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Progress in the Producer-Scrounger Game : Information Use and Spatial Models

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Progress in the Producer-Scrounger Game : Information Use and Spatial Models UNNERSITÉ DU QUÉBEC À MONTRÉAL PROGRESS IN THE PRODUCER-SCROUNGER GAME: INFORMATION USE AND SPATIAL MODELS THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIRElVIENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN BIOLOGY BY STEVEN HAMBLIN SEPTEMBER 2011 UNIVERSITÉ DU QUÉBEC À MONTRÉAL Service des bibliothèques Avertissement La diffusion de cette thèse se fait dans le respect des droits de son auteur, qui a signé le formulaire Autorisation de reproduire et de diffuser un travail de recherche de cycles supérieurs (SDU-522 - Rév.01-2006). Cette autorisation stipule que «conformément à l'article 11 du Règlement no 8 des études de cycles supérieurs, [l'auteur] concède à l'Université du Québec à Montréal une licence non exclusive d'utilisation et de publication de la totalité ou d'une partie importante de [son] travail de recherche pour des fins pédagogiques et non commerciales. Plus précisément, [l'auteur] autorise l'Université du Québec à Montréal à reproduire, diffuser, prêter, distribuer ou vendre des copies de [son] travail de recherche à des fins non commerciales sur quelque support que ce soit, y compris l'Internet. Cette licence et cette autorisation n'entraînent pas une renonciation de [la] part [de l'auteur] à [ses] droits moraux ni à [ses] droits de propriété intellectuelle. Sauf entente contraire, [l'auteur] conserve la liberté de diffuser et de commercialiser ou non ce travail dont [il] possède un exemplaire.» UNIVERSITÉ DU QUÉBEC À MONTRÉAL PROGRÈS DANS LE JEU PRODUCTEUR-CHAPARDEUR: L'UTILISATION DES INFORMATIONS ET DES MODÈLES SPATIAUX THÈSE PRÉSENTÉE COMME EXIGENCE PARTIELLE DU DOCTORAT EN BIOLOGIE PAR STEVEN HAMBLIN SEPTEMBRE 20 Il Dedicated to Brandy and Dawn. One saw the end ofthis and one didn't, but if wouldn't existed without both. FOREWORD In this doctoral thesis, five chapters are presented as scientific articles. The first ar­ ticle (Chapter II) was published in Animal Behaviour (Vol. 78, pp. 1343-1350) un­ der the title "Finding the evolutionarily stable learning mie for freguency-dependent foraging". This article was co-authored by Steven Hamblin and Luc-Alain Giraldeau. Chapter III is an article published in Oikos (Vol. 119, pp. 286-291) entitled "Predator inadvertent social information use favors reduced clumping of its prey", and was co-authored by Steven Hamblin, Kimberley Mathot, Julie Morand-Ferron, Joseph Nocera, and Luc-Alain Giraldeau. In Chapter IV is the article "Scroungers are shy and producers are either: The co-evolution of boldness and foraging tactics", co­ authored by Ralf Kurvers, Steven Hamblin, and Luc-Alain Giraldeau and submitted ta the journal Behavioral Ecology. Chapter V contains the article "A mie of thumb for social foraging", co-authored by Steven Hamblin and Luc-Alain Giraldeau and submitted ta The American Naturalist. FinaIly, Chapter VI is a single-author article by Steven Hamblin in preparation for journal submission (e.g. to Proceedings ofthe Royal Academy, Series B or Behavioral Ecology). During the time of this thesis, Steven Hamblin was supported by an NSERC CGS­ D scholarship as weIl as une bourse d'exemption des frais majorés from UQAM, and from an NSERC Discovery grant ta Luc-Alain Giraldeau. viii Acknowledgements It takes a village ta raise a child. African Proverb My growth as a scientist has been shaped in both subtle and spectacular ways by many people; my successes are shared with them, but my failures are my own. Among the greatest influences on this work is my supervisor, Luc-Alain Giraldeau. The circumstances of my degree have been at times peculiar, but his support has never wavered and he never stapped pushing me to look at biological questions as a behavioural ecologist would; 1believe that this has made me a better biologist. His financial support kept me going for the first two years of the degree, and 1appreciate the support for conference travel that he has given me (particularly the opportunity to attend the MEl conference in Ohio). 1have learned more than 1could have hoped under his supervision. 1 would also like to thank my Master's advisor, Pete Hurd, for giving me a place to work when 1was in Edmonton and for invaluable discussion along the way. Pete started me on the path l've taken, and l'Il always be grateful for his help. Along the way, many people were gracious enough to help me with various stages of the Ph.D. process. Bill Vickery and Pedro Peres-Neto provided valuable feedback on my Ph.D. proposai, and 1 want to thank Pedro as weil as Louis Lefebvre for be­ ing a part of my examen de synthèse committee. Ginette Lozeau, François Dragon, and Louise Brisette were also a great help in navigating the administrative aspects of pursuing a Ph.D. at UQAM in the absence of usable French skills. The Psychology Department at the University of Alberta gave me an office