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Desert Tortoises (Gopherus spp.) Fact Sheet: Bibliography & Resources

Desert Tortoise (Gopherus spp.)


Alberts, A., D. Rostral, V. Lance 1994. Studies on the chemistry and social significance of chin gland secretions in the desert tortoise, Gopherus agassizii. Herpetological Monographs 8:116-124.

Allison LJ, McKuckie AM. 2018. Population trends in Mojave desert tortoises (Gopherus agassizii). Herpetological Conservation and Biology. 13(2):433–452.

Auffenberg, W. 1974. Checklist of fossil land tortoises (Testudinidae). Bulletin of the Florida State Museum, Biological Sciences. 18(3): 121-251.

Auffenberg, W. 1977. Display behavior in tortoises. American Zoologist. 17:241-250.

Averill-Murray. R.C., B.E. Martin, S.J. Bailey and E.B. Wirt 2002. Activity and behavior of the Sonoran Desert Tortoise in Arizona. In: T. Van Devender (Ed.) The Sonoran Desert Tortoise. University of Arizona Press. Tucson, Arizona. pp. 135-159.

Averill-Murray RC, Darst CR, Field KJ, Allison LJ. 2012. A New Approach to Conservation of the Mojave Desert Tortoise. BioScience. 62(10):893–899. doi:10.1525/bio.2012.62.10.9.

Berry, K.H. (Ed.) 1984. The status of the desert tortoise (Gopherus agassizii) in the United States. Report to U.S. Fish and Wildlife Service from the Desert Tortoise Council. U.S. Department of Interior, Bureau of Land Management. Riverside, California.

Berry, K.H. 1986. Desert tortoise (Gopherus agassizii). Relocation: implications of social behavior and movements. Herpetologica 42:113-125.

Berry, K.H. 1990 (as amended). The status of the desert tortoise in California in 1989.U.S. Bureau of Land Management, Riverside, California; amended to include data from 1990, 1991, and 1992.

Berry KH, Murphy RW. 2019. Gopherus agassizii (Cooper 1861) – Mojave Desert Tortoise, Agassiz’s Desert Tortoise. In: Rhodin, A.G.J., Iverson, J.B., van Dijk, P.P., Stanford, C.B., Goode, E.V., Buhlmann, K.A., Pritchard, P.C.H., and Mittermeier, R.A. (Eds.). Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian Research Monographs 5(13):109.1–45. doi: 10.3854/crm.5.109.agassizii.v1.2019;

Berry KH, Allison LJ, McLuckie AM, Vaughn M, and Murphy RW. 2021. Gopherus agassizii. The IUCN red list of threatened species [Internet]. Version 2021-2; eT97246272A3150871 [species assessed 2020 Oct 01; page accessed 2021 Nov 16]. doi: 10.2305/IUCN.UK.2021-2RLTS.T97246272A3150871.en.

Berry ,K.H., B.L. Homer, W. Alley, M. Chaffee, G. Haxel 2001. Health and elevated mortality rates in desert tortoise populations: the role of arsenic and other potential toxins. Abstract for Arsenic Group Meeting, February 21022m 2001 Denver Federal Center.

Berry K, Murphy R. 2019. Gopherus agassizii (Cooper, 1861) - Agassiz’s Desert Tortoise, Mojave Desert Tortoise. Chelonian Research Foundation and Turtle Conservancy. 5(13):1–43. doi:10.3854/crm.5.109.agassizii.v1.2019.

Boarman, W.I. 2002. Threats to desert tortoise populations: a critical review of the literature. U.S. Geological Survey Western Ecological Research Center, San Diego, CA

Bonin, F.B. Devaux, A. Dupre. 2006. Turtles of the World. Johns Hopkins University Press. Baltimore, Maryland.

Bramble, D.M. 1982. Scaptochelys: Generic revision and evolution of gopher tortoises. Copia 1982:852-867.

Brown, M.B. Schumacher, I.M. P.A. Klein, K. Harris, T. Corell, E.R. Jacobson. 1994. Mycoplasma agassizii causes upper respiratory tract disease in the desert tortoise. Infection and Immunity. 62:4580--86.

Brown, D.R., J.L.Merritt, E.R. Jacobson, P.A. Klein, J.G. Tully, M.B. Brown 2004. Mycoplasma testudineum sp.nov., from a desert tortoise (Gopherus agassizii) with upper respiratory tract disease. International Journal Systematic and Evolutionary Microbiology 54:1527-1529.doi: 10.1099/ijs.0.63072-0 

Burge, B. 1977. Daily and seasonal behavior, and areas utilized by the desert tortoise Gopherus agassizi in southern Nevada. Proceedings of Symposium The Desert Tortoise Council 1977:59-94.

