Mating System and Dispersal Patterns in the Diamondback Terrapin (Malaclemys Terrapin)
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Mating system and dispersal patterns in the diamondback terrapin (Malaclemys terrapin) A Dissertation Submitted to the Faculty of Drexel University by Claire Marie Sheridan in partial fulfillment of the requirements for the degree of Doctor of Philosophy April 2010 ©Copyright 2010 Claire Marie Sheridan. All Rights Reserved ii ACKNOWLEDGEMENTS I would like to take this opportunity to thank all those whose abilities, efforts, facilities or funds allowed the completion of this project. First, I would like to thank my husband, Michael, for all of his love and support over the past 5 years. Without him, I would not have made it this far. I would like to thank the rest of my family, especially my mother and my parents-in-law, for all the emotional support that they have provided me. I thank my colleagues and friends in Drexel’s Biology Department: John Wnek for all the great discussions and help in selecting field sites; Jules Winters, Lori Lester, Abby Dominy, Karen Sullam, and Emily Basile for all hard work you helped me to accomplish and the good times that came along with it; Steve Pearson and Julia Stone for always being there to listen to me ramble on in the office; and Laurie Cotroneo for being there as a great friend and the one person with whom I could talk about genetics. I am also grateful to Ashley Lavender who has been a great listener and friend, regardless of moving miles away from Drexel. I am grateful to the other people who helped me in the field: Earthwatch volunteers, MATES students, Chaz Sineri, Sam Campos, Bryan Vorbach, Julie Fekete, Chris Wilson, and Mallory Watson. I am grateful to Jacquie Walters, Andy Harrison, and Dr. Ed Standora from Buffalo State for helping me during various field seasons. I am also grateful to James Houck for helping me in the lab. I thank Dr. Willem Roosenburg for his willingness to collaborate, facilitating sample collection at Poplar Island, and critically reviewing the mating system manuscript. iii I am very grateful to my advisors, Dr. Harold Avery and Dr. James Spotila for the years of guidance they have given me. They have both been great mentors and friends. I’ll never forget our many great field adventures. I am grateful to Dr. Walter Bien for his advice throughout the years and for showing me around the Pine Barrens. I would also like to thank the rest of the members of my committee, Dr. Kim Scribner, Dr. Jacob Russell, and Dr. Shivanthi Anandan for providing invaluable comments, suggestions, and thought provoking questions throughout my dissertation project. This research was funded by the Earthwatch Institute and the Betz Chair of Environmental Science at Drexel University. Scientific Collecting and Holding Permits were issued yearly by the NJDEP Division of Fish and Wildlife and Special Use Permits were issued yearly by the Edwin B. Forsythe National Wildlife Refuge. Research protocols were approved by Drexel University’s Institutional Animal Care and Use Committees. iv TABLE OF CONTENTS LIST OF TABLES ..................................................................................................... vi LIST OF FIGURES .....................................................................................................x ABSTRACT ............................................................................................................... xii CHAPTER 1: Introduction 1.1 Habitat fragmentation decreases biodiversity ....................................................1 1.2 Estuaries as important ecosystems .....................................................................2 1.3 The diamondback terrapin as a model vertebrate ..............................................4 1.4 Background on the diamondback terrapin .........................................................6 1.5 Terrapin dispersal patterns measured by direct analysis ...................................8 1.6 Terrapin dispersal patterns measured by genetic analysis .................................9 1.7 Effects of habitat fragmentation on mating behavior ......................................12 1.8 Diamondback terrapin mating behavior...........................................................13 1.9 Research questions and dissertation structure..................................................14 CHAPTER 2: Landscape genetic structure of the diamondback terrapin (Malaclemys terrapin) in a highly fragmented ecosystem 2.1 Abstract ............................................................................................................18 2.2 Introduction ......................................................................................................19 2.3 Methods............................................................................................................25 2.4 Results ..............................................................................................................35 2.5 Discussion ........................................................................................................38 CHAPTER 3: Sex-biased dispersal, natal philopatry, and home range movements of the diamondback terrapin, Malaclemys terrapin 3.1 Abstract ............................................................................................................53 3.2 Introduction ......................................................................................................54 3.3 Methods............................................................................................................58 3.4 Results ..............................................................................................................69 3.5 Discussion ........................................................................................................75 v CHAPTER 4: Inter-population variation of multiple paternity in the diamondback terrapin, Malaclemys terrapin 4.1 Abstract ............................................................................................................89 4.2 Introduction ......................................................................................................90 4.3 Methods............................................................................................................93 4.4 Results ............................................................................................................101 4.5 Discussion ......................................................................................................106 CHAPTER 5: Constraints on egg size, optimal egg size theory, and latitudinal reproductive variation in the diamondback terrapin (Malaclemys terrapin) 5.1 Abstract ..........................................................................................................121 5.2 Introduction ....................................................................................................122 5.3 Methods..........................................................................................................125 5.4 Results ............................................................................................................133 5.5 Discussion ......................................................................................................137 CHAPTER 6: Dissertation summary and applications 6.1 Introduction ..............................................................................................155 6.2 Summary of major findings .....................................................................155 6.3 Conservation implications .......................................................................159 6.4 Directions for future research ..................................................................168 LIST OF REFERENCES ........................................................................................175 CIRRICULUM VITA ..............................................................................................204 vi LIST OF TABLES 2-1. Genetic and geographical distances between the six diamondback terrapin populations sampled in Barnegat Bay, NJ. Genetic distance (upper diagonal) is represented by FST (and G'ST) and geographical distance in km (lower diagonal). Significant genetic differentiation is indicated by an asterisk (Fisher's exact test p < 0.05). ...........................................................................46 2-2. Correlation coefficients between genetic differentiation and the least cost paths generated for the range of cell cost values assessed. Correlation coefficients with significant p-values (<0.05) after bootstrapping (N = 1000) are shown in bold. The 'optimal' cell cost value chosen for each of the landscape features with significant p-values is shown with an *. The analysis values highlighted in gray added significantly to the null models (straight line distance and shoreline distance) based on AICc. Data are presented for 6 of the 18 tested cost values, but results are similar with the full dataset .............47 2-3. AICc, ∆AIC, and wAIC values for each of the best fit least cost models. For each landscape type, AICc values were calculated based on the individual landscape feature model with the highest correlation coefficient (r) ..............48 2-4. Correlation coefficients between genetic differentiation and the least cost paths of seven mixed models. The cell cost values for each landscape feature in the model are given. Correlation coefficients with significant p-values (<0.05) after bootstrapping (N = 1000) are shown in bold. The 'optimal' mixed model with the highest correlation coefficient