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University of Nevada, Reno Effects of Fire on Desert Tortoise (Gopherus Agassizii) Thermal Ecology a Dissertation Submitted in P

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University of Nevada, Reno Effects of Fire on Desert Tortoise (Gopherus Agassizii) Thermal Ecology a Dissertation Submitted in P University of Nevada, Reno Effects of fire on desert tortoise (Gopherus agassizii) thermal ecology A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Ecology, Evolution, and Conservation Biology by Sarah J. Snyder Dr. C. Richard Tracy/Dissertation Advisor May 2014 THE GRADUATE SCHOOL We recommend that the dissertation prepared under our supervision by SARAH J. SNYDER Entitled Effects of fire on desert tortoise (Gopherus agassizii) thermal ecology be accepted in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY C. Richard Tracy, Ph.D., Advisor Kenneth Nussear, Ph.D., Committee Member Peter Weisberg, Ph.D., Committee Member Lynn Zimmerman, Ph.D., Committee Member Lesley DeFalco, Ph.D., Graduate School Representative David W. Zeh, Ph. D., Dean, Graduate School May, 2014 i ABSTRACT Among the many threats facing the desert tortoise (Gopherus agassizii) is the destruction and alteration of habitat. In recent years, wildfires have burned extensive portions of tortoise habitat in the Mojave Desert, leaving burned landscapes that are virtually devoid of living vegetation. Here, we investigated the effects of fire on the thermal ecology of the desert tortoise by quantifying the thermal quality of above- and below-ground habitat, determining which shrub species are most thermally valuable for tortoises including which shrub species are used by tortoises most frequently, and comparing the body temperature of tortoises in burned and unburned habitat. To address these questions we placed operative temperature models in microhabitats that received filtered radiation to test the validity of assuming that the interaction between radiation and radiation-absorbing properties of the model can result in a single, mean radiant absorptance regardless of whether the incident solar radiation is direct unfiltered or filtered by plant canopies, using the desert tortoise as a case study. We found that operative temperatures were nearly identical within microhabitats no matter the absorptance used in the model, which supports the use of a single mean absorptance in modeling operative temperature for animals in a variety of habitats. Using validated models, we calculated indices of thermal habitat quality, and also the hours tortoises could be active within their preferred body temperature range each day across the tortoise activity season. The thermal quality index was similar between burned and unburned habitat, but unburned habitat was more thermally heterogeneous, and it provided slightly longer activity times for tortoises within their preferred body temperature range as long as they could access all thermal microhabitats in their home range, and they were not ii constrained by habitat configuration or microhabitat abundance. However, the differences found were small and likely not biologically significant. Living Yucca species provided the best thermal microhabitats for tortoises during the summer and were used most frequently by tortoises in burned and unburned habitat. We found that burrows in burned and unburned habitats were of similar thermal quality regardless of whether shrubs were present near the burrow mouth. Finally, body temperatures of tortoises using burned and unburned habitat were similar, but tortoises in burned habitat had slightly higher minimum body temperatures. The small magnitude differences detected in minimum body temperature would likely not affect physiological performance. Taken together, these results suggest that burned habitat may be thermally suitable for desert tortoises but direct mortality from fire, and change in nutritional resources, should also be considered when evaluating the effects of fire, and the suitability of burned habitat for tortoise population persistence. iii ACKNOWLEDGMENTS This project was truly a collaborative effort made possible by a partnership between the University of Nevada, Reno (UNR) and the United States Geological Survey (USGS). In addition to funding, USGS provided me with countless helpers and hours of manpower, prudent advice, and inspiration during all stages of this project. I would especially like to thank Ken Nussear, Todd Esque, Lesley DeFalco, Kristina Drake, Rich Inman, Katherine Nolte, and Andrew Modlin for their guidance and patience throughout my tenure at UNR. I would like to thank UNR, USGS, and Coyote Springs Investment, LLC for generous funding that supported this research. This project required the help of many people, from painting 300 metal bowls, to tracking tortoises in the hot desert from sundown to sunup, and I have many people to thank for their time and