Bury, R. B. (Ed) 1982. North American tortoise conservation and ecology. Wildlife Research Reports 12. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service.

California Department of Fish and Game 2010. Species Explorer Details for desert tortoise. Downloaded Aug. 10, 2010.

Collins, J. T., and T. W. Taggart. 2002. Standard common and current scientific names for North American amphibians, turtles, reptiles, and crocodilians. Fifth edition. The Center for North American Herpetology, Lawrence, Kansas. iv + 44 pp.

Cooper JG 1861. New California Animals. Proceedings of the California Academy of Sciences. 2:118–123. PDF

Corn, PS 1994. Recent trends of desert tortoise populations in the Mojave Desert. In: R. B. Bury and D. J. Germano, (Eds) The Biology of North American Tortoises National Biol. Survey, Washington, DC. Fish and Wildlife Research 13 pp. 85-93.

Crawford NG, Faircloth BC, McCormack JE, Brumfield RT, Winder K, Glenn TC. 2012. More than 1000 ultraconserved elements provide evidence that turtles are the sister group of archosaurs. Biology Letters. doi: 10.1098/rsbl.2012.0331

Crumly, CR 1984. The genus name for North American gopher tortoises. Proceedings of the Symposium of the Desert Tortoise Council 1984:147-148.

Curtin, J.J., G.R. Zug, J.R. Spotila 2009. Longevity and growth strategies of the desert tortoise (Gopherus agassizii) in two American deserts. Journal of Arid Environments. doi:10.1016/j.jaridenv.2008.11.011.

Desert Tortoise Life History. [accessed 2022 Jan 26].

Duda, J. J., A. J. Krzysik, and J. E. Freilich. 1999. Effects of drought on desert tortoise movement and activity. Journal of Wildlife Management

Edwards, T., C. Schwalbe, D. Swann, C. Goldberg 2004. Implications of anthropogenic landscape change on inter-population movements of the desert tortoise (Gopherus agassizii). Conservation Genetics 5:485-499.

Edwards T, Karl A, Vaughn M, Rosen P, Meléndez Torres C, Murphy R. 2016. The desert tortoise trichotomy: Mexico hosts a third, new sister-species of tortoise in the Gopherus morafkai–G. agassizii group. ZK. 562:131–158. doi:10.3897/zookeys.562.6124.

Edwards T, Karl AE, Rosen PC, Vaughn M, Melendez Torres C, Kiester AR, Goode EV. 2018. Gopherus evgoodei (errata version published 2019). The IUCN red list of threatened species [Internet]. Version 2021-2; e.T90377823A144764819 [species assessed 2018 Jan 25; page accessed 2021 Nov 16]. doi: 10.2305/IUCN.UK.2018-2RLTS.T90377823A144764819.en.

Ernst, C.H. and J.E. Lovich. 2009. Turtles of the United States and Canada. Johns Hopkins University Press, Baltimore. pp. 540-570.

Federal Register 2009. 90-Day Finding on a Petition to List the Sonoran Population of Desert Tortoise (Gopherus agasizzii) as a Distinct Population Segment (DPS) With Critical Habitat. August 28, 2009. Vol. 74, No. 166.

Germano, D. F.H. Poughy, D. Morafka, E. Smith, M. Demlong 2002. Growth of Desert tortoises. In: T. Van Devender (Ed.) The Sonoran Desert Tortoise. University of Arizona Press. Tucson, Arizona. pp. 265-288.

Grover, M.C. and L.A. DeFalco 1995. Desert Tortoise (Gopherus agassizii): Status-of-Knowledge Outline With References. U.S.D.A. Forest Service Intermountain Research Station. General Technical Report INT-GTR-316. 1-140 pp.

Haines, D. F., T. C. Esque, and C. R. Tracy. 1999. Hibernation temperatures, timing, and site characteristics for desert tortoises in the northeast Mojave Desert (poster presentation). The 24th Annual Desert Tortoise Council Symposium. St. George, UT.

Hansen, R.M., M.K. Johnson, T. R. Van Devender 1976. Foods of the desert tortoise, Gopherus agassizii, in Arizona and Utah. Herpetologica 32(3):247-251.

Harper, D. 2010. Gopher. On-Line Etymology Dictionary. Downloaded October 2010.

