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Ammospermophilus Harrisii) Behavior and Spatial Ecology of the Harris' Antelope Ground Squirrel (Ammospermophilus harrisii) Item Type text; Electronic Thesis Authors Burnett, Alexandra D. Publisher The University of Arizona. Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction, presentation (such as public display or performance) of protected items is prohibited except with permission of the author. Download date 28/09/2021 10:13:42 Link to Item http://hdl.handle.net/10150/636507 BEHAVIOR AND SPATIAL ECOLOGY OF THE HARRIS’ ANTELOPE GROUND SQUIRREL (Ammospermophilus harrisii) by Alexandra Burnett ____________________________ Copyright © Alexandra Burnett 2019 A Thesis Submitted to the Faculty of the SCHOOL OF NATURAL RESOURCES & THE ENVIRONMENT In Partial Fulfillment of the Requirements For the Degree of MASTER OF SCIENCE NATURAL RESOURCES – WILDLIFE CONSERVATION & MANAGEMENT In the Graduate College THE UNIVERSITY OF ARIZONA 2019 2 THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE As members of the Master’s Committee, we certify that we have read the thesis prepared by: Alexandra Burnett titled: and recommend that it be accepted as fulfilling the thesis requirement for the Master’s Degree. _________________________________________________________________ Date: ____________Sep 9, 2019 John L Koprowski _________________________________________________________________ Date: ____________Sep 10, 2019 Michael Bogan _________________________________________________________________ Date: ____________Sep 9, 2019 bret pasch Final approval and acceptance of this thesis is contingent upon the candidate’s submission of the final copies of the thesis to the Graduate College. I hereby certify that I have read this thesis prepared under my direction and recommend that it be accepted as fulfilling the Master’s requirement. _________________________________________________________________ Date: ____________Sep 9, 2019 John L Koprowski T hesis Committee Chair School of Natural Resources and the Environment 3 Acknowledgments I would like to sincerely thank my graduate advisor, Dr. John Koprowski for providing the opportunity for me to complete my master’s degree at the University of Arizona, as well as his support and guidance throughout my thesis project. I would further like to thank my committee members, Dr. Bret Pasch and Dr. Bill Mannan, for their availability and support along the way. Special thanks to Sarah Hale, Melissa Merrick, and Vicki Greer for training support and field assistants, Alexis Blair, Sandy Slovikosky, Helena Yomantas, Anne-Laure Blanche, Megan Bethel, Nicole Bokanowski, Brandon Mayer, and Eduardo Gracia for countless hours of field support and data input. Thanks to M. Merrick and M. Mazzamuto for guidance with software and data analysis, and V. Greer for logistical support with permitting, equipment, and vehicles. I am grateful for the support and guidance provided by the Koprowski Conservation Research Laboratory throughout my time in Arizona. Finally, I thank my partner, Kevin Wagner, and my parents, friends, and family, for their support and patience throughout my thesis project. Alexandra D. Burnett 4 Table of Contents List of Figures……………………………………………………………………………………..5 Abstract……………………………………………………………………………………………8 Chapter 1: Introduction…………………………………………………………………………..10 Chapter 2: Present Study…………………………………………………………………………15 Summary of Conclusions…………………………………………………………………………15 Literature Cited……………………………………………………………………………….….17 Appendix A: On using a nonhibernating squirrel to inform models of complexity……..………23 Introduction………………………………………………………………………………………24 Methods…………………………………………………………………………………………..30 Results……………………………………………………………………………………………33 Discussion………………………………………………………………………………………..36 Appendix B: Effects of shrub encroachment on antipredator behavior…………….……………59 Introduction………………………………………………………………………………………60 Methods………………………………………………………………………………………..…62 Results……………………………………………………………………………………………66 Discussion………………………………………………………………………………………..67 5 List of Figures A.1 Figure 1: Average home range size (ha) of Ammospermophilus harrisii in southeastern Arizona throughout 2017 and 2018. Data are divided by the mating season (Jan-Apr), juvenile season (May-Aug), and nonbreeding season (Sep-Dec). There are no statistical differences of home ranges sizes among season or sex.………………………………………………………54 A.1 Figure 2: 95% and 50% kernel densities of individual antelope squirrels (Ammospermophilus harrisii) in southeastern Arizona. Home ranges of a) all individuals found within study site 2 and 3 of the Santa Rita Experimental Range in which we collected enough points for a home range. Males are represented by warmer colors (red, orange, yellow) and females are represented by cooler colors (green, blue, purple). Individuals overlapped intra- and intersexually: b) female (green)-male (red) overlap during the months of June-August 2018, b) female-female overlap during the months of June-August 2017, and c) male-male overlap during the months of October and November 2017…………………………………...