Diptera: Anthomyiidae)
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ECOLOGICAL AND PHYSIOLOGICAL FACTORS AFFECTING MATE-FINDING AND MATING BEHAVIOUR OF DELIA ANTIQUA (MEIGEN) (DIPTERA: ANTHOMYIIDAE) R. S. MCDONALD B.Sc., University of Lethbridge, 1977 M.Sc. Universrty of Guelph, 1986 Post Baccularate Diploma, Simon Fraser University, 1991 THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in the Department of Biological Sciences Q Robert Stuart McDonald 1995 SIMON FRASER UNIVERSITY August 1995 All rights reserved. This work may not be reproduced in whole or in part, by photocopy or other means, without permission of the author. Frontispiece: Mating sequence of Delia antiqua (Meigen), (Diptera: Anthomyiidae) APPROVAL NAME: Robert Stuart McDonald DEGREE: DOCTOR OF PHILOSOPHY TITLE OF THESIS: ECOLOGICAL AND PHYSIOLOGICAL FACTORS AFFECTING MATE-FINDING BEHAVIOUR OF DELIA ANTIQUA (MEIGAN) (DIPTERA: ANTHOMYIIDAE) Examining Committee: Chair: Dr. M. Mackauer, Professor Dr. J.'Borden, Professor, Senior Supervisor, Department of Biological Sciences, SFU A Dr. WHaunerland, Associate Professor Department of Biological Sciences, SFU Dr. H. Pierce jr., Research Associate Department of Chemistry, SFU //L --I -- Dr. R.Vernon, Research Scientist Agriculture Canada Public examiner r~iller;kdftkor ' Department of Entomology, Michigan State University VExternal Examiner PARTIAL COPYRIGHT LICENSE I hereby grant to Simon Fraser University the right to lend my thesis, project or extended essay (the title of which is shown below) to users of the Simon Fraser Univeristy Library, and to make partial or single copies only for such users or in response to a request from the library of any other univerisrty, or other educational institution, on its own behalf or for one of its users. I further agree that permission for multiple copying of this work for scholarly purposes may be granted by me of the Dean of Graduate Studies. It is understood that copying or publication of this work for financial gain shall not be allowed without my written permission. Title of ThesisfProjectJExtended Essay: Ecological and Physiological Factors Affecting Mate-finding and Mating Behavlour of Delia anfiqua (Meigen) (Diptera: Anthomyiidae) Author: (signature) Robert Stuart McDonald (name) August 14, 1995 (date) ABSTRACT Adult onion flies, Delia antiqua (Meigen), were studied from eclosion to sexual maturity, to assess the impact of ecological and phys~ologicalfactors on mate-finding, and mating behaviour. Protandry (male eclosion in advance of females) resulted from disparate pupal development times between the sexes. The postulation that protandry is an adaptive strategy providing reproductive benefits to either sex is equivocal for D. antiqua because eclosion curves of pupae were normally distributed and mean time-lags in eclosion peaks between the sexes were too short (<4 d) to optimize any reproductive benefits from early sexual maturation or mating. The alternative hypothesis is presented that protandry is a non-adaptive outcome of selection for other life history traits that is unbiased by female size dimorphism. Mating behaviour was age- dependent for both sexes. Few males or females were sexually mature at 3-4 d post edosion, however, >50% of adults aged 6-7 d mated. Although at this time, oocytes were normally 150% of their final egg volume, ovarian development was only weakly correlated (r = 0.48) with frequency of mating. Previtellogenic females, however, were rarely inseminated, suggesting a dietary association with sexual receptivity. In contrast, sucrose feeding by adult males over 10 d had no effect on the proportion of gravid females inseminated over 24 h (-5 females per male), the magnitude of the ovipositional response, or the total numbers of eggs deposited in comparison with males fed protein-rich diet. Upwind response by virgin females aged 410 d to onion dour in a wind tunnel was comparable to that of gravid, mated 10-d-old females, but male upwind response increased linearly with age (? = 0.98), and was inhibited in sexually mature males by the presence of mature females in the wind tunnel. In the absence of host dour, males, but not females, were attracted upwind to conspecifics. Since upwind response to onion dour was independent of ovarian development or mating status, the host-plant probably serves to situate females to their oviposition sites, and males near females. Male courtship and mating behaviour fdlowed a sequence that apparently involves visual recognition of a potential mate, and then species- and sex-specific chemical recognition. Solvent extracts of cuticular volatiles from mature, gravid females, elicited significantly more contact and mating attempts than extracts from immature females or sobent controls. Unique, age-related, C28 