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CHARACTERIZING TERRITORIALITY AND THE MECHANISMS THAT MEDIATE IT IN FEMALE ANOLIS GUNDLACHI LIZARDS _______________________________________ A Dissertation presented to the Faculty of the Graduate School at the University of Missouri-Columbia _______________________________________________________ In Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy _____________________________________________________ by ELLEE G. COOK Dr. Manuel Leal, Dissertation Supervisor JULY 2019 The undersigned, appointed by the dean of the Graduate School, have examined the dissertation entitled CHARACTERIZING TERRITORIALITY AND THE MECHANISMS THAT MEDIATE IT IN FEMALE ANOLIS GUNDLACHI LIZARDS Presented by Ellee G. Cook, a candidate for the degree of Doctor of Philosophy, and hereby certify that, in their opinion, it is worthy of acceptance. ___________________________________ Professor Manuel Leal ___________________________________ Professor Reginald Cocroft ___________________________________ Professor Kevin Middleton ___________________________________ Professor Johannes Schul ACKNOWLEDGEMENTS Countless people have contributed to this dissertation and to the successful culmination of my graduate work at the University of Missouri. I have had the tremendous good fortune to have an incredible support system, both in terms of my academic and scientific development, and the emotional support that is critical for success in this field. I have learned more from Manuel Leal than from any other teacher or mentor I have had the fortune to work with. His mentorship challenged me to develop as a critical thinker, and has instilled in me a confidence in facing new challenges. I will be forever grateful for his perpetual willingness to talk and laugh, his tremendous investment in me as a student and scientist, and in always encouraging me to pursue lines of inquiry of interest to me in science and beyond. The other members of my committee have also invested substantially in my success, and I thank Rex Cocroft, Kevin Middleton, and Johannes Schul for taking the time to teach me, talk with me, and make this dissertation what it is. The Division of Biological Sciences has been a hub of support. No day was complete without my visiting with Nila Emirich and Rebecca Ballew, and I would not have survived this process nearly as gracefully without the support of Melody Kroll, Alan Marshall, Nick Valentine, and all the other staff that make this place great. I also thank Lori Eggert and Candi Galen for their mentorship in teaching, and thank Barb Sonderman and Richard Daniel for always being ready to meet to have executive discussions about our classes. And last but certainly not least, my fellow graduate students and lab mates. I can’t possibly mention all of the individuals I’ve spoken to about science, life, and life- after-graduate-school, but I thank you all! In particular, I thank Jake Burkhart, Ronnie ii Lacombe, Arianne Messerman, Erin Sellner, and Levi Storks for always being my people both in science and beyond. This work would not have been possible without the help of field assistants who went above and beyond what I asked of them. Deborah Lopez-Nieves, Karen Patterson, Jessica Pita-Aquino, and Phil Pearson, your thoughtfulness and hard work made these projects better and I am so grateful for the adventures we had. Furthermore, my collaborators Miguel Acevedo, Matthew B. Lovern, David Clark so graciously agreed to collaborate with us, and I thank them for the tools that made these projects what they are. Finally, none of this would have been possible without the support of my family. Doug and Paula Cook have been my cheerleaders for the last six years, never once doubting my ability to succeed. I am forever grateful to them for reminding me that I can do anything I set out to do. iii TABLE OF CONTENTS ACKNOWLEDGEMENTS ................................................................................................ii LIST OF TABLES ...............................................................................................................vi LIST OF FIGURES .............................................................................................................vii ABSTRACT ..........................................................................................................................viii CHAPTER 1 • INTRODUCTION INTRODUCTION ...........................................................................................................1 LITERATURE CITED .....................................................................................................7 CHAPTER 2 • TERRITORIALITY: A SURVEY OF THE PROGRESSION FROM OBSERVATION TO EXPERIMENTATION TO UNDERSTAND THE EVOLUTION, FUNCTION, AND REGULATION OF TERRITORIAL BEHAVIOR ABSTRACT ..................................................................................................................10 INTRODUCTION ...........................................................................................................10 CASE STUDY ...............................................................................................................18 CONCLUSIONS AND FUTURE DIRECTIONS ...................................................................27 REFERENCES ...............................................................................................................29 CHAPTER 3 • SIMILAR BEHAVIOR, SHARED MECHANISM? EVALUATING THE INLFUENCE OF BODY CONDITION ON TERRITORIAL BEHAVIOR IN FEMALE ANOLIS GUNDLACHI LIZARDS ABSTRACT ..................................................................................................................43 INTRODUCTION ...........................................................................................................44 MATERIALS AND METHODS ........................................................................................49 RESULTS .....................................................................................................................56 DISCUSSION ................................................................................................................60 ACKNOWLEDGEMENTS ................................................................................................69 iv REFERENCES ...............................................................................................................71 CHAPTER 4 • EVALUATING THE POTENTIAL FOR TESTOSTERONE TO MEDIATE AGGRESSIVE BEHAVIOR IN WILD FEMALE ANOLIS GUNDLACHI LIZARDS ABSTRACT ..................................................................................................................91 INTRODUCTION ...........................................................................................................92 MATERIALS AND METHODS ........................................................................................97 RESULTS .....................................................................................................................106 DISCUSSION ................................................................................................................110 ACKNOWLEDGEMENTS ................................................................................................117 REFERENCES ...............................................................................................................118 CHAPTER 5: TESTOSTERONE AND PARASITE LOAD: AN INVESTIGATION OF THE IMMUNOCOMPETENCE HYPOTHESIS IN WILD ANOLIS GUNDLACHI LIZARDS ABSTRACT ..................................................................................................................141 INTRODUCTION ...........................................................................................................141 MATERIALS AND METHODS ........................................................................................146 RESULTS .....................................................................................................................149 DISCUSSION ................................................................................................................150 ACKNOWLEDGEMENTS ................................................................................................153 REFERENCES ...............................................................................................................155 CHAPTER 6 • CONCLUSIONS CONCLUSIONS .............................................................................................................166 SUMMARY OF RESULTS AND FUTURE DIRECTIONS ......................................................167 VITA......................................................................................................................................171 v LIST OF TABLES CHAPTER 2. Table 1. Results of literature search for lizard territoriality.......................................40 CHAPTER 3. Table 1. Descriptions of observed territorial behaviors .............................................80 Table 2. Results of t-tests comparing variables across sampling years .....................81 CHAPTER 4. Table 1. Description of territorial behaviors ..............................................................127 Table 2. Description of aggressive behaviors observed.............................................128 Table 3. Model comparison for probability of undetectable concentrations of testosterone in males and females ..............................................................................130 Table 4. Model comparison for probability of undetectable