Behaviour and Life History of a Large Carpenter Bee (Xylocopa Virginica) in The

1 Behaviour and Life History of a Large Carpenter Bee (Xylocopa virginica) in the Northern Extent of its Range by Sean Michael Prager, B.A. A Thesis submitted to the Department of Biological Sciences in partial fulfilment of the requirements for the degree of Doctor of Philosophy Brock University St. Catharines, Ontario © Sean M. Prager, 2008 Library and Bibliotheque et 1*1 Archives Canada Archives Canada Published Heritage Direction du Branch Patrimoine de I'edition 395 Wellington Street 395, rue Wellington Ottawa ON K1A0N4 Ottawa ON K1A0N4 Canada Canada Your file Votre reference ISBN: 978-0-494-46634-6 Our file Notre reference ISBN: 978-0-494-46634-6 NOTICE: AVIS: The author has granted a non L'auteur a accorde une licence non exclusive exclusive license allowing Library permettant a la Bibliotheque et Archives and Archives Canada to reproduce, Canada de reproduire, publier, archiver, publish, archive, preserve, conserve, sauvegarder, conserver, transmettre au public communicate to the public by par telecommunication ou par Plntemet, prefer, telecommunication or on the Internet, distribuer et vendre des theses partout dans loan, distribute and sell theses le monde, a des fins commerciales ou autres, worldwide, for commercial or non sur support microforme, papier, electronique commercial purposes, in microform, et/ou autres formats. paper, electronic and/or any other formats. The author retains copyright L'auteur conserve la propriete du droit d'auteur ownership and moral rights in et des droits moraux qui protege cette these. this thesis. Neither the thesis Ni la these ni des extraits substantiels de nor substantial extracts from it celle-ci ne doivent etre imprimes ou autrement may be printed or otherwise reproduits sans son autorisation. reproduced without the author's permission. In compliance with the Canadian Conformement a la loi canadienne Privacy Act some supporting sur la protection de la vie privee, forms may have been removed quelques formulaires secondaires from this thesis. ont ete enleves de cette these. While these forms may be included Bien que ces formulaires in the document page count, aient inclus dans la pagination, their removal does not represent il n'y aura aucun contenu manquant. any loss of content from the thesis. Canada 2 Abstract Large carpenter bees (Hymenoptera: Apidae: Xylocopa) have traditionally been thought of as exhibiting solitary or occasionally communal colony social organization. However, studies have demonstrated more complex forms of social behaviour in this genus. In this document, I examine elements of behaviour and life history in a North American species at the northern extreme of its range. Xylocopa virginica was found to be socially polymorphic with both solitary and meta-social or semi-social nests in the same population. In social nests, there is no apparent benefit from additional females which do not perform significant work or guarding. I found that the timing of life-history events varies between years, yet foraging effort only differed in the coldest and wettest year of 2004 the study. Finally, I that maleX. virginica exhibit female defence polygyny, with resident and satellite males. Resident males maintain their territories through greater aggression relative to satellites. 3 Acknowledgements A project like this does not happen without the assistance of those who choose to contribute some of their intellectual fitness. I only hope they received some indirect fitness. Fiona Hunter was amazing in fighting for me and getting me to defence, I could not have done this without your aid. Jean Richardson provided advice, consul and many hours of data checking. To the other members of my committee: Jeff Stuart, Jon Middleton, Miriam Richards and early on Bob Caroline, your help is much appreciated. I would especially like thank the external reviewer for this thesis; Dr. Jon Barthell not only contributed his time and comments but also donated his bees. Numerous field assistants helped observe and mark bees, without them this would have been difficult. Thank you to the machine shop for lending me tools and to physical plant for your varied assistance. Sandra Rehan and Amy Rutgers-Kelly, you both started as lab mates and have become great friends and comrades. Candice Kerling, there are countless ways in which I appreciate you and your karate kicking self. I would like to that thank all those who have helped me by reading drafts. I would like to thank my friends; Canadian, American and other, you are all great. Finally, I would like to thank all the members of my family, immediate and extended, every one of you has contributed to this in some way and I never could have done this without you. 4 Table of Contents Abstract 2 Acknowledgements 3 Table of Contents..... 