methyl-branched alkanes in the extracts of sexually mature females may act as a contact recognition pheromone. An understanding of age-related phenomena affecting mate-finding and mating behaviour under laboratory conditions may enhance the management of D. antiqua in commercial onion production if D. antiqua adopt a land-mark based, male-controlled mating system in nature. Integration of chernosensory cues from the host-plants and sex pheromone could lead to improved techniques for monitoring or attracting adults. With incorporation of control measures, such synthetic cues could potentially disrupt mate-finding and mating, especially to the spring generation of aduRs, when competition from naturally-occurring attractants is lowest. DEDICATION To E. ti., J. A. and S. ACKNOWLEDGMENTS I wish to express my appreciation to my senior supervisor John H. Borden for generous support and assistance during the course of study and preparation of this thesis. Furthermore, I am especially grateful to the following technical staff at Simon Fraser University: Akbar Syed (Department of Biological Sciences) for providing consistently high quality D. antiqua and rearing supplies on demand; the late Roger Turner and Les Wakida (Science Technical Centre) for construction and modifications to the design of the wind tunnel and glassware, respectively; Greg Owen (Department of Chemistry) for mass spectrometry; Elizabeth K. Carefoot (Instructional Media Centre) for preparation of the frontispiece; and Victor Bourne (Biological Sciences) for assistance with photography. Jay Whistlecraft (Research Branch, Agriculture Canada, London, Ont.) generously provided cultures of D. antiqua for comparison during the course of this study. The technical assistance of undergraduate students Michelle A. Thon, Joanne Scherba, Christa Scott, Carmen Pon, and Paul Mah during the development stages of many bioassays, and referencing much of the literature for the appendix is most appreciated. I also thank my friends and colleagues of the Chemical Ecology Research Group and environs surrounding 86220, whose affabilrty made my time here so enjoyable. Finally, I would especially like to acknowledge my committee members, in particular Harold D. Pierce Jr. (Department of Chemistry) who provided technical advice and assistance for many of the pheromonal investigations and James R. Miller (Michigan State University, East Lansing, MI) and Robert S. Vernon (Research Branch, Agriculture Canada, Vancouver, B.C.) for critical review of the thesis. This study was supported in part by a grant from the Natural Sciences and Engineering Research Council of Canada. TABLE OF CONTENTS Page APPROVAL ............................................................................................. ii ABSTRACT .............................................................................................. iii DEDICATION ........................................................................................... v ACKNOWLEDGEMENTS .......................................................................... vi TABLE OF CONTENTS .......................... ................................................. vii LIST OF TABLES ...................................................................................... xi LIST OF FIGURES .................................................................................. xii 1. INTRODUCTION ......................................................................... 1 1.1 BIOLOGY AND MANAGEMENT OF DELIA ANTIQUA ..................... 2 1.2 THESIS OBJECTIVES ................................................................... 8 2. THE ROLE OF PROTANDRY AS AN ADAPTIVE REPRODUCTIVE STRATEGY INTRODUCTION ...................... .. ...... .. .................................... 10 MATERIALS AND METHODS ..................................................... 13 RESULTS ................................................................................... 16 DISCUSSION ............................................................................. 32 vii 3 . THE RELATIONSHIP OF AGE AND OVARIAN DEVELOPMENT TO MATING 3.1 INTRODUCTION ............................. .. .................................... 3.2 MATERIALS AND METHODS ..................................................... 3.3 RESULTS ................................................................................... 3.4 DISCUSSION ............................................................................. 4 . DIETARY CONSTRAINTS ON SEXUAL RECEPTIVITY. MATING SUCCESS. AND MALE SURVNORSHIP 4.1 INTRODUCTION ................................ ...................................... 4.2 MATERIALS AND METHODS ............................................... 4.3 RESULTS ..................................................................................