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Natural Enemies of Wood-Boring Beetles in Northeastern Temperate

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Natural Enemies of Wood-Boring Beetles in Northeastern Temperate Natural Enemies of Wood-boring Beetles in Northeastern Temperate Forests and Implications for Biological Control of the Emerald Ash Borer (Coleoptera: Buprestidae) in North America by Justin Michael Gaudon A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy in Forestry Faculty of Forestry University of Toronto © Copyright by Justin M. Gaudon 2019 Natural Enemies of Wood-boring Beetles in Northeastern Temperate Forests and Implications for Biological Control of the Emerald Ash Borer (Coleoptera: Buprestidae) in North America Justin Michael Gaudon Doctor of Philosophy in Forestry Faculty of Forestry University of Toronto 2019 Abstract The emerald ash borer (hereafter EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is a wood-boring beetle accidentally introduced into North America during the 1990s, and has since been killing millions of ash trees, Fraxinus spp. L. (Lamiales: Oleaceae), as it spreads across Canada and the USA. Native North American natural enemies, especially parasitoid wasps, are important mortality factors of EAB, but little information is available on their arrival and detection in EAB-infested regions, and their feasibility for augmentative biological control against EAB is uncertain. Two important native parasitoid groups, Phasgonophora sulcata Westwood (Hymenoptera: Chalcididae) and Atanycolus spp. Foerster (Hymenoptera: Braconidae), were investigated to determine (1) factors influencing their capacity to disperse, (2) vegetation and habitat characteristics influencing their local abundance and role in EAB mortality, (3) whether their populations can be augmented to increase parasitism of EAB, and (4) how best to detect and monitor their populations as EAB continues to spread. The weak dispersal capacity of P. sulcata suggests it should be released as pupae close to EAB if used in an augmentative biological control program. Forest vegetation and habitat structure determine local abundance of P. sulcata and Atanycolus spp., and tree biomass, tree condition, and floral resource availability are important predictors of high parasitism on EAB. Relocating parasitoid-infested ash logs to EAB-infested ii sites can significantly augment populations of native parasitoids to increase EAB parasitism by 64.8 ± 18.1 % three years after introduction. Purple prism traps can be used to detect and monitor changes in populations of relocated P. sulcata and Atanycolus spp. These findings improve our understanding of the role of native natural enemies in suppressing EAB population growth and slowing ash tree mortality in North America. iii Acknowledgements First, I thank my academic supervisor, Prof. Sandy Smith, for accepting my application to do graduate work in her laboratory at the Faculty of Forestry, University of Toronto. Her strong mentorship throughout my PhD program has helped me grow as an independent researcher and a team leader in both a field and laboratory setting. She has shown me what it takes to produce rigorous, high-quality scientific research; how to effectively communicate my work; and has even calmed my anxiety during some of those trying times that many of us encounter at some point while completing a degree. I also thank my supervisory committee members, Profs. Jeremy Allison, Chris Darling, and Sean Thomas for providing valuable advice and feedback throughout all stages of this work, including experimental planning and design, data collection, data analysis, interpretation of my results, and writing. Thank you to Dr. Taylor Scarr for also sitting on my supervisory committee as a “non-voting” member and providing the same support. Special thanks go to Dr. Danijela Puric-Mladenovic for her immense support throughout my studies, her involvement with my research, and for participating as an external examiner on my qualifying exam committee and both departmental and final oral exam committees. Thank you to Dr. Richard Westwood for participating as the external appraiser on my final oral exam committee. Early in my PhD program, I received mentorship and help starting one of my research projects from Dr. Lucas Roscoe. Later, I had help growing my knowledge of statistics, especially using R, and collecting parasitoids used for some of this work from Dr. Chris MacQuarrie. Thank you both for your direction and assistance during such critical times in my career. Much of this research would not have been possible without the various in-kind support received from federal and provincial governments, municipalities, conservation authorities, and iv other land owners. Thank you to Kristjan Vitols (City of Toronto), Stacey Bowman (York Region), Phillip Davies (Lake Simcoe Region Conservation Authority), Tys Theysmeyer and Dr. David Galbraith (Royal Botanical Gardens), Curtis Marcoux and Candace Karandiuk (Town of Oakville) for permission to collect data; and Kristjan Vitols (City of Toronto), Mary Orr (Canadian Food Inspection Agency), and Al Foley and Sarah Drabble (Ontario Tree Seed Plant) for storage space. Thank you to Gene Jones and Drs. Krista Ryall and Barry Lyons at the Canadian Forest Service for sharing their insight, especially with respect to collecting parasitoids. It would have been difficult to complete such a large body of work without help from research assistants. Thank you to Jing (Iris) Hu and Charlotte De Keyzer for their help in the laboratory and especially in the field when we worked through hot or rainy weather or began working before sunrise. Special thanks to my colleague and good friend, Dr. M. Lukas Seehausen. This research would certainly not be what it is without our fruitful conversations about invasive forest insects, parasitoids, and statistics. Many thanks to Richard Dickinson for both resources and assistance with plant identification throughout my thesis work, especially Chapter 3. Thanks to Rhoda DeJonge, Susan Frye, Janani Sivarajah, Nigel Gale, Dr. Nurul Islam, Eric Davies, Nicolas Tanguay, Kenneth Dearborn, Ian Jones, and others who have been excellent colleagues that challenged me to do the best that I can. I appreciate that you were there to celebrate the highs and help me through the lows of completing a PhD. I express my gratitude to my mother-in-law, Susan Ingram; Aunt Geni; parents, Raymond and Judy Gaudon; and grandparents, Donald and Inez Gannett, for their support throughout my academic career; and my dogs, Marley, Macey, and the late Abby, for always giving me the time to take a walk. v Last, my biggest thank you goes to my wife and best friend, Michelle Gaudon, for her continued love and endless support throughout my PhD program and as I proceed throughout my career. She is and will always be my rock. I dedicate this to you. vi Table of Contents Acknowledgements........................................................................................................................iv Table of Contents ......................................................................................................................... vii List of Tables .................................................................................................................................ix List of Figures ................................................................................................................................. x Appendices .................................................................................................................................. xii Chapter 1 General Introduction.................................................................................................. 1 1.1 Invasive Species and their Success in Novel Environments ................................................. 1 1.2 Emerald Ash Borer in North America .................................................................................. 2 1.3 Parasitoid Definitions and Natural History ........................................................................... 5 1.4 North American Natural Enemies of Agrilus (Coleoptera: Buprestidae) ............................. 6 1.5 Biological Control Using Parasitoid Wasps ........................................................................ 10 1.6 Adult Parasitoid Dispersal ................................................................................................. 13 1.7 Spatial Distribution of Ash Trees, Emerald Ash Borer, and its North American Parasitoids ................................................................................................................................................... 15 1.8 Sampling Parasitoid Populations ........................................................................................ 17 1.9 Thesis Objectives and Outline ............................................................................................ 18 Chapter 2 Factors Influencing the Dispersal of a Native Parasitoid, Phasgonophora sulcata, Attacking the Emerald Ash Borer: Implications for Biological Control............................... 24 2.1 Abstract ............................................................................................................................... 24 2.2 Introduction ......................................................................................................................... 25 2.3 Materials and Methods ........................................................................................................ 29 2.4 Results ................................................................................................................................
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