Semiochemical-Mediated Interactions of an Invasive Leaf Miner, Caloptilia Fraxinella (Lepidoptera: Gracillariidae), on a Non-Native Host, Fraxinus Spp
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Semiochemical-Mediated Interactions of an Invasive Leaf miner, Caloptilia fraxinella (Lepidoptera: Gracillariidae), on a Non-Native Host, Fraxinus spp. (Oleaceae) and its Native Parasitoid, Apanteles polychrosidis (Hymenoptera: Braconidae) by Tyler Jonathan Wist A thesis submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Ecology Department of Biological Sciences University of Alberta ©Tyler Jonathan Wist, 2014 Abstract The ash leaf-coneroller, Caloptilia fraxinella (Ely) (Lepidoptera: Gracillariidae) is an introduced moth to the urban forest of Western Canada that specializes on ash (Fraxinus spp.) (Oleaceae). It uses green ash, F. pensylvannica Marsh. var. subintegerrima (Vahl) Fern, and black ash, F. nigra Marsh in its expanded range but prefers black over green ash for oviposition. The oviposition preference for black ash was not linked to enhanced performance of offspring on black ash as green ash supported higher larval survival and faster development than black ash. Younger leaflets are preferred by female moths over older leaflets and green ash elicits more host location flight by gravid females than black ash in wind tunnel experiments. Caloptilia fraxinella adults are attracted to volatile organic compounds (VOCs) released from ash seedlings. The antennae of mated female C. fraxinella consistently detect six VOCs released from black ash and green ash, four of which are common to both species. Synthetic copies of these VOCs elicited as much oriented upwind flight in mated female C. fraxinella as green and black ash seedlings but did not elicit contact with the VOC lure. Lures do not attract more female C. fraxinella than unbaited control traps in field experiments. These experiments lay the foundation for further development of semiochemical lures based on host VOCs to monitor and potentially control populations of C. fraxinella. Apanteles polychrosidis Vierek (Hymenoptera: Braconidae), is identified here as the dominant parasitoid of C. fraxinella in its expanded range. It exhibits protandrous adult emergence and adults mate soon after females emerge. The remainder of the parasitoid complex affecting C. fraxinella in Edmonton is, in order of parasitism rate, Diadegma Förster (near D. fenestrale (Holmgren)) (Hymenoptera: Ichneumonidae), three Sympiesis Förster species (Hymenoptera: Eulophidae) (S. sericeicornis (Nees von Esenbeck), one near S. viridula (Thomson) and one unknown Sympiesis species, and Pteromalus phycidis (Ashmead) ii (Hymenoptera: Pteromalidae). Of these parasitoids, only A. polychrosidis is present in numbers high enough to control populations of C. fraxinella. Parasitism rate by A. polychrosidis of C. fraxinella on black ash is higher than on green. Parasitism is positively dependent on C. fraxinella density on black ash but is negatively density dependent on green ash. Apanteles polychrosidis exhibits differential attraction to the VOCs of each ash species which may mediate the differential parasitism response. In olfactometer experiments, A. polychrosidis use host location cues induced by feeding damage on black ash but not on green ash. Thirteen ash VOCs are detected by the antennae of A. polychrosidis. iii Preface Chapter four of this thesis has been published as Wist T.J. and Evenden M.E. 2013. Parasitoid complex and bionomics of Apanteles polychrosidis on the ash leaf cone-roller. The Canadian Entomologist 145: 416-429. I was responsible for the data collection and analysis as well as the manuscript composition. Maya Evenden was the supervisory author and was involved with concept formation and manuscript editing. iv Acknowledgements A tremendous thank you goes out to my thesis supervisor, Dr. Maya Evenden, for her dedication and patience to guiding and revising this thesis and all of its many projects. Thank you to my advisory committee members Drs. JC Cahil and Nadir Erbilgin for their guidance and suggestions. The members of the Evenden lab at the University of Alberta, both past and present have been excellent companions along the way (Artem Abdukarov, Marius Aurelian, Ronald Batallas, Brad Jones, Jessica Kwon, Boyd Mori, Amanda St. Onge, Chaminda Weeraddana and Caroline Whitehouse). Thanks especially to Joelle Lemmen-Lechelt for her expertise in Caloptilia fraxinella collecting and handling and for much moth discussion. I am indebted to José Fernandez and Henri Goulet (AAFC Ottawa) for identification of Apanteles polychrosidis. Thank you to Chris Saunders of the City of Edmonton for the initial inspiration and push to study this tritrophic relationship in the urban forest. Thank you to Dr. Wilbert Ronald and Jeffries Nurseries of Portage la Prairie, Manitoba for donation of ash seedlings. Thanks to Dr. Stefan Bartram of the Max Plank Institute for Chemical Ecology (Jena, Germany) for donation of (E) 4,8-dimethyl 1, 3, 7 nonatriene. My gratitude is extended to Regine Gries at Simon Fraser University for her skill and knowledge of GC-EAD techniques and for applying them in the search of electrophysiologically-active volatiles from ash trees. This thesis was helped immensely by the field and lab work of many summer students (Brady Chase, Emily Dombowskie, Elizabeth Hodges, Ilona Houston, Jessica Kwon, Andrew Perri, and Jocelyn Walker). Financial support was provided by the National Science and Engineering Research Council (NSERC), Alberta Innovates, the Alberta Government and the University of Alberta for scholarships to TJW and the Alberta Crop Industry Development Fund (ACIDF) and Landscape Alberta Nursery Trades Association (LANTA) for their generous grants to MLE. I thank Calvin and Gloria Sutela for their tireless hospitality that I needed to complete this degree. Finally, thank you to my wonderful wife and family for their sacrifice and patience during this chapter of our lives. v Table of Contents Abstract ..................................................................................................................................ii Preface ................................................................................................................................... iv Acknowledgements ................................................................................................................. v List of Tables ......................................................................................................................... xi List of Figures ......................................................................................................................xiii List of Abbreviations ........................................................................................................... xvii Chapter 1: Introduction ........................................................................................................... 1 Herbivore host plant preference and offspring performance ................................................. 1 Odour mediated host location by herbivorous insects ........................................................... 2 Host location by lepidopterans ............................................................................................. 3 Tritrophic interactions ......................................................................................................... 8 Chemically-mediated host location by parasitoids ................................................................ 9 VOC-mediated differential parasitism................................................................................ 12 The tritrophic system ......................................................................................................... 13 First trophic level: Ash trees (Fraxinus spp.) .................................................................. 13 Second trophic level: Caloptilia fraxinella ..................................................................... 14 Third trophic level: Parasitoids, with a focus on Apanteles polychrosidis ....................... 16 Thesis overview and objectives ......................................................................................... 18 Chapter 2: The volatile organic compound host location cues of Caloptilia fraxinella ............ 46 Introduction .......................................................................................................................... 46 Microscopic analysis of sensilla......................................................................................... 48 Volatiles from Fraxinus spp. ............................................................................................. 48 Ash leaf cone-roller ........................................................................................................... 49 Objectives ......................................................................................................................... 50 Materials and Methods .......................................................................................................... 50 vi Field sites .......................................................................................................................... 50 Insects ............................................................................................................................... 50 Plant material .................................................................................................................... 51 Microscopic analysis of sensilla......................................................................................... 51 Detection