Chapter 1), followed by four experimental chapters (Chapters 2 – 5), and a final general discussion chapter which synthesises the key findings, limitations and outlines future research directions (Chapter 6). The four experimental chapters have been presented in the format of stand-alone manuscripts for submission to peer-reviewed international journals and thus some repetition will be present. I am the primary author of the entire thesis, including of all experimental chapters, and was principally responsible for the concept, experimental design, data collection, analysis and writing of each chapter, with guidance from my supervisory panel (Markus Riegler, James M. Cook and Scott N. Johnson) as well as other co-authors. Some of these chapters have already been published, or have been submitted to peer- reviewed journals, as outlined below. Chapter 2: Hall, A.A.G., Gherlenda, A.N., Hasegawa, S., Johnson, S.N., Cook, J.M. & Riegler, M. (2015). Anatomy of an outbreak: the biology and population dynamics of a Cardiaspina psyllid species in an endangered woodland ecosystem. Agricultural and Forest Entomology, 17, 292-301. Chapter 3: Hall, A.A.G., Johnson, S.N., Cook, J.M. & Riegler, M. High nymphal host density and mortality negatively impact parasitoid complex during an insect outbreak. Chapter 4: Hall, A.A.G., Morrow, J.L., Fromont, C., Steinbauer, M.J., Taylor, G.S., Johnson, S.N., Cook, J.M. & Riegler, M. (2016). Codivergence of the primary bacterial endosymbiont of psyllids versus host switches and replacement of their secondary bacterial endosymbionts. Environmental Microbiology, 18, 2591–2603. Chapter 5: Hall, A.A.G., Steinbauer, M.J., Taylor, G.S., Johnson, S.N., Cook, J.M. & Riegler, M. Cryptic diversity, host specificity, trophic and coevolutionary interactions in Australian psyllid – parasitoid food webs. III Table of contents Acknowledgements ....................................................................................................... I Statement of authentication .......................................................................................... II Preface ........................................................................................................................ III Abstract ........................................................................................................................ 1 Chapter 1 ...................................................................................................................... 4 1.1 Insect population dynamics .......................................................................... 5 1.1.1 Processes influencing population dynamics and community structure.... 5 1.1.2 Psyllids and their significance .................................................................. 6 1.1.3 Effect of temperature on insects ............................................................... 7 1.1.4 Effect of temperature, rainfall and nutrient fluxes on psyllids................. 8 1.1.5 Competition and resource limitation ........................................................ 9 1.1.6 Psyllid competition and resource limitation ........................................... 10 1.1.7 Impact of natural enemies ...................................................................... 11 1.1.8 Natural enemies of psyllids .................................................................... 13 1.1.9 Significance of bacterial symbioses of insects ....................................... 15 1.1.10 Bacterial endosymbionts of psyllids .................................................. 16 1.2 Molecular tools in community and evolutionary ecology.......................... 17 1.2.1 Food web studies .................................................................................... 17 1.2.2 Coevolution studies ................................................................................ 18 1.3 Research scope and aims ............................................................................ 18 1.3.1 Study system: Cardiaspina psyllids and their associated parasitoids and endosymbiotic bacteria ....................................................................................... 19 1.3.2 Population dynamics of a Cardiaspina sp. outbreak ............................. 21 1.3.3 Constraints on parasitoid control of a Cardiaspina sp. outbreak ........... 21 1.3.4 Evolutionary relationships of bacterial endosymbionts of psyllids ....... 22 1.3.5 Diversity and host specificity of Cardiaspina parasitoids ..................... 23 Chapter 2 .................................................................................................................... 24 2.1 Abstract ...................................................................................................... 25 2.2 Introduction ................................................................................................ 26 2.3 Materials and methods ............................................................................... 28 2.3.1 Data collection period and site description ............................................ 28 2.3.2 Psyllid identification and biology .......................................................... 30 IV 2.3.3 Survey of parasitisation rates and parasitoid characterisation ............... 31 2.3.4 Survey of tree health and density ........................................................... 31 2.3.5 Statistical analysis .................................................................................. 32 2.4 Results ........................................................................................................ 33 2.4.1 Psyllid identification and biology .......................................................... 33 2.4.2 Parasitisation rates and parasitoid identification .................................... 34 2.4.3 Temperature profile and psyllid abundance ........................................... 35 2.4.4 Tree health and density .......................................................................... 37 2.5 Discussion .................................................................................................. 39 2.5.1 Area-wide defoliation of E. moluccana in CPW ................................... 39 2.5.2 Grey box Cardiaspina ............................................................................ 40 2.5.3 Parasitoids and other natural enemies .................................................... 41 2.5.4 Temperature and Grey Box Cardiaspina population dynamics ............ 41 2.5.5 Psyllid population crash as a result of resource depletion and extreme temperature events .............................................................................................. 42 2.5.6 Other potentially contributing factors .................................................... 43 2.5.7 Conclusions ...........................................................................................
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