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In the Hermit Crab, Pagurus Samuelis Loma Linda University TheScholarsRepository@LLU: Digital Archive of Research, Scholarship & Creative Works Loma Linda University Electronic Theses, Dissertations & Projects 9-2008 Behavioral Evidence for "Contextual Decision Hierarchies" In the Hermit Crab, Pagurus samuelis Wendy Lee Billock Follow this and additional works at: https://scholarsrepository.llu.edu/etd Part of the Biology Commons Recommended Citation Billock, Wendy Lee, "Behavioral Evidence for "Contextual Decision Hierarchies" In the Hermit Crab, Pagurus samuelis" (2008). Loma Linda University Electronic Theses, Dissertations & Projects. 665. https://scholarsrepository.llu.edu/etd/665 This Dissertation is brought to you for free and open access by TheScholarsRepository@LLU: Digital Archive of Research, Scholarship & Creative Works. It has been accepted for inclusion in Loma Linda University Electronic Theses, Dissertations & Projects by an authorized administrator of TheScholarsRepository@LLU: Digital Archive of Research, Scholarship & Creative Works. For more information, please contact [email protected]. LOMA LINDA UNIVERSITY School of Science and Technology in conjunction with the Faculty of Graduate Studies Behavioral Evidence for "Contextual Decision Hierarchies" In the Hermit Crab, Pagurus samuelis by Wendy Lee Billock A Dissertation submitted in partial satisfaction of the requirements for the degree of Doctor of Philosophy in Biology September 2008 © 2008 Wendy Lee BiHock All rights reserved Each person whose signature appears below certifies that this thesis, in his opinion, is adequate in scope and quality as a dissertation for the degree Doctor of Philosophy. erson Stephen G. Dunbar, Assistant Professor of Biology eonard R. Brand, Professor of Biology and Paleontology Ronald L. Carter, Professor of Biology, Vice Chancellor of Academic Affairs - L. J6 es Gibson, Professor of Biology, Director Geoscience Research Institute William K. Hayes, Professor of Biolo y 111 ACKNOWLEDGMENTS First and foremost, I would like to thank my advisor, Dr. Steve Dunbar, for his help, guidance, and encouragement throughout the entire graduate school experience. Through countless hours of discussions, laboratory training, and reviewing my writing, he has walked me through the process to develop this dissertation. He will always be an example to me of how to be an instructor, mentor, and friend to one's students. 1 also want to thank my committee members: Dr. Leonard Brand; Dr. Ron Carter; Dr. Jim Gibson; and Dr. Bill Hayes. As department chair, Dr. Brand not only advised me on the administrative aspects of graduate school, but he also showed genuine interest in my project and frequently discussed my research with me in the hallways. Although he was very busy as Vice Chancellor, Dr. Carter made time to discuss ideas and always inquired about my research when he would see me around campus. Dr. Gibson provided invaluable suggestions to improve this manuscript and his thoughtful comments on my research were a great help. I am also indebted to Dr. Hayes not only for his expertise in research design and statistical analysis, but also for the discussions and advice that led to the development of this line of research. Although not officially on my committee, Dr. Ernie Schwab has taken an interest in my research and provided a great deal of suggestions and encouragement that have improved the writing of this dissertation. In addition, discussions with Dr. Chris Tudge and Dr. Dan Fong led to the research of Chapter 3. iv This research was supported by grants from The Crustacean Society, the Southern California Academy of Sciences, and the Marine Research Group of Loma Linda University. I am also grateful to the Department of Earth and Biological Sciences of LLU for financial support of my research and graduate education. I also thank my colleagues at LLU who not only helped with various aspects of my research, but also encouraged me with their friendship. JaneIle Shives and April Sjoboen joined me for trips to the beach for animal collection, and helped with the maintenance of animals in the laboratory. Zia Nisani was a tremendous help in analyzing my data and navigating the waters of graduate school. Suggestions for improving this manuscript were made by Melissa Berube and April Sjoboen. I am forever grateful for the love and encouragement that I received from friends and family. The friendship and prayers of my church family has meant a great deal to me. My brother, Dr. Phil Sandefer, provided the inspiration to pursue my terminal degree. I also could not have completed this undertaking without the love and support of my mother, Jodie Fife. Finally, I want to thank my number one fan, my husband, Joe Billock. He has encouraged me throughout my