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Interactions Between the Woodwasp Sirex Noctilio and Co-Habiting Phloem- and Woodboring Beetles, and Their Fungal Associates in Southern Ontario

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Interactions Between the Woodwasp Sirex Noctilio and Co-Habiting Phloem- and Woodboring Beetles, and Their Fungal Associates in Southern Ontario Interactions between the Woodwasp Sirex noctilio and Co-habiting Phloem- and Woodboring Beetles, and their Fungal Associates in southern Ontario by Kathleen Ryan A thesis submitted in conformity with the requirements for the degree of Doctor of Philosophy Faculty of Forestry University of Toronto ©Copyright by Kathleen Ryan 2011 Interactions between the Woodwasp Sirex noctilio and Co-habiting Phloem- and Woodboring Beetles, and their Fungal Associates in southern Ontario Kathleen Ryan Doctor of Philosophy Faculty of Forestry University of Toronto 2011 Abstract In its introduced southern hemisphere range, Sirex noctilio causes considerable mortality in non-native pine forests. In its native Eurasian range however, S. noctilio is of little concern perhaps due to interactions with a well-developed community of pine-inhabiting insects and their associated microorganisms. If such interactions occur, they may limit the woodwasp’s impact in its newly introduced range in North America. My research addresses two broad questions: 1) Does S. noctilio share its habitat with other insects and if so, with whom? 2) Is there evidence that co-habitants affect S. noctilio , and if so how might such interactions occur? Field studies undertaken to describe the woodwasp’s host-attack ecology in Pinus sylvestris showed S. noctilio activity occurred between mid-July and late August, and other phloem- and woodborers sometimes entered the tree after the woodwasp. Tree mortality occurred from two weeks to several months after initial woodwasp symptoms. Suppressed or intermediate trees, those with ≤ 25% residual foliage, or those with stem injury or previous woodwasp symptoms were most likely to have symptoms of woodwasp attack. ii A second field study conducted to identify associated insect species in S. noctilio -infested Pinus sp. showed the wasp was sometimes found alone, but usually shared the tree with other phloem- or woodboring insects, most commonly the curculionids Tomicus piniperda , Pissodes nemorensis and Ips grandicollis and the cerambycid Monochamus carolinensis . I found no indication that wasps were absent when beetles were present, but there was evidence that woodwasps were less abundant, but larger, when beetles were present. Experiments showed that indirect interactions can occur between the two insect groups via fungal associates of one or both. In the laboratory, the woodwasp symbiont was outcompeted by two beetle-associated fungi, Leptographium wingfieldii and Ophiostoma minus, over a range of temperatures. Under field conditions the woodwasp was able to detect and avoid ovipositing in P. sylvestris inoculated with L. wingfieldii, but its oviposition was unaffected by O. minus . My results show that insects co-habiting pine with S. noctilio have potential to exert a measure of biological control on the woodwasp and may help to limit its impact in North America. iii In memory of Peter de Groot Mentor and friend iv Acknowledgements As with any endeavour of this scope, many people were instrumental in the completion of this project. Special thanks to my advisors Peter de Groot who provided guidance, and who shared his wisdom throughout the completion of project and Sandy Smith who taught me that anything is possible. Thanks to my advisory committee members Jean-Marc Moncalvo, Peter Kotanen and Taylor Scarr for advice and assistance, and especially for their enthusiasm about this project. Thank you to Chuck Davis for providing assistance and guidance on all aspects of this project, and whose contribution was immeasurable. Data was collected with the assistance of Sheung Au, Madelaine Danby, Sarah Drabble, Megan Evers, Sean Strong and Kate Surowiak. I am grateful for their conscientious efforts. Data collection was facilitated by the outstanding technical assistance provided by Ian Kennedy and John McCarron. Many people provided essential advice for the fungal component of this project including Simona Margaritescu, Martin Hubbes, James Reid, and Tony Ung. Ed Czerwinski provided training and suggestions on tree assessment, but more importantly showed me how to see a forest through an insect’s eyes. Hugh Evans gave advice, helped with finding sites, and patiently answered a myriad of questions. Thanks to Isabelle Ochoa for teaching me bark beetle and woodborer identification. Isabelle, Katherine Nystrom, Reg Nott and David Langor identified many of the insects used in this project, and Serge Laplante verified identifications. v Many students at the Faculty of Forestry helped with advice or field work, of those I’d especially like to thank Laura Timms, Smith Sundar, Jeff Boone, James Dennis and Nick Rudzik. Thank you to my husband Sean for his love and support through this endeavour, for reminding me of the big picture, and for helping with every aspect of this project. I am forever grateful to you. Thanks to my family and friends for their patience, support and perseverance through my time completing this project, and especially for never asking, why? Several private landowners, Sandbanks Provincial Park (Don Bucholz), Toronto Regional Conservation Authority (Tom Hildebrand), Grey-Bruce Conservation Authority (Ken Goldsmith), Simcoe County (Graeme Davis) and Canadian Forces Base Borden (Bill Huff) provided sites and access to trees. The Ontario Tree Seed Plant (Al Foley) provided facilities. Funding was provided by Natural Resources Canada - Alien Invasive Species Program, the National Sciences and Engineering Council of Canada and the Ontario Ministry of Natural Resources. vi Table of Contents Abstract..................................................................................................................................... ii Acknowledgements.................................................................................................................. iv Table of Contents.................................................................................................................... vii List of Tables ........................................................................................................................... ix List of Figures........................................................................................................................... x List of Appendices .................................................................................................................. xii Introduction............................................................................................................................... 1 Chapter 1: Literature Review.................................................................................................... 5 1 Insect-fungus complex....................................................................................................... 5 2 The life history of Sirex noctilio........................................................................................ 8 3 Host trees ......................................................................................................................... 12 4 Population factors ............................................................................................................ 13 4.1 Natality Factors......................................................................................................... 13 4.2 Population limiting factors........................................................................................ 14 5 Interaction among woodboring insects and their fungal associates................................. 15 6 Fungal interactions........................................................................................................... 18 7. Summary......................................................................................................................... 20 Chapter 2: Aspects of Sirex noctilio host colonization ecology of P. sylvestris; preferred host tree condition, timing of attack, host death, and spatial and temporal variation ................................................................................................................................................. 21 Abstract............................................................................................................................... 21 1 Introduction...................................................................................................................... 22 2 Methods............................................................................................................................ 24 2.1 Field sites .................................................................................................................. 24 2.2 Symptoms of S. noctilio oviposition activity............................................................ 27 2.3 Field methods............................................................................................................ 28 2.4 Data Analysis............................................................................................................ 30 3 Results.............................................................................................................................. 31 4 Discussion........................................................................................................................ 39 Chapter 3: Effect of two phloem- and woodborers-vectored fungi on the on-host search and oviposition behaviour of Sirex noctilio on Pinus sylvestris trees and logs............................. 42 Abstract..............................................................................................................................
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