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(The Psyllid, Aphalara Itadori Shinji) for Invasive Knotweeds

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(The Psyllid, Aphalara Itadori Shinji) for Invasive Knotweeds Manipulating Phenotypic Plasticity to Improve Population Establishment of a Classical Biological Control Agent (the Psyllid, Aphalara itadori Shinji) for Invasive Knotweeds by Timothy Owen Skuse A thesis submitted in conformity with the requirements for the degree of Master of Science, Forestry Faculty of Forestry University of Toronto © Copyright by Timothy Owen Skuse 2018 Manipulating Phenotypic Plasticity to Improve Population Establishment of a Classical Biological Control Agent (the Psyllid, Aphalara itadori Shinji) for Invasive Knotweeds Timothy Owen Skuse Master of Science, Forestry Faculty of Forestry University of Toronto 2018 Abstract The psyllid Aphalara itadori was approved for release as a biocontrol agent for invasive knotweeds (Fallopia sp.) in Canada, but has had limited success establishing in the field. Establishing biocontrol agent populations in the field is a common problem and one unexplored possibility in improving establishment is the manipulation of an agent’s phenotype prior to release. We developed a series of laboratory bioassays to assess the psyllid’s fecundity and offspring fitness in response to host plants of varying quality. Results show an increase in fitness measures that appear unrelated to the host plant quality and more closely determined by temperature or humidity. This suggests phenotypic plastic responses in A. itadori populations are less sensitive to changes in their host and more sensitive to environmental variables. Determining which environmental factors drive changes in these fitness variables and rearing A. itadori in these conditions prior to release may improve population establishment. ii Acknowledgments I would like to thank my supervisor Prof. Sandy M. Smith for accepting me into the MScF program, for guiding me through the process to complete this thesis, for financial support, and for her valuable edits needed to finish this document. I would also like to thank Dr. Rob Bourchier for his support as a supervisor, welcoming me into his lab at AAFC Lethbridge, and providing me with the opportunity to work with A. itadori. His guidance was critical in helping to develop the ideas and planning out the experiments that make up this thesis. His help with statistics and edits were invaluable as well as the financial support provided. My thanks to Janine Brooke and Karma Tiberg for their help in all things related to working at AAFC Lethbridge. Their help and patience with preparing for and carrying out my (sometimes frustrating) experiments was immensely appreciated and I could not have finished this document without them. I would also like to acknowledge Prof. Megan Frederickson and Prof. Peter Kotanen for accepting to be on my MScF committee and providing feedback on experimental design and interpretation of results. A big thank you to the students Becky Bedell, Johnathan Reid, and Diana Wilches who helped with setting up and checking experiments. Financial support was provided through Queen Elizabeth II/Buell Graduate Scholarships in Science and Technology and facilitated me completing this work. iii Table of Contents Acknowledgments ......................................................................................................................... iii List of Tables .................................................................................................................................. vi List of Figures ................................................................................................................................. ix List of Appendices ........................................................................................................................ xii Introduction .................................................................................................................................... 1 The invasive knotweed complex ........................................................................................................... 1 Challenges of knotweed management ................................................................................................. 5 Biological control using the psyllid Aphalara itadori Shinji .................................................................. 6 Establishment of classical biological control agents ........................................................................... 10 The role of phenotypic plasticity ........................................................................................................ 13 Research goals .................................................................................................................................... 14 Materials and Methods ................................................................................................................ 16 Plant and insect material .................................................................................................................... 16 Preliminary host quality experiments ................................................................................................. 16 Bioassay experimental design ............................................................................................................. 17 Experiment 1: Damaged versus intact host plants ............................................................................. 19 Experiment 2: Juvenile versus mature host plants ............................................................................. 21 Experiment 3: Hardened leaves .......................................................................................................... 22 Experiment 4: Psyllid density manipulation........................................................................................ 23 Statistical analysis ............................................................................................................................... 24 Results ........................................................................................................................................... 26 Preliminary host quality tests ............................................................................................................. 26 Experiment 1: Damaged versus intact plants ..................................................................................... 26 Experiment 2: Juvenile versus mature plants ..................................................................................... 32 Experiment 3: Hardened leaves .......................................................................................................... 36 Overall Morphometrics ....................................................................................................................... 39 Experiment 4: Density manipulation experiment ............................................................................... 40 Further modelling ............................................................................................................................... 44 iv Discussion ..................................................................................................................................... 48 Conclusions, recommendations, and future directions .............................................................. 62 References .................................................................................................................................... 64 Appendix A ................................................................................................................................... 74 v List of Tables Table 1. Factors and corresponding levels used in models predicting Aphalara itadori fitness responses in Experiment 1. Each F0 A. itadori used in the experiment came from the same natal environment (laboratory colony) and F1 and F2 A. itadori were reared on either damaged or intact Japanese knotweed (Fallopia japonica) plants. ................................................................... 21 Table 2. Factors and corresponding levels used in models predicting Aphalara itadori fitness responses in Experiment 2. Each F0 A. itadori used in the experiment came from the same natal environment (laboratory colony) and F1 A. itadori were reared on either juvenile or mature Japanese knotweed (Fallopia japonica) plants. Psyllids from each generation and natal treatment were exposed to either juvenile or mature knotweed resulting in six possible generation*parent natal treatment*treatment combinations. ...................................................................................... 22 Table 3. Factors and corresponding levels used in models predicting Aphalara itadori fitness responses in Experiment 3. Each F0 A. itadori used in the experiment came from the same natal environment (laboratory colony) and F2 A. itadori were reared on mature Japanese knotweed (Fallopia japonica) plants from Experiment 2. Psyllids from each generation and natal treatment were exposed to either hardened or unhardened knotweed resulting in four parent natal treatment*treatment combinations. ........................................................................... 23 Table 4. GLM results of Experiment 1 quasipoisson model testing whether Aphalara itadori oviposition is affected by host plant treatment, generation number, natal environment and the interaction term between treatment and natal environment. Asterisk indicates significance of P<0.05. .........................................................................................................................................
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