PHEROMONE-BASED ARRESTMENT BEHAVIOUR OF THREE SPECIES OF THYSANURA (LEPISMATIDAE) by Nathan Woodbury BSc, Simon Fraser University 2000 THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF PEST MANAGEMENT In the Department of Biological Sciences © Nathan Woodbury 2008 SIMON FRASER UNIVERSITY Spring 2008 All rights reserved. This work may not be reproduced in whole or in part, by photocopy or other means, without permission of the author. APPROVAL Name: Nathan Woodbury Degree: Master of Pest Management Title of Thesis: Pheromone-Based Arrestment Behaviour of Three Species of Thysanura (Lepismatidae) Examining Committee: Chair: Dr. J. Reynolds, Professor, S.F.U. Dr. G. Gries, Professor, Senior Supervisor Department of Biological Sciences, S.F.U. Dr. C. Lowenberger, Associate Professor Department of Biological Sciences, S.F.U. Dr. G. Judd, Research Scientist, Entomologist Pacific Agri-food Research Center, Summerland, B.C. Date Approved: APRIL 07 7000 11 SIMON FRASER UNIVERSITY LIBRARY Declaration of Partial Copyright Licence The author, whose copyright is declared on the title page of this work, has granted to Simon Fraser University the right to lend this thesis, project or extended essay to users of the Simon Fraser University Library, and to make partial or single copies only for such users or in response to a request from the library of any other university, or other educational institution, on its own behalf or for one of its users. The author has further granted permission to Simon Fraser University to keep or make a digital copy for use in its circulating collection (currently available to the public at the "Institutional Repository" link of the SFU Library website <www.lib.sfu.ca> at: <http://ir.lib.sfu.ca/handle/1892/112>) and, without changing the content, to translate the thesis/project or extended essays, if technically possible, to any medium or format for the purpose of preservation of the digital work. The author has further agreed that permission for multiple copying of this work for scholarly purposes may be granted by either the author or the Dean of Graduate Studies. It is understood that copying or publication of this work for financial gain shall not be allowed without the author's written permission. Permission for public performance, or limited permission for private scholarly use, of any multimedia materials forming part of this work, may have been granted by the author. This information may be found on the separately catalogued multimedia material and in the signed Partial Copyright Licence. While licensing SFU to permit the above uses, the author retains copyright in the thesis, project or extended essays, including the right to change the work for subsequent purposes, including editing and publishing the work in whole or in part, and licensing other parties, as the author may desire. The original Partial Copyright Licence attesting to these terms, and signed by this author, may be found in the original bound copy of this work, retained in the Simon Fraser University Archive. Simon Fraser University Library Burnaby, BC, Canada Revised: Fall 2007 ABSTRACT Aggregations of the common silverfish, Lepisma saccharina L., giant silverfish, Ctenolepisma longicaudata (Escherich), and firebrat, Thermobia domestica (Packard), are mediated by non-volatile, species-specific pheromones. In dual-choice olfactometer experiments, filter paper previously exposed to male, female, or juvenile L. saccharina or C. longicaudata arrested conspecifics regardless of developmental stage or gender. Lepisma saccharina did not respond to the C. longicaudata pheromone, nor to the T. domestica pheromone. However, C. longicaudata responded to the pheromones of both L. saccharina and T. domestica, whereas T. domestica responded to the C. longicaudata but not L. saccharina pheromone. Female T. domestica were significantly arrested by (i) loose, insect-derived debris brushed from shelters, (ii) a frass mixture manually separated from loose debris, and (iii) specific amber-type frass manually separated from the frass mixture, but did not respond to other types of shelter debris or insect-altered cellulose, suggesting that T. domestica pheromone is present in amber-type frass. Keywords: Lepisma saccharina, Ctenolepisma longicaudata, Thermobia domestica, Thysanura, Zygentoma, Lepismatidae, pheromone, aggregation, arrestment, frass, feces. III ACKNOWLEDGEMENTS I would like to thank my senior supervisor, Dr. Gerhard Gries, for his invaluable advice, patience and endless enthusiasm throughout the course of this (still ongoing) work. In addition, Mrs. Regine Gries, Dr. Grigori Khaskin, Dr. Robert Britton and my Gries Laboratory colleages deserve heartfelt thanks for helping me surmount many scientific obstacles. I thank my surpervisory committee member, Dr. Carl Lowenberger for review of my thesis and for constructive suggestions during the course of this research. I also thank the public examiner, Dr. Gary Judd for his valuable suggestions which have lead to many