years of graduate school and joined me in the field whenever he could. CONTENTS Approval Page iii Acknowledgements iv Table of Contents vi List of Tables List of Figures xi Abstract xii Chapter 1. Introduction to Hermit Crab Behavior 1 Objectives 1 Hermit Crab Behavior 4 Behavioral Ecology of Hermit Crabs Shell usage 5 Shell selection 6 Trophic level 9 Reproduction 9 Hermit Crab Classification 10 Pa gurus samuelis 11 Hermit Crab Sensory Apparati 12 Visual 12 Chemical 13 Tactile 14 Decision-Making and Cognition 15 Literature Cited 18 2. Contextual Decision Hierarchies in Crustaceans 24 Abstract 24 Introduction 25 Cognition defined 25 Cognitive processes 26 vi Attributes of Cognition 27 Attention 28 Representation 31 Learning 34 Problem solving 37 Contextual modulation 39 Crustacean Sensory Processing 41 Visual 42 Chemical 43 Tactile 47 Decision Hierarchies 49 Information processing 49 Sequential Decision Hierarchies 50 "Contextual Decision Hierarchies" 53 CDH model 58 Future of CDH Research 61 Acknowledgements 62 Literature Cited 63 3. Influence of Motivation on Behavior in the Hermit Crab, Pagurus samuelis 72 Abstract 72 Introduction 73 Materials and Methods 76 Animal collection and maintenance 76 Test protocol 76 Statistical analysis 79 Results 79 Discussion 82 Acknowledgements 89 Literature Cited 90 4. Shell and Food Acquisition Behaviors: Evidence for Contextual Decision Hierarchies in Hermit Crabs 92 vii Abstract 92 Introduction 93 Materials and Methods 95 Animal collection and maintenance 95 Shell acquisition 98 Food acquisition 99 Statistical analysis 100 Results 100 Shell acquisition 100 Food acquisition 103 Discussion 107 Shell acquisition 107 Food acquisition 111 Contextual Decision Hierarchies 113 Acknowledgments 117 Literature Cited 118 5. Influence of Sensory Cues on Predator Avoidance Behavior in the Hermit Crab, Pagurus samuelis 121 Abstract 121 Introduction 122 Materials and Methods 123 Animal collection and maintenance ..123 Experimental procedure 125 Statistical analysis 127 Results 128 Discussion 131 Contextual Decision Hierarchies 134 Acknowledgments 136 Literature Cited 137 6. Conclusions on Hermit Crab Behavior 140 Chapter II 140 viii Chapter III 143 Chapter IV 145 Chapter V 147 Conclusions 148 Literature Cited 152 TABLES Table Page 2-1. Summary of Sequential Decision Hierarchies found in a range of taxa 73 2-2. Summary of Contextual Decision Hierarchies found in a range of taxa 73 3-1. Description of the factorial treatment arrangements used to test hermit crab motivation to acquire shell and food resources 92 3-2. The number of hermit crabs that first made contact with either food or shells based on treatment 92 3-3. The number of hermit crabs that decided to insert into shells, feed, or take no action during 15 minute sessions based on treatment 92 4-1. Factorial treatment organization for Shell Acquisition and Food Acquisition experiments 121 4-2. P values of post-hoc test using Bonferroni adjustment for multiple comparisons based on the number of contacts with the shell in the Shell Acquisition experiment 122 4-3. P values of post-hoc test using Bonferroni adjustment for multiple comparisons based on decision time in the Shell Acquisition experiment 123 4-4. P values of post-hoc test using Bonferroni adjustment for multiple comparisons based on the number of contacts with gastropod flesh in the Food Acquisition experiment 124 4-5. P values of post-hoc test using Bonferroni adjustment for multiple comparisons based on decision time in the Food Acquisition experiment 125 5-1. Factorial treatment organization for predator avoidance experiment . .148 FIGURES Figures Page 2-1. Diagram of Contextual Decision Hierarchy model 74 3-1. Linear regression showing the relationship between hermit crab body weight and the preferred shell aperture width 93 3-2. The mean time to initial contact with objects (food or shell) based on treatment 94 3-3. The mean number of contacts with objects (food or shell) based on treatment 95 3-4. The mean time to initiate behavior based on objects (food or shell) and treatment 96 3-5. Proportion of hermit crabs exhibiting behaviors based on treatment during 15 minute test sessions 97 4-1. Diagram of test arena showing acrylic cylinder, hermit crab starting box, object placement, and pulley system 126 4-2. The mean number (± 1.0 SE) of shell contacts during Shell Acquisition treatments 127 4-3. The mean decision time (± 1.0 SE) during Shell Acquisition treatments 128 4-4. The mean number of shell contacts (± 1.0 SE) during Food Acquisition treatments 129 4-5. The mean decision time (± 1.0 SE) during Food Acquisition treatments 130 5-1. Mean number of contacts (± 1.0 SE) with a crab/model crab per session 149 5-2. The mean time (± 1.0 SE ) to initial shell contact 150 5-3. The mean
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