Homer, B. L., K. H. Berry, M. B. Brown, G. Ellis, and E. R. Jacobson.  1998.  Pathology of diseases in wild desert tortoises from California.  J. Wildl. Dis. 34(3):508-523.

Integrated Taxonomic Information System (ITIS). Desert Tortoise (Gopherus agassizii). Downloaded Sept 8, 2010.

Integrated Taxonomic Information System database (ITIS) [Internet]. Desert Tortoise (Gopherus agassizii); [accessed 06 Jan 2017].

Jacobson, E.R., T.J. Wronski, J.Schumacher, C. Reggiardo, K. Berry 1994. Cutaneous dyskeratosis in free-ranging desert tortoises, Gopherus agassiziii, in the Colorado Desert of Southern California. Journal of Zoo and Wildlife Medicine 25(1):68-81.

Jarchow, J.L., H.E. Lawler, T.R. Van Devender, C.S. Ivanyi 2002. Care and diet of captive Sonoran desert tortoises. In: T. Van Devender (Ed.) The Sonoran Desert Tortoise. University of Arizona Press. Tucson, Arizona. pp. 289-311.

Johnson, A.J., D.J. Morafka, E.R. Jacobson 2006. Seroprevalence of Mycoplasma agassizii and tortoise herpesvirus in captive desert tortoises (Gopherus agassizii) from the Greater Barstow Area, Mojave Desert, California. Journal of Arid Environments 67:192-201.

Kelly EC, Cypher BL, Westall TL. 2021. Predation on desert tortoises (Gopherus agassizii) by desert canids. Journal of Arid Environments. 189:104476. doi:10.1016/j.jaridenv.2021.104476.

Lamb, T., J. C. Avise, J. W. Gibbons: Phylogeographic patterns in mitochondrial DNA of the desert tortoise (Xerobates agassizi) and evolutionary relationships among the north American gopher tortoises. Evolution, 43: 76-87, 1989.

Le, M., C. Raxworthy, W. McCord, L. Mertz. 2006 A molecular phylogeny of tortoises (Testudines:Testudinidae) based on mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 40(2):517-531.

Lemm, J. 2006. Field Guide to Amphibians and Reptiles of the San Diego Region California Natural History Guides. University of California Press. Berkeley, California.

Lovich, J. E. and D. Bainbridge. 1999. Anthropogenic degradation of the Southern California desert ecosystem and prospects for natural recovery and restoration. Environ. Mgmt. 24:309-326.

Lovich JE, Edwards T, Berry KH, Puffer SR, Cummings KL, Ennen JR, Agha M, Woodard R, Brundige KD, Murphy RW. 2020. Refining genetic boundaries for Agassiz’s desert tortoise (Gopherus agassizii) in the western Sonoran Desert: the influence of the Coachella Valley on gene flow among populations in southern California. Frontiers of Biogeography. 12(3). doi:10.21425/F5FBG46888. [accessed 2022 Jan 25].

Luckenbach, R. Ecology and management of the desert tortoise (Gopherus agassizii) in California. In: North American Tortoises: Conservation and Ecology. United States Department of Interior Fish and Wildlife Service Wildlife Research Report 12. pp. 1-37.

McCord, R. 2002. Fossil history and evolution of the gopher tortoises (Genus Gopherus). In: T. Van Devender (Ed.) The Sonoran Desert Tortoise. University of Arizona Press. Tucson, Arizona. pp. 29-51

Marlow, R.W. and K. Tollestrup 1982. Mining and exploitation of natural mineral deposits by the desert tortoise, Gopherus agassizii. Animal Behavior 30:475-478.

Miller, J.D. and S.A. Dinkelacker 2008. Reproductive structures and strategies of turtles. In: Wyneken, J., M. Godfrey , V. Bels (Eds.) Biology of Turtles. CRC Press: Taylor and Francis Group. Boca Raton, London, New York. pp. 225-278.

Murphy, R.W., K.H. Berry, T. Edwards, A.M. McLuckie 2007. A genetic assessment of the recovery units for the Mojave population of the desert tortoise, Gopherus agassizii. Chelonian Conservation and Biology 6:229-251.

Murphy R, Berry K, Edwards T, Leviton A, Lathrop A, Riedle JD. 2011. The dazed and confused identity of Agassiz’s land tortoise, Gopherus agassizii (Testudines: Testudinidae) with the description of a new species and its consequences for conservation. ZK. 113:39–71. doi:10.3897/zookeys.113.1353.

Murray, R.C. 1997. Gopherus agassizii (desert tortoise). Diet. Herpetological Review 37:77-78.

