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Life-History Traits and Temperature-Dependent Life-History Traits and Temperature-Dependent Performance of Tranosema rostrale (Hym.: Ichneumonidae), a Parasitoid of Low-Density Spruce Budworm (Lep.: Tortricidae) Populations by Martin Lukas Seehausen A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy Faculty of Forestry University of Toronto © Copyright by M. Lukas Seehausen 2017 Life-History Traits and Temperature-Dependent Performance of Tranosema rostrale (Hym.: Ichneumonidae), a Parasitoid of Low-Density Spruce Budworm (Lep.: Tortricidae) Populations M. Lukas Seehausen Doctor of Philosophy Faculty of Forestry University of Toronto 2016 Abstract The eastern spruce budworm Choristoneura fumiferana (Clemens) (Lepidoptera: Tortricidae) is one of the most important outbreaking defoliator in conifer forests of eastern North America. In low-density populations, the larval endoparasitoid Tranosema rostrale (Brischke) (Hymenoptera: Ichneumonidae) is known as an important mortality factor, but relatively little information is available about the factors influencing its ability to keep spruce budworm populations low. A series of studies were conducted to investigate T. rostrale’s: 1) reproductive biology and behaviour; 2) seasonal pattern of parasitism and host instar preference; 3) developmental, survival, and reproductive response to temperature; 4) ability to abrogate spruce budworm’s immune response at high temperatures; and 5) spatiotemporal biology in response to changing temperature. Three traits of the parasitoid’s reproductive biology contribute to its successful parasitism: its lack of a pre-mating and preoviposition period, the rapid maturity of its eggs soon after emergence despite being synovigenic, and its efficacy in host searching and oviposition behaviour. As T. rostrale does not prefer to attack any one particular host instar, its seasonal pattern of parasitism is likely influenced by its phenology or competition with the ectoparasitoid Elachertus cacoeciae (Howard) ii (Hymenoptera: Eulophidae). Overall performance by T. rostrale was severely reduced at high temperatures. The negative correlation between the parasitoid’s survival and rearing temperatures above 20°C was associated with a downregulation of T. rostrale’s polydnavirus gene transcription and an enhancement of the accumulation of host immunity gene transcripts. An individual-based simulation model incorporating the parasitoid’s response to temperature allowed insight into unknown facts about its seasonal biology, such as phenology, voltinism, and overwintering strategy. The findings improve our understanding of the influence of temperature on spruce budworm population dynamics via its parasitoids, and our ability to predict pest outbreaks, which is important to develop more effective pest management strategies under scenarios of changing climate. iii Acknowledgements I thank my supervisor Prof. Sandy M. Smith for accepting me into the PhD program in Forestry. Her open, friendly, and calm personality gave me the necessary environment to do a PhD and to write the present document. I appreciate her positive feedback and encouragement to pursue higher goals. A special thanks goes to her for her lessons in biological control and integrated pest management, that widened my horizon beyond the work of this dissertation. I wish to thank Dr. Jacques Régnière for his enormous support as supervisor, for inviting me to work in his lab at the Laurentian Forestry Centre in Québec, for letting me use his resources, and for financial support. I am especially grateful for his patient introduction into statistics and advanced modeling approaches and for correcting numerous first drafts of manuscripts. His everlasting help and encouragement pushed me to ever higher goals without losing sight of the ground. Many thanks to Dr. Véronique Martel for supervision and sharing her resources and knowledge with me. The frequent discussions with her had a marked influence on the experiments I did and the interpretation of my results. I thank her for organizing social events that brought the entomology research groups at the Laurentian Forestry Centre in Québec so close together, allowing for this huge amount of work to get done with so much fun. I also wish to acknowledge Prof. Marie-Josée Fortin, Prof. Marc W. Cadotte, Prof. D. Chris Darling, and Dr. Taylor Scarr for accepting to be on my PhD committee and providing feedback on experimental design, interpretation of results, and writing throughout my studies. A special thanks goes to Dr. Michel Cusson, who so openly welcomed me to do part of my work in his lab and introduced me to the world of molecular biology. His door was always open for me to have very fruitful discussions about T. rostrale, PDVs, spruce budworm, and science in general. I will