………………...55 A.1 Figure 3: Average number of antipredator vocalizations given by Harris’ antelope ground squirrels (Ammospermophilus harrisii) detected per hour in each month of 2017 and 2018 (pooled) in southeastern Arizona. Dashed lines represent seasonal boundaries. Both years start with a low hourly rate of antipredator vocalizations until April in the mating season (months 1- 4), remain at a moderate level through the juvenile season (months 5-8), and peak in the nonbreeding season after juvenile dispersal (months 9-12) …………………………………...56 6 A.1 Figure 4: Spectrogram of an adult Harris’ antelope squirrel (Ammospermophilus harrisii) call given in southeastern Arizona, looking at frequency (kHz) over time(s). Color represents amplitude; warmer colors represent higher amplitudes. Calls feature a trill with high-frequency overtones that descends in amplitude over time. Squirrels frequently give calls repeatedly, in bouts……………………………………………………………………………………………...57 A.1 Figure 5: Spectrogram of a juvenile Harris’ antelope squirrel (Ammospermophilus harrisii) call in the presence of a Western diamondback rattlesnake (Crotalus atrox) in southeastern Arizona. Frequency (kHz) is represented on the y-axis and time (s) is represented on the x-axis. Chirps consist of a short, broadband syllable that is repeated.…………………………………..58 A.2 Figure 1: Selection indices of burrow locations of Ammospermophilus harrisii in southeastern Arizona within each vegetation class based on direct observations compared to average availability across study sites. We included all burrow locations that were used by marked individuals at least once (n=22 animals, 192 total burrow locations). Negative values indicate selection against, positive values indicate selection for the vegetation class. Burrows beneath Opuntia were used most commonly. Prosopis and Celtis were the next most frequent genera for burrowing beneath, though at a significantly lower frequency. Individuals also used burrows beneath Senegalia and Cylindropuntia, as well as Ferocactus, Ephedra, and Heteropogon……………………………………………………………………………………78 A.2 Figure 2: Selection indices of foraging items within each vegetation class based on direct observations of feeding compared to average availability across study sites (n=47 direct 7 observations of feeding). Negative values indicate selection against, positive values indicate selection for the vegetation class. We observed Ammospermophilus harrisii feeding on flowers and fruits of Opuntia the vast majority of the time in southeastern Arizona. The next most frequently observed food source was grass seeds (Poaceae)…………………………………...79 A.2 Figure 3: Selection indices of alarm calling locations within each vegetation class compared to average availability of each vegetation class across study sites. Only observations in which the plant the individual was calling from was positively identified were included in the analyses (n=18 observations). Negative values indicate selection against, positive values indicate selection for the vegetation class. Ammospermophilus harrisii called from mesquite trees (Prosopis) in 38.8% of known alarm call locations in southeastern Arizona. Prickly pear (Opuntia) were the next most common observed alarm call locations (33.3%), and then cholla (Cylindropuntia, 11.1%) and hackberry (Celtis, 11.1%) …………………………………………………………...80 A.2 Figure 4: Selection indices of active individuals within each vegetation class compared to average availability across study sites (n=22 individuals, 257 observations). Negative values indicate selection against, positive values indicate selection for the vegetation class. Ammospermophilus harrisii in southeastern Arizona selected for bare ground slightly avoided shrub and cacti………………………………………………………..…………………81 8 Abstract Ground squirrels provide a valuable model system for studying both theoretical and applied questions in behavioral ecology. High variability among species ecology imposes selective forces that result in vast differences in social and communication systems. Resource distribution, body size, and kin selection drive social evolution, leading to hypotheses linking larger body size and shorter active periods to greater sociality, which in turn has implications for strategies that may have interesting implications for models of social and communicative complexity. We conducted a two-year study of a nonhibernating squirrel,
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