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Personality, Habitat Selection and Territoriality Kathleen Church A Habitat Complexity and Behaviour: Personality, Habitat Selection and Territoriality Kathleen Church A Thesis In the Department of Biology Presented in Partial Fulfilment of the Requirements For the Degree of Doctor of Philosophy (Biology) at Concordia University Montréal, Québec, Canada July 2018 © Kathleen Church, 2018 iii Abstract Habitat complexity and behaviour: personality, habitat selection and territoriality Kathleen Church, Ph.D. Concordia University, 2018 Structurally complex habitats support high species diversity and promote ecosystem health and stability, however anthropogenic activity is causing natural forms of complexity to rapidly diminish. At the population level, reductions in complexity negatively affect densities of territorial species, as increased visual distance increases the territory size of individuals. Individual behaviour, including aggression, activity and boldness, is also altered by complexity, due to plastic behavioural responses to complexity, habitat selection by particular personality types, or both processes occurring simultaneously. This thesis explores the behavioural effects of habitat complexity in four chapters. The first chapter, a laboratory experiment based on the ideal free distribution, observes how convict cichlids (Amatitlania nigrofasciata) trade-off the higher foraging success obtainable in open habitats with the greater safety provided in complex habitats under overt predation threat. Dominants always preferred the complex habitat, forming ideal despotic distributions, while subordinates altered their habitat use in response to predation. The second chapter also employs the ideal free distribution to assess how convict cichlids within a dominance hierarchy trade-off between food monopolization and safety in the absence of a iv predator. Dominants again formed ideal despotic distributions in the complex habitat, while dominants with lower energetic states more strongly preferred the complex habitat. For both laboratory experiments, personality did not predict habitat preference. The third chapter, a field study with juvenile Atlantic salmon (Salmo salar), tested whether stream restorations that increase habitat complexity will also select for particular personality traits, and we again found that complexity did not favour any particular personality types. A broader range perspective regarding the effects of habitat complexity on behaviour was addressed in the fourth chapter via a meta-analysis on a wide range of territorial and non-territorial taxa. Territoriality modified the effects of complexity on behaviour, likely due to the strong reliance of territorial species on visual cues. Taken together, all four chapters demonstrate the high context dependency of the effects of complexity on behaviour. Nevertheless, whether or not an individual is territorial emerged as an important predictor of how habitat complexity is likely to affect its behaviour. v Acknowledgements Above all, I would like to thank my supervisor, Dr. James Grant, for his endless support, guidance, and kindness throughout this entire process. I will be forever grateful for everything he has taught me over these last few years, and could not have asked for a better supervisor. I also would like to thank the other members of my committee, Dr. Grant Brown and Dr. Pedro Peres-Neto. I am grateful to Grant Brown for helpful discussion, especially in regards to everything to do with predation, and am grateful to Pedro Peres-Neto for assistance with the analysis, especially in regards to analysis with multivariate statistics. I am also grateful to Dr. Guillaume Larocque for his advice and help with the statistical analysis whenever I got stuck. For Chapters 1 and 2, I am grateful to the many people who assisted with the feeding trials, including Talia Ke, Ashlee Prévost, Sean Devine, Michelle Le Blanc, Tammy Brunet, Nathalie Jreidini, Dr. Anne Almey, Guillaume Roullet, Dr. Pierre Chuard, and Jean-Michel Matte; I am especially grateful to Jasmine Roberts, who helped with the experimental design of chapter 1, and to Wyatt Toure, who assisted with the vast majority of the feeding trials. For chapter three, I am grateful to Dr. Rick Cunjak for use of the field station at Catamaran Brook, and to Dr. Jess Ethier, Adam Trapid, Yifan Ling, Elise McDowell and Aaron Fraser for field assistance, and for also being so entertaining during the long days in the field. I am also grateful to Dr. Denis Réale for his assistance with the experimental design and statistics, and for taking the time to share his expertise on personality and personality analysis with me. For chapter 4, I would also like to thank everyone who assisted in the literature search for the meta-analysis, including Sean Devine, Wyatt