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Desert Tortoise United States Department of Agriculture Desert Tortoise Forest Service Intermountain (Gopherus agassizii): Research Station General Technical Report INT-GTR-316 Status-of-Knowledge July 1995 Outline With References Mark C. Grover Lesley A. DeFalco The Authors Acknowledgments Mark C. Grover has Bachelor (1988) and Master B. Campbell and T. Day of the U.S. Department of Science (1992) degrees in zoology from Brigham of the Interior, Bureau of Land Management Li- Young University. He is now a Ph.D. degree candi- brary in Denver, CO, provided literature search as- date in biology at the University of Virginia. He sistance and timely correspondence necessary for was a biological technician for the Intermountain completion of this manuscript. J. F. Douglass, J. B. Research Station during the 1988 field season. His Iverson, and D. J. Morafka provided constructive re- research interests center on herpetologic ecology. views of the earlier version of the manuscript. T. A. Duck of Arizona Strip Bureau of Land Management Lesley A. DeFalco completed her Bachelor’s de- provided many valuable and sometimes obscure ref- gree program at Colorado State University in May erences. J. Jenson, P. Grover and S. Grover typed 1989. She began research on the desert tortoise in the original manuscript. The Utah State Office of 1989 with the U.S. Fish and Wildlife Service; she the Bureau of Land Management provided encour- now works on the same project as a research biolo- agement and essential financial support. Project gist with the National Biological Survey in Fort Leader E. D. McArthur of the Intermountain Re- Collins, CO, and is a graduate student in biology at search Station’s Shrubland Biology and Restoration Colorado State University. She has conducted ex- Research Work Unit provided leadership and sup- tensive field studies, concentrating on the nutrition port from the beginning of the project to its conclu- and foraging ecology of the population of desert tor- sion. D. J. Pietrzak and J. W. Sites provided en- toises inhabiting the northeastern Mojave Desert. couragement and support that helped the authors complete this manuscript. The desert tortoise on the cover was drawn by Lesley DeFalco. Intermountain Research Station 324 25th Street Ogden, UT 84401 Contents Page Page Introduction ...............................................................1 Commensals ..........................................................45 Taxonomy ...................................................................3 Reproduction ............................................................46 Taxonomic classification ........................................3 Characteristics at sexual maturity......................46 Original description ...............................................3 Breeding season ....................................................46 Synonyms ................................................................3 Courtship and mating behavior ...........................46 Relatedness to similar species ............................... 4 Nests and egg deposition .....................................47 Description .................................................................4 Egg development, incubation, and hatching ....... 49 Physical description ...............................................4 Growth and population structure ...........................50 Similar species ........................................................5 Size and growth rate ............................................51 Morphology ................................................................5 Longevity ..............................................................51 Adults ......................................................................5 Age determination ................................................52 Hatchlings ...............................................................8 Physiology ................................................................53 Sexual dimorphism ................................................9 Thermoregulation .................................................53 Anomalies ...............................................................9 Water balance .......................................................55 Regional variation in morphology .......................11 Metabolism ...........................................................58 Genetics ....................................................................12 Hematology ...........................................................59 General..................................................................12 Bone and scute regeneration ...............................60 Regional genetic variation ...................................12 Feeding behavior, diet, and nutrition .....................61 Hybridization ........................................................14 Feeding behavior ..................................................61 Paleontology and paleoecology ................................14 Diet ........................................................................64 Evolution ...............................................................14 Nutrition ...............................................................69 Paedomorphosis ....................................................15 Scats ........................................................................72 Os transiliens .......................................................15 Description ............................................................72 Prehistoric range ..................................................16 Frequency of defecation .......................................73 Fossil sites beyond the current range .................16 Use as markers .....................................................73 Fossil sites within the current range ..................17 Mortality factors ......................................................73 Habitat and climate .............................................17 Disease ..................................................................73 Distribution and population status ........................18 Parasitism .............................................................75 General..................................................................18 Injury ....................................................................76 Arizona ..................................................................19 Predation ..............................................................76 California ..............................................................20 Human-related mortality .....................................79 Nevada ..................................................................22 Environmental factors .........................................81 Utah ......................................................................23 Estimating the years since death Mexico ...................................................................24 by examining carcasses .....................................81 Habitat .....................................................................24 Other causes .........................................................81 Habitat type and vegetation associations ...........24 Behavior ...................................................................82 Vegetation .............................................................27 Courtship behavior ...............................................82 Climate ..................................................................28 Sex and species discrimination ............................82 Precipitation .........................................................28 Social behavior .....................................................82 Soil ........................................................................30 Defensive behavior ...............................................84 Elevation ...............................................................31 Agonistic behavior ................................................84 Terrain ..................................................................31 Feeding behavior ..................................................86 Habitat deterioration ..............................................33 Drinking behavior ................................................86 Livestock grazing .................................................33 Sleeping behavior .................................................86 Off-road vehicles (ORV’s) .....................................39 Digging behavior ..................................................86 Urbanization and agricultural development ......40 Thermoregulatory behavior .................................86 Roads .....................................................................40 Audition ................................................................86 Other factors .........................................................41 Olfaction ................................................................86 Burrows and dens ....................................................41 Vision ....................................................................87 Cover sites ............................................................41 Vocalization ..........................................................87 Summer burrows or dens .....................................41 Righting reflex ......................................................88 Winter dens or burrows .......................................43 Locomotion ............................................................89
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