improvements of this thesis. I would also like to thank Dave Booth for supplying Lepisma saccharina, Michelle Tremblay for advice on the capture and rearing of all thysanuran species, Eberhard Kiehlmann for contructive comments on manuscripts, and Colin Zhang, Aleksander Miroshnychenko, Allen Haddrell and Michael Katz for all of their help with chemical analyses. I also thank Kate McLellan, Landon Woodbury, my parents and my friends for years of unfaltering support and wisdom. This research was supported by a Professor Thelma Finlayson Fellowship and a Simon Fraser University Graduate Fellowship to N.W., and a Natural Sciences and Engineering Research Council of Canada (NSERC)-Industrial IV Research Chair to G.G. with SC Johnson Canada, Pherotech International Inc. and Global Forest Science (GF-18-2007-226 and GF-18-2007-227) as the industrial sponsors. v TABLE OF CONTENTS Approval ii Abstract iii Acknowledgements iv Table of Contents vi List of Figures viii 1 THYSANURAN BIOLOGY AND AGGREGATION OF PHYLOGENETICALLY ANCESTRAL INSECTS 1 1.1 THYSANURAN BIOLOGY & ECOLOGY 1 1.1.1 Phylogeny, Distribution and Morphology of Thysanura 1 1.1.2 Abiotic Conditions and Food Preferences of Thysanura 5 1.1.3 Mating Behaviour & Development of Thysanura 7 1.2 AGGREGATION BEHAVIOUR OF PHYLOGENETICALLY ANCESTRAL INSECTS 9 1.2.1 Aggregation Behaviour of Thysanura 9 1.2.2 The Role of Pheromones in Aggregations of Hexapods and Orthopteroid Insects 12 1.2.3 The Role of Pheromones in Thysanuran Aggregations .15 1.3 THE ECONOMIC IMPORTANCE OF THYSANURA 16 1.4 RESEARCH OBJECTIVES 17 2 EVIDENCE FOR AN ARRESTMENT PHEROMONE IN L. SACCHARINA AND C. LONGICAUDATA 19 2.1 INTRODUCTION 19 2.2 MATERIALS AND METHODS 21 2.2.1 Collecting and Rearing of Experimental Insects 21 2.2.2 General Bioassay Procedures 22 2.2.3 Specific Experiments 23 2.2.4 Statistical Analyses 25 2.3 RESULTS 25 vi 2.4 DiSCUSSiON 37 3 SHELTER DEBRIS AND THE ARRESTMENT PHEROMONE OF T. DOMESTICA 41 3.1 INTRODUCTION 41 3.2 MATERIALS AND METHODS 44 3.2.1 Collecting and Rearing of Experimental Insects .44 3.2.2 General Bioassay Procedures .45 3.2.3 Specific Experiments .46 3.2.4 Statistical Analyses 50 3.3 RESULTS 50 3.4 DiSCUSSiON 54 4 CONCLUSiON 60 References 61 vii LIST OF FIGURES Figure 1.1 Photographic illustrations of the common silverfish, Lepisma saccharina (top), giant or long-tailed silverfish, Ctenolepisma longicaudata (middle), and the firebrat, Thermobia domestica (bottom) 3 Figure 2.1 Number of male, female, or juvenile L. saccharina responding to a piece of filter paper previously exposed to male, female, or juvenile L. saccharina. Numbers near bars indicate the number of insects responding to the test stimulus. An asterisk (*) indicates a significant preference for a particular test stimulus (X2 test; *P :5 0.05, **P :5 0.01). Numbers in brackets indicate numbers of non-responding insects 27 Figure 2.2 Number of male, female, or juvenile C. longicaudata responding to a piece of filter paper previously exposed to male, female, or juvenile C. longicaudata. Numbers near bars indicate the number of insects responding to the test stimulus. An asterisk (*) indicates a significant preference for a particular test stimulus (X2 test; *P :5 0.05, **P :5 0.01). Numbers in brackets indicate numbers of non-responding insects 29 Figure 2.3 Number of female L. saccharina (left) or C. longicaudata (right) responding to a piece of filter paper previously exposed to male, female, or juvenile conspecifics when stimulus contact was prohibited or allowed. Numbers near bars indicate the number of insects responding to the test stimulus. An asterisk (*) indicates a significant preference for a particular test stimulus (l test; *P :5 0.05, **P :5 0.01). Numbers in brackets indicate numbers of non-responding insects 31 Figure 2.4 Number of female L. saccharina responding to conspecific frass, scales, antennae and caudal filaments, or salivary gland contents. Numbers near bars indicate the number of insects responding to the test stimulus. An asterisk (*) indicates a significant preference for a particular test stimulus (x2 test; *P:5 0.05, **P :5 0.01). Numbers in brackets indicate numbers of non-responding insects 33 viii Figure 2.5 Number of female L. saccharina, C. longicaudata, or T. domestica responding to a piece of filter paper previously exposed to conspecific or heterospecific males, females, and juveniles. Numbers near bars indicate the number of insects responding to the test stimulus. An asterisk (*) indicates a significant preference for a particular test stimulus (X2 test; *P ~ 0.05, **P ~ 0.01). Numbers in brackets indicate numbers of non-responding insects 35 Figure 3.1 Number of female T. domestica (A) responding to shelter paper exposed to conspecifics for 3 days (experiment 1) or insect-derived debris accumulating in those shelters; and (8) responding to insect-exposed paper shelter tips, paper shelter perimeters, glass surfaces exposed to conspecifcs for 3 days, silk removed from shelters, or silk squeezed from male T.