4 List of Tables 8 List of Figures 9 General Introduction 11 Hypothesis to explain the evolution of altruism 12 Kin selection 12 Mutualism 14 Queen control 14 Social organization in insects 15 Evolution of eusociality in insects 17 Reproductive skew models 18 Subfamily Xylocopinae 20 Tribe Xylocopini 21 Genus Xylocopa 22 Subgenus Xylocopoides 22 Xylocopa virginica virginica 23 Aims of this research 24 General background and methods 26 Study sites-St. Catharines 26 Phenology and life cycle 28 Bees obtained from other sources 30 Destructive sampling of nests and collection of contents 31 Paint marking for identification 33 Videoscope observations of nests 34 Observations of females at nest entrances 35 Terminology 36 Chapter One: Foundress associations in Xylocopa (Xylocopoides) virginica 37 Introduction 37 Methods 39 Nest contents and productivity 39 Brood parasites 40 Foundress survival and colony longevity 41 Behavioural roles of females 42 Comparisons of females from multiple foundress nests 46 Statistical analyses 49 Results 50 Nest contents and productivity 50 Foundress survival and colony longevity 56 Behavioural roles of females 57 Comparisons of females from multiple-foundress nests 70 Behavioural comparisons 75 Discussion 80 5 The social organization of Xylocopa virginica 80 Adaptive significance of solitary versus multi-foundress nesting in Xylocopa virginica 83 Broader implications ofXylocopa virginica social structure 86 Conclusions 88 Chapter Two: Relationships between nest architecture and behaviour in Xylocopa virginica 90 Introduction 90 Methods 93 Study sites 93 Nest architecture as it relates to brood laying order and size 93 The relationship between nest architecture and sex allocation 94 Relationships between nest architecture and parasitism 95 Relationships between nest architecture and number of females 96 Sample availability considerations 97 Results 97 Relationships between nest architecture and brood 97 The relationship between nest architecture and sex allocation 98 Relationships between nest architecture and parasitism 98 Relationship between nest architecture and number of females 103 Discussion 106 Brood development and nest architecture 106 Relationships between nests and sex allocation 107 Relationships between parasitism and nesting 108 Relationships between nests and social structure 109 Conclusions Ill Chapter Three: Annual variation in demography and activity of Xylocopa virginica in southern Ontario 112 Introduction 112 Methods 115 Study sites 115 Phenology and life cycle of Xylocopa virginica 115 Demographic indicators of phenology 116 Analyses and quantification of weather and climate trends 117 Relationships between weather and bee phenology 124 Evaluation of female foraging effort 126 Annual variation in morphological size 128 Results 128 Phenology and life cycle ofXylocopa virginica 128 Demographic indicators of phenology 132 Relationships between weather and bee phenology 137 Evaluation of female foraging effort 137 Annual variation in morphological size 139 Discussion 142 Annual variation in phenology 142 Geographic variation in life cycle and phenology 143 6 Analyses of foraging effort 143 Weather and nest re-use 146 Taxonomic variation in phenology 147 Consequences of poor weather 148 Conclusions 149 Chapter 4: Male behaviour, mating tactics and aggression in association with female traits in the large carpenter bee, Xylocopa virginica 150 Introduction 150 Methods 153 Study Site 153 Marking for identification 153 Behavioural observations and censuses 153 Analytical Methods 157 Results 162 Evidence for polygyny 162 Evidence of territoriality, resident and satellite males 162 Territorial vs. satellite male behaviour 169 Male-female interactions 171 Characteristics of territory quality 172 Discussion 177 Mating system in Xylocopa virginica 177 Behavioural differences between residents and satellites 178 Male-female interactions 179 Territory quality 181 Mating strategies in Hymenoptera 181 Conclusions 184 General Discussion and Conclusions 185 Associations between males and females 185 Anomalous 2004 season 185 Sociality in Xylocopa 186 A proposed sequence for the evolution of sociality in Xylocopa 189 Further Research 193 Literature Cited 195 Appendices 208 Appendix 10: A descriptive and comparative study of nest architecture in Xylocopa virginica .218 Introduction 218 Methods 218 Study sites, nesting site and substrate selection 218 Evaluation of nest architecture 220 Results 223 Nesting site and substrate selection 223 Nest architecture 226 Geographic variation 231 Discussion 234 Nest site and substrate selection 234 7 Factors influencing nest architecture 236 Geographical variation in nest architecture 237 Interspecific variation in nest architecture in the genus Xylocopa

Load more