to work in while 1was in Edmonton, and 1 appreciate it greatly. The Giraldeau lab mernbers provided support and assistance over theyears: Kim­ berely Mathot, Guillaume Rieucau, Melanie Dappen, Stephanie Surveyer, François Racine, Marie-Pierre Bonin, Mewen Le Ho, Simone Tommasina, Sophie Godde, Mo­ hammad Afshar and Neeltje Boogert. l'd also like ta thank the people who made Montréal an entertaining place to be: Caterina Ferrari, Astrid Vik Stronen, Sarah Overington, and especially Neeltje Boogert and Jeremy Schwartzentruber, who gave me a place to stay on my many trips back. Pedro Peres-Neto provided welcome dis­ cussion of technical issues and helped with questions in spatial analysis, and one of ix my chapters owes its existence to Ralf Kurvers, who has been an incredibly patient coauthor over the past year. Throughout the Ph.D., l was supported financially by an NSERC CGS-D schol­ arship and NSERC Discovery grants to Luc-Alain Giraldeau as weil as une bourse d'exemption de frais majorés from UQAM. The research was also supported by NSERC grants ta Luc-Alain Giraldeau. Finally, l'd like to thank my friends and family, who kept me going while l pursued this dream. Anjie Khalid, Saira Khalid, Sheta Patel, Jaimini Patel, and Samina Ullah gave me a space to unwind in Edmonton and kept my wife sane while l was gone, and members of the Hurd lab and the Psych department provided companionship and the opportunity to chat science. Finally, Brandy Williams, my wife, put up with this adventure through four years of ups and downs; words could never express the debt l have to you, but l'Il end with 'thank you'. CONTENTS FOREWORD ...................................... .. vii LIST OF FIGURES. xiv LIST OF TABLES. xvii RÉSUMÉ ....................................... xix ABSTRACT . .. xxi CHAPTER l GENERAL INTRODUCTION 1 LI Frequency dependent ('social') foraging 2 1.1.1 Evolutionary game theory . 2 1.1.2 Frequency dependent selection 4 1.1.3 Evolutionary and ecological time . 4 UA Social foraging models . 6 1.1.5 Empirical work . 8 1.1.6 Personality and social foraging 9 1.2 Information use and learning . la 1.2.1 Personal and social information use while learning . Il 1.2.2 Social information use beyond the individual . 12 1.2.3 Empirical work on learning in producer-scrounger games . 12 1.3 Social foragers in space . 13 1.4 Modelling methodology ..... 15 1.4.1 Individual-based models 15 1.4.2 Genetic algorithms . 16 xi 1.4.3 Cellular automata .. 17 1.5 Thesis overview o •• 18 1.6 Bibliography 21 CHAPTER II LEARNING RU LES FOR SOCIAL FORAGING . · .... 31 ILl Abstract ... · .... 33 II.2 Introduction ...... · .... 34 II.3 Methods ....... · .... 37 11.3.1 The model .. .... 37 IIA Results .......... 42 IIA.l Base model .. 42 11.4.2 Environmental variability 47 11.5 Discussion ..... 47 II.6 Acknowledgements 55 11.7 Bibliography · ..... 56 CHAPTERIII COEVOLUTION OF SOCIAL INFORMATION USE. 61 IILl Abstract ... 63 III.2 Introduction . 64 II1.3 Methods . 65 III.3.1 Genetic algorithms . 67 III.3.2 Simulation types 68 IIIA Results . 69 III.5 Discussion . 74 II1.6 Acknowledgements 76 II1.7 Bibliography 77 CHAPTERIV SOCIAL FORAGING AND PERSONALITY .. 81 IVI Abstract . 83 IV2 Introduction . 84 IV3 The model. ... 85 IV3.1 The foraging simulation . 85 xii IV3.2 The genetic algorithm .................... 87 IV3.3 Measuring the evolution of personality and scrounging 88 IV4 Results. 89 IV5 Discussion 92 IV6 Acknowledgements 96 IV7 Bibliography 97 CHAPTERV SOCIAL LEARNING FOR SOCIAL FORAGING 105 VI Abstract... 107 V2 Introduction 108 V3 Methods... III V3.1 Neighbourhoods III V3.2 Payoffs... III V3.3 Updating.. 113 V3.4 Simulations 115 V3.5 Comparison to predictions of deterministic rate-maximizing model. 116 V4 Results . 117 V4.1 Equilibrium outcomes . 117 V4.2 Invasion........ 120 V4.3 Population dynamics . 120 V4.4 Effective group size . 122 V5 Discussion .... 125 V6 Acknowledgments 128 V7 Bibliography 129 CHAPTERVI LANDSCAPE GEOMETRY AND PRODUCER-SCROUNGER 135 VI.l Abstract . 137 VI.2 Introduction 138 VI.3 Methods. 140 VI.4 Results . 144 VI.5 Discussion 147 xiii VI.6 Acknowledgments 151 VI.7 Bibliography 152 CHAPTER VII GENERAL CONCLUSION 157 VII.1 Summary and Synthesis 157 VII.l.1 Advancing the models 157 VII.l.2 Information use . 159 VII.1.3 Spatial social foraging 160 VII.2 Future research . 161 VII.2.1 Thoughts on models 161 VII.2.2 Integrating information use . 163 VII.2.3 Foraging ahead, spatially 163 VII.3 Conclusion . 164 VIlA Bibliography 165 LIST OF FIGURES I.l [The Hawk-Dove game] . 3 1.2 [Frequency dependent selection.] . S 1.3 [Figure 1 ofMottley and Giraldeau (2000).] . 7 lA [Figure 1 of Gardner (1970).] .. 18 II.1 [Typical evolution oflearning rules in agents playing a producer-scrounger game.] ...................................... .. 43 IL2 [Mean optimal parameter values selected for by the genetic algorithm as group size increases.] 44 IL3 [Mean proportion ofindividuals that are specialist producers or scroungers as a function of group size.] .........................
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