assistance. Many Student Conservation Association (SCA) interns and USGS employees provided hours of work in the field including C. Aiello, P. Babbin, S. Bajwa, T. Barwise, P. F. Berntson, F. Chen, J. Curran, Cosman, K. Drake, E. Driver, E. Frankel, J. Gleave, G. Howerton, R. Inman, B. Jacobs, E. J. Jay, Jeffreys, A. Jones, M. Kang, A. Klapperich, R. Kunicki, D. Martin, A. Modlin, S. Ng, J. Niedbala, K. Nolte, K. Oddenino, C. Phillips, B. Reyonlds, J. Steiner, A. Terry, and V. Vanzerr. I’d also like to thank Rich Inman for his help using the spectroradiometer, Martyn Drabik- Hamshare for his help in the field collecting irradiance data, and Fran Sandmeier, Dick Tracy, and Chava Weitzman for collecting iButtons from the field. I’d like to acknowledge all of the help I had from friends who attended my infamous “bowl sanding iv parties” and especially Jenny Todd for her help constructing operative temperature models. I sought much-needed statistical advice from many individuals including Ken Nussear, Rich Inman, Kevin Burls, and Matt Forister. My wonderful committee members, Dick Tracy, Ken Nussear, Lesley DeFalco, Lynn Zimmerman, and Peter Weisberg have given me valuable feedback in the form of reading earlier drafts of manuscripts to giving advice on my research design. Their guidance has helped me to become a competent ecologist and conservation biologist, and for that I am very grateful. The professors I worked for as a teaching assistant, including Elena Pravasudova, Charlotte Borgeson, Tom Kidd, Pamela Sandstrom, Dick Tracy, and Patricia Berninsone have been exceptionally understanding as I have tried to juggle dissertation work with my teaching responsibilities. These individuals and my teaching experiences have inspired me to pursue a career in undergraduate education. Administrative staff, faculty, and students in the EECB program and Biology Department at UNR have been influential in my success as a graduate student by filling many capacities over the past seven years. I would like to thank the Tracy lab members, new and old, for their camaraderie, humor, and continual support. I believe Tracy lab members share a unique and unrivaled bond that is everlasting. Thanks to Bridgette Hagerty, Fran Sandmeier, Chris Geinger, Rich Inman, Tia Pilikian, Mike Pesa, Amy Barber, Stephanie Wakeling, Nichole Maloney, Lee Lemenager, Chava Wietzman, Jenny Todd, Michelle Gordon, John Gray, and honorary member Molly Bechtel. Additionally, I don’t think I would have survived my early years as a graduate student without the sage advice from Cynthia Downs on how to navigate paperwork, and advice from Bridgette Hagerty, Fran Sandmeier, and v Chris Geinger on the inner workings of the Tracy lab. My dwindling cohort of graduate students and new students that have entered the program since I started have been my Nevada family and, while they are too numerous to list in full, I can’t begin to thank them enough. I would like to especially thank Kevin Badik and Kevin Burls who were my first wonderful roommates in Reno and have remained close buddies ever since. Over the past several years Cynthia Scholl, Chava Weitzman, and Kevin Burls have seriously kept me going and introduced new joy into my life and for that I will be eternally grateful. My best friends from undergrad, Alli Clearwater and Shane Eaton, have always provided me with necessary distraction and adventure when work became overwhelming, which allowed me to persevere. Of special note I would first like to thank my major professor, Dick Tracy, who has challenged, inspired, and guided me as a graduate student. Dick has been my greatest advocate and has pushed me to become a critical thinker and passionate scientist and teacher. No student could ask for a more supportive, generous, and caring mentor. No thanks would be complete without acknowledging my truly wonderful family. My parents and grandparents have been my greatest supporters and have believed in me when I had doubts. Thanks to my parents for introducing me to the beauty of the natural world as a child, to my father for his unconditional understanding, and to my mother who first planted the seed that I ought to pursue a career in wildlife biology. Finally, I would like to thank the majestic tortoise, which ambles across the dry, desolate scrub as a symbol of the rugged Nevada desert, that I called home. vi TABLE OF CONTENTS ABSTRACT ……………………........................................................................................i ACKNOWLEDGEMENTS …………………………...…………………………...…….iii LIST OF TABLES …………………...…………………………………………………..ix Chapter 1…………………………………………...…………………………………ix Chapter 2 ……………………………………………..………………………………ix Chapter 3 …………………………………………...…………………………………x LIST OF FIGURES ……………………………………………………………..……….xi Chapter 1……………………………………………………………………...………xi Chapter 2 ………………………………………………………………..……………xi Chapter 3 ……………………………………………………………………………xiii
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