Nagy, K.A., B.T. Henen, and D.B. Vyas. 1998. Nutritional quality of native and introduced food plants of wild desert tortoises. Journal of Herpetology 32:260-267.

Nagy, K.A. and P.A. Medica. 1986. Physiological ecology of desert tortoises in southern Nevada. Herpetologica 42(1):73-92.

National Fish and Wildlife Foundation. 2022. Mojave Desert Tortoise Recovery Partnership 2022. [accessed 2022 Jan 25].

Nowak, L., . G. Nowak, R. Parker, G.R. Stewart 1999. Desert Tortoise. In: Thelander, C. (Ed) Life on the Edge; A Guide to California's Endangered Natural Resources. Biosystems Books, Santa Cruz, CA and Heyday Books, Berkeley

Oftedal, O. 2002. Nutritional ecology of the desert tortoise in the Mohave and Sonoran Deserts. In: T. Van Devender (Ed.) The Sonoran Desert Tortoise. University of Arizona Press. Tucson, Arizona. pp. 194-241.

Oftedal, O. 2003. Desert tortoises nutritionally constrained by a shortage of high PEP plants, and if so, what do we do? 28th Annual meeting and symposium of the desert tortoise council, Feb. 21-23, 2003. Abstract.

Osorio, S. and R. Bury 1982. Ecology and Status of the Desert Tortoise (Gopherus agassizii) on Tiburón Island, Sonora. In: Bury, R. Bruce, (Ed.) North American Tortoises:Conservation and Ecology. United States Department of the Interior Fish and Wildlife Service. Wildlife Research Report 12. Washington, D.C.

Pritchard, P.C.H. 2008. Evolution and structure of the turtle shell. In: Wyneken, J., M. Godfrey , V. Bels (Eds.) Biology of Turtles. CRC Press: Taylor and Francis Group. Boca Raton, London, New York. pp. 45-83.

Renous, S., Lapparent de Broin, F. M. Depecker, Davenport, J. Bels, V.2008. Evolution of locomotion in aquatic turtles. In: Wyneken, J., M. Godfrey , V. Bels (Eds.) Biology of Turtles. CRC Press: Faylor and Francis Group. Boca Raton, London, New York. pp. 97-139

Reynoso, V-H. and Montellano-Ballesteros, M. 2004. A new giant turtle of the genus Gopherus (Chelonia: Testudinidae) from the Pleistocene of Tamaulipas, Mexico, and a review of the phylogeny and biogeography of gopher tortoises. Journal of Vertebrate Paleontology 24(4):822-837.

Rhodin AGJ, Stanford CB, Dijk PPV, Eisemberg C, Luiselli L, Mittermeier RA, Hudson R, Horne BD, Goode EV, Kuchling G, et al. 2018. Global Conservation Status of Turtles and Tortoises (Order Testudines). Chelonian Conservation and Biology. 17(2):135. doi:10.2744/CCB-1348.1.

Riedle, J.D., R.C. Averill-Murray, C.Lutz, D.K. Bolen 2008. Habitat use by desert tortoises (Gopherus agassizii) on alluvial fans in the Sonoran Desert, south-central Arizona. Copeia 2008 Nol 2. 414-420.

Rosmarino & Connor 2008. Petition to list the Sonoran Desert tortoise (Gopherus agassizii) under the U.S. Endangered Species Act, Office of Endangered Species. U.S. Fish and Wildlife Service, U.S. Dept. of Interior. Wild Earth Guardians and Western Watersheds Project

Ruby , D.E. and H.A. Niblick 1994. Behavioral inventory of the desert tortoise: development of an ethogram. Herpetological Monographs. 8:88-102.

SDZWA Public Relations. 2024 (14 Apr). 70 critically endangered desert tortoises reintroduced into wild emerge from brumation. San Diego (CA): San Diego Zoo Wildlife Alliance. [accessed 2024 Apr 16].

Shaffer HB, McCartney-Melstad E, Ralph P, Bradburd G, Lundgren E, Vu J, Hagerty B, Sandmeier F, Weitzman C, Tracy R. 2017. Desert Tortoises in the Genomic Age: Population Genetics and the Landscape. [accessed 2021 Oct 12]. doi:10.1101/195743

Stanford, C.B. The Last Tortoise: A Tale of Extinction in Our Lifetime. Belknap Press, Harvard University Press, Cambridge, Massachusetts.

Stebbins, R.C. 2003. Field Guide to Western Reptiles and Amphibians. Houghton Mifflin Co. Boston