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Sex-Specific Behavioral Strategies for Thermoregulation in the Comon Chuckwalla (Sauromalus Ater) ______

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Sex-Specific Behavioral Strategies for Thermoregulation in the Comon Chuckwalla (Sauromalus Ater) ______ SEX-SPECIFIC BEHAVIORAL STRATEGIES FOR THERMOREGULATION IN THE COMON CHUCKWALLA (SAUROMALUS ATER) ____________________________________ A Thesis Presented to the Faculty of California State University, Fullerton ____________________________________ In Partial Fulfillment of the Requirements for the Degree Master of Science in Biology ____________________________________ By Emily Rose Sanchez Thesis Committee Approval: Christopher R. Tracy, Department of Biological Science, Chair Sean E. Walker, Department of Biological Science Paul Stapp, Department of Biological Science Spring, 2018 ABSTRACT Intraspecific variability of behavioral thermoregulation in lizards due to habitat, temperature availability, and seasonality is well documented, but variability due to sex is not. Sex-specific thermoregulatory behaviors are important to understand because they can affect relative fitness in ways that result in different responses to environmental changes. The common chuckwalla (Sauromalus ater) is a great model for investigating sex differences in thermoregulation because males behave differently from females while they actively defend distinct territories while females may not. I recorded body temperatures of wild adult chuckwallas continuously from May to July 2016, as well as operative environmental temperatures in crevices and aboveground sites used by chuckwallas for basking. I compared the effect of sex on indices of thermoregulatory accuracy and effectiveness, aboveground activity, and the time chuckwallas selected body temperatures relative to their preferred temperature range (34-39˚C) for May-July. Males and females achieved the same thermoregulatory accuracy and effectiveness, and amount of time at preferred body temperatures, but males were more active aboveground than females, especially in June. These results indicate that different behavioral strategies can accomplish the same thermoregulatory outcomes. Further, chuckwallas might be able to cope with predicted changes in climate that result in a shorter amount of time they could be active at preferred body temperature because these lizards were only active aboveground for one to four hours per day, a fraction of the time available. ii TABLE OF CONTENTS ABSTRACT ................................................................................................................... ii LIST OF TABLES ......................................................................................................... iv LIST OF FIGURES ....................................................................................................... v ACKNOWLEDGMENTS ............................................................................................. vi Chapter 1. A FIELD COMPARISON OF MALE AND FEMALE CHUCKWALLA THERMOREGULATION .................................................................................... 1 Introduction ........................................................................................................... 1 Methods ................................................................................................................ 4 Study Site ....................................................................................................... 4 Field Operative Environmental Temperatures (Te) ........................................ 5 Field Body Temperatures (Tb) ....................................................................... 8 Assessment of Thermoregulation .................................................................. 11 Behavioral Observations ................................................................................ 15 Estimation of Aboveground Activity ............................................................. 16 Results ................................................................................................................... 18 Thermal Quality of the Environment ............................................................. 18 Indices of Lizard Thermoregulation .............................................................. 22 Behavioral Observations of Aboveground Activity ...................................... 29 Aboveground Activity of Vested Lizards ...................................................... 31 Discussion ............................................................................................................. 33 Sex Differences in Habitat Use ...................................................................... 33 Causes of Sex Differences in Chuckwalla Habitat Use ................................. 39 Implications ................................................................................................... 41 APPENDICES ............................................................................................................... 45 A. ANNOTATED FOCAL ANIMAL OBSERVATIONS ................................ 45 B. METHODS FOR ABOVEGROUND ACTIVITY ........................................ 54 C. INDIVIDUAL LIZARD DATA .................................................................... 101 REFERENCES .............................................................................................................. 110 iii LIST OF TABLES Table Page 1. Commonly Used Measurements of Thermoregulation Established by Hertz et al. (1993) ................................................................................................ 14 2. Mean Thermal Quality of the Environment (de) .................................................. 22 3. Mean Daily Thermoregulatory Precision, Accuracy (db) and Effectiveness of Male and Female Chuckwallas ........................................................................ 25 4. Mean Thermoregulatory Accuracy (db) and Effectiveness During Diurnal Period of Male and Female Activity .................................................................... 26 iv LIST OF FIGURES Figure Page 1. Representative study site habitat and terrain of the study site, to the south of Coyote Canyon..................................................................................................... 6 2. An operative temperature (Te) model in full sun ................................................. 8 3. Male and female chuckwallas with vests housing iButton temperature loggers and radio transmitters .............................................................................. 10 4. Visual representation of method used for estimating chuckwalla emergences from crevices and amount of time they spent aboveground ................................ 19 5. Operative environmental temperatures, female body temperatures, and male body temperatures ............................................................................................... 21 6. The average number of hours that male and female chuckwalla body temperatures, and operative temperatures were within three temperature categories relative to preferred body temperature range ...................................... 27 7. The average number of hours that chuckwallas spent at different body temperatures during periods of diurnal activity ................................................... 28 8. The relative abundance of male and female chuckwalla observed aboveground per person per 15-minute interval ....................................................................... 30 9. The average amount of time that males and females spent active aboveground per day, estimated from changes in body temperature over time ........................ 32 10. Average number of male and female emergences from belowground ................ 32 v ACKNOWLEDGMENTS I would like to thank Dr. Chris Tracy for being a wonderful advisor to me over the past five years, and for enthusiastically jumping on board with this field project. Your gentle guidance, and passion for research has inspired me and helped me mature as a researcher (and person). I would like to thank my parents for supporting me throughout my academic endeavors, in endless ways. You, along with the rest of my incredible family, always understood when I had to miss out on family activities, and never stopped cheering me on. Without all of you I would have never come this far. A special thank you goes to my fiancé, Patrick, for being my main source of emotional support and my biggest cheerleader. Thank you for lifting me up when I would get down on myself, for spending countless hours helping me with R, for making sure I was still having fun, and so much more, and for continuing to do it all from over 1,300 miles away. To my sister, Erin, thank you for being there whenever I needed a burrito, a coffee break, a study buddy, or some laughter. To Melissa Goger, Amanda Hernandez, Cristy Rice, Velvet Park, and Melissa Gonzalez, thank you for always inspiring me to keep going and having my back. Your friendship means the world to me. To all my amazing CSUF friends, especially my lab mates (past and present), thank you for always encouraging me, spending time with me, and offering a helping hand whenever you could. I will always be grateful to have been surrounded by so many supportive people. I would like to give a special thanks to Dr. Dick Tracy for providing valuable input, and funding for this vi project, and for always being so positive. Your words of encouragement have meant a lot to me. Thank you to Deep Canyon Research Center (DOI: 10.21973/N3V66D) and Dr. Allan Muth, Mark Fisher, and Mr. Dave for providing a wonderful environment from which to conduct my field work. I would also like to thank Trevor Ruppert and Tomas Kneppers
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