Toure, Nathalie Jreidini, Anika Forget, and especially Naomi Safir, who conducted her own meta-analysis on the effects of habitat complexity on behaviour for her undergraduate research project. vi I am also grateful to my parents, Dr. Dana Williams and Dr. Michael Church, for inspiring me to pursue a PhD way back in my early childhood, and for their lifelong support of my scholastic endeavors and interest in the natural world. Thank you for giving me such a thorough education on psychology throughout my entire life, information which has helped immensely when studying the behaviour and personality of fish. I am also grateful for their continued support, along with my brother, Eric Church, and sister in law Tamara Zderic, throughout this process. I am grateful to the friends that have helped me along in this process, especially Dr. Anne Almey, who is always available with no-nonsense advice on everything academic and otherwise, and to my boyfriend, Paelo Gonzalez, for his support and practical assistance during the writing process. Finally, I would like to thank my funding sources. I am grateful to Concordia University for awarding me the Frederick Lowy Scholarship fund, as well as providing me with two Teaching Assistantships. I am also grateful to the Natural Sciences and Engineering Research Council of Canada for financing this research through my supervisor, Dr. James Grant. vii Contributions of Authors I contributed to the experimental design, conducted the experiments, analyzed the data, and wrote all four chapters of the thesis. Dr. Grant contributed to the experimental design, assisted with the interpretations of results and edited all four chapters. Each chapter within this thesis was prepared as a manuscript for submission to a peer-reviewed journal for publication. Chapter 1 has been submitted to Behavioral Processes as: Church, K. D., & Grant, J. W. A. Ideal despotic distributions in convict cichlids (Amatitlania nigrofasciata)? Effects of predation risk and personality on habitat preference. Chapter 2 is being prepared for submission to a behavioural ecology journal as: Church, K. D., & Grant, J. W. A. Effects of habitat complexity, dominance and personality on habitat selection: ideal despotic cichlids. Chapter 3 has been published as: Church, K. D., & Grant, J. W. A. 2018. Does increasing habitat complexity favour particular personality types of juvenile Atlantic salmon, Salmo salar? Animal Behaviour, 135: 139-146. Chapter 4 will be submitted to an ecological journal as: Church, K. D., & Grant, J. W. A. A meta-analysis of the effects of habitat complexity and territoriality on behaviour. viii Table of Contents List of Figures………………………………………………………………….………………..xiv List of Tables…………………………………………………………………………………..xviii General Introduction…………………………………………………………………….………...1 Chapter 1. Ideal despotic distributions in convict cichlids (Amatitlania nigrofasciata)? Effects of predation risk and personality on habitat preference.………………….…………….…………....6 Abstract…………………………………………………………………………………..………..7 1. Introduction……………………………………………………………………………..……....8 2. Material and methods…………………………………………………………………….……10 2.1. Feeding treatment……………………………………………………………...…….10 2.2. Individual behavioural assay…………………………………………………...……11 2.3. Statistics………………………………………………………………………..……14 2.3.1. Group trials……………………………………………………………..…14 2.3.2. Individual behavioral assay………………………………………………..14 3. Results………………………………………………………………………………………....15 3.1. Habitat Choice……………………………………………………………….……...15 ix 3.2. Foraging Success…………………………………………………………….……...16 3.3. Aggression…………………………………………………………………………..17 3.4. Growth………………………………………………………………………………17 3.5. Individual Behaviour………………………………………………………………..18 4. Discussion………………………………………………………………………….………….19 4.1. Conclusions……………………………………………………….…………………22 Figure Legends………………………………………………………………………………..…24 Tables…………………………………………………………………………………………….25 Figures……………………………………………………………………………………………27 Supplementary Figure Legend..……………………………………………………………….…34 Supplementary Tables………………………………………………………………………...….35 Supplementary Figure…………………………………………………………………………....36 Chapter 2. Effects of habitat complexity, dominance and personality on habitat selection: ideal despotic cichlids………………………………………….…………………………………...….38 Abstract…………………………………………………………………………………….….…39 Introduction…………………………………………………...………………………………….40 Methods…………………………………………………………..………………………..……..43 x Feeding experiment………………………………………………………………………43 Individual behavioural assay……………………………………………………………..45 Statistics………………………………………………………………………………….47 Group trials………………………………………………………………………47 Individual behavioural assay……………………………………………………..48 Results………………………………………………………………..………………………......49 One-Patch Treatments……………………………………………………………………49 Two-Patch Treatments………….………………………………………………………..51
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