never forget his advice for good scientific practices and the art of writing a scientific article. A huge thank you to the students Pascale Bolduc, Sophie Ouellet, Michael Leblanc, Myriam Labrecque, Anne-Marie Dion, Shannie Parent, Maxence Bory, and Hedvig K. Nenzén, as well as the technicians Simon Trudeau, Maxime Moisan, and Paule Huron for helping me so diligently with my experiments, be it for long hours in the lab on weekdays or weekends, or outside in the forest during sunshine, rain or snow. Thanks to Maria Toledo-Garibaldi for doing Fig. 1.2, and to the volunteers Charles Heller, Nina Socolov, and Jennifer Baici for going through the numerous iv Malaise trap samples. Without you all, it would have been impossible to conduct the many experiments presented in this dissertation! Many people from the Laurentian Forestry Centre helped me during the summers and beyond with my experiments. In this context, I want to acknowledge Catherine Béliveau, Don Stewart, and Abdelmadjid Djoumad for helping me with the molecular experiments, Dr. Lucy Royer, Dr. Johanne Delisle, and Dr. Deepa Pureswaran for good discussions and help with experiments, and Ariane Bechard for taking care of all the little things that usually slip my mind. I also thank Rémi Saint-Amant for his help with BioSIM and the model described in Chapter 6, your programming skills are truly incredible! I am grateful to Pierre Therrien, Denis Simoneau, Guylaine Trudel, and Claudine Dussault of the Ministère des Forêts, Faune et Parcs Québec (MFFPQ) for insect rearing, for providing me with T. rostrale and spruce budworm larvae, as well as other in-kind support for my experiments. My colleague and friend Justin Gaudon deserves special acknowledgements here for all the fruitful discussions, for sharing my passion about parasitoids and statistics, and for always being there when help was needed. With you, I always remember that knowledge is powerful only when it is shared; thanks for that! Thank you Rhoda deJonge, Amany Mansour, Susan Frye, Pei-Yu Kuo, Janani Sivarajah, Nigel Gale, John Schurman, Lucas Roscoe, Graham Watt, Richard Dickinson, Dr. Nurul Islam, Eric Davies, Nicolas Tanguy, and Rana Roy for being good colleagues that helped me to grow professionally and as a person. I also thank Nigel Gale and Prof. Sean Thomas for introducing me into the world of plant ecology and biochar. Thanks to all members of the Faculty of Forestry staff for helping with the necessary paper work during my PhD program and to all members of the Forestry Graduate Students’ Association for their help and trust in me. Financial support was granted through SERG-International by the provincial governments of Ontario, Quebec, Newfoundland, Nova Scotia, Saskatchewan and the Société de protection des forêts contre les insectes et maladies (SOPFIM), as well as by the Atlantic Canada Opportunities Agency and Natural Resources Canada (Canadian Forest Service). Additional funding was provided through a Natural Sciences and Engineering Research Council Cananda (NSERC) Discovery Grant to SMS. Funding for myself was provided by the Ontario Trillium Scholarship through the University of Toronto, for which I’m eternally thankful because I could never had covered the high school fees and living expenses in Toronto without it. v My friends Dr. Alvaro Fuentealba, Maria Constanza Diaz, and Dr. Roberto Quezada deserve a mention here for always having an open door during my long summers in Quebec; thank you! I extend everlasting gratitude to my parents Hildegard and Udo Seehausen, who always provided me with the best possible education and supported me in all my decisions with such positive attitude and good advice. I am also thankful to my grandparents Irmgard Schnellbächer and Herta Seehausen, as well as to my sisters and brothers, who supported me on my way to this dissertation. The biggest thank-you goes to my lovely wife Dr. Julieta Caballero, my son Elias, and the little Sam for always being by my side, supporting me and letting me see every day for what it is all worth it. All my work is dedicated to them! vi Table of Contents Acknowledgements ........................................................................................................................ iv Table of Contents .......................................................................................................................... vii List of Tables ................................................................................................................................. ix List of Figures ..................................................................................................................................x Chapter 1 General Introduction ...................................................................................................1
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