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Facultative Group Living in the Western Black Widow Spider, Latrodectus Hesperus: an Evolutionary Approach

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Facultative Group Living in the Western Black Widow Spider, Latrodectus Hesperus: an Evolutionary Approach FACULTATIVE GROUP LIVING IN THE WESTERN BLACK WIDOW SPIDER, LATRODECTUS HESPERUS: AN EVOLUTIONARY APPROACH by Maxence F. Salomon M.Sc. Swiss Tropical Institute, 2000 B.Sc. University ofGeneva, 1999 THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY In the Department of Biological Sciences © Maxence Salomon 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 ofthe author. APPROVAL Name: Maxence F. Salomon Degree: Doctor of Philosophy Title ofThesis: Facultative group living in the western black widow spider, Latrodectus hesperus: an evolutionary approach Examining Committee: Chair: Dr. R.A. Nicholson, Associate Professor Dr. B.D. Roitberg, Professor, Senior Supervisor Department of Biological Sciences, S.F.U. Dr. A.S. Harestad, Professor Department of Biological Sciences, S.F.U. Dr. R.G. Bennett, Seed Pest Management Officer B.C. Ministry of Forests Dr. W.P. Maddison, Professor Departments of Zoology and Botany, University of British Columbia Public Examiner Dr. I.A. Stamps, Professor Emeritus Section of Ecology and Evolution, University of California, Davis External Examiner 4 April 2008 Date Approved ii 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 A major goal ofevolutionary research is to elucidate the processes involved in the evolution ofgroup versus solitary living, by examining the selective forces driving or constraining a particular type ofsocial system. Species with intermediate group living tendencies are particularly interesting because they offer insight into evolutionary transitions and, in particular, how different selective environments may modulate group living behaviours. This dissertation explores the different factors that shape the social strategies ofthe western black widow spider, Latrodectus hesperus, a facultatively group­ living spider. Over a period offour years, I characterized the social structure of coastal British Columbia populations of1. hesperus, and show that individuals live either solitarily or in groups depending on the time ofyear, their reproductive status, and age. I then used an experimental approach to investigate the dynamics and adaptive value of facultative group-living behaviours, by testing different hypotheses about the decisions involved in social interactions, web building, microhabitat settlement, movement, and foraging. Several factors were manipulated and shown to influence the patterns of group living in 1. hesperus, including individual nutritional state, prey availability, group size, population density, and neighbour proximity. Spiders adjusted their group living behaviours according to changes in these factors, and responded strategically to the presence and proximity of conspecifics. Based on the results ofthese experiments, I developed a model ofgroup living in spiders that specifically considers the dynamic and strategic nature ofsocial interactions in the context offrequency-dependent selection. The research presented in this thesis furthers our understanding ofthe evolution of social behaviour by providing new evidence on the mechanisms that promote and regulate facultative group-living behaviours. Keywords: aggregation; foraging; group living; social; spiders; web building iii ACKNOWLEDGEMENTS First and foremost, I thank Bernie Roitberg for giving me the opportunity to conduct this project and for being an outstanding supervisor. His enduring trust and mentorship allowed me to flourish as a researcher and find my way through challenging times, and he was inspirational. I also thank Robb Bennett for his keen support and mentorship, for sharing his passion of spider biology, and for welcoming me into his home. I thank Alton Harestad for all ofhis feedback and advice throughout this project and for his enthusiasm. I also thank Judy Stamps and Wayne Maddison for reading my dissertation and providing a lot of constructive criticism and ideas. Samantha Vibert provided invaluable help and advice during all phases ofthis project and her contribution was instrumental to the success ofmost ofthis research, as was her close friendship. I also thank all the members ofthe Roitberg lab from 2002­ 2008 for providing such a stimulating research environment. I thank Clea Moray for patiently sharing her knowledge ofscientific writing and computer programming. I thank Gwylim Blackburn for a lot ofhelpful discussions, and also good times in the field. The Bennett-Terpenning family have improved my quality oflife by providing generous hospitality and much support during deamnding times. I thank Sarbjeet Mann for her patience and love, which no doubt helped me complete this project. I thank Nick Charette, Angela Mah, Carolyn Duckham, Nadia Brinkman, Melanie Hart, Coltilde Salomon, Jason Peterson and Gwylim Balckburn for assistance in the field and laboratory. I also thank numerous co-workers and friends for intellectual input and help, namely Jay Biernaskie, Richard Ring, Bruce Leighton, David Green, Laurie Marczak, Deepa Pureswaran, Dave Gillespie, Ian Gordon, Peter Midford, Gerta Moray, Gerhard and Regine Gries, various members ofthe Gries lab, Larry Dill, Heather Higo, IV Scott Pavey, Peter Belton, Dave Raworth, Kefyn Catley, Carl Schwarz, Nadia Sartoretti, Nick Page, Patricio Salazar and Esperanza, Patrik Nosi!, Marcela Olguin, Andres Valle­ Domenech, the BISC technicians, various TA instructors and fellow TAs, the BISC office staff, especially Marlene Nguyen, and other people in BERG and BISC whom I am surely forgetting. I thank former housemates and flatmates for all the good times we had: Scott Pavey, Tom Cowan, Marie-Helene Burle, Frankie Doodle, Graham and Pam Humphreys, and Anders Odeen. Finally, I am grateful to the Blackburn family ofEsquimalt, the Mann family of Duncan, and the Johnston family ofCrofton for their oustanding hospitality. I also thank the Tsawout First Nation for allowing me to conduct my research on their beautiful land on the Saanich Peninsula. Funding was provided by NSERC, SFU, the B.c. Ministry of Forests, the Entomological Society of Canada, the Entomological Society ofBritish Columbia, and the Highlands Biological Station. v TABLE OF CONTENTS Approval ii Abstract iii Acknowledgements iv Table of Contents vi List of Figures viii List of Tables xii Chapter 1: General Introduction 1 1.1 General Context ofThesis Research 2 1.2 Focus ofThesis Project and Definitions 4 1.3 Overview ofThesis Chapters 5 1.4 References 6 Chapter 2: Microhabitat use by Latrodectus hesperus spiders in coastal British Columbia: evidence of facultative group living * 9 2.1 Abstract 10 2.2 Introduction 10 2.3 Materials and Methods 13 2.4 Results 19 2.5 Discussion 27 2.6 Acknowledgements 34 2.7 References 34 2.8 Appendices 45 Chapter 3: Cohabitation dynamics among web-building spiders as a function of prey availability and population density * 50 3.1 Abstract 51 3.2 Introduction 51 3.3 Materials and Methods 55 3.4 Results 63 3.5 Discussion 73 3.6 Acknowledgements 79 3.7 References 79 Chapter 4: Combined influence of neighbours and feeding state on movement decisions made by a web-building spider 91 4.1 Abstract 92 4.2 Introduction 92 4.3 Experiment 1: Microhabitat Tenacity in Established Spiders 95 vi 4.4 Experiment 2: Effect ofNeighbours on Microhabitat Residence Time 97 4.5 Experiment 3: Effects ofMultiple Factors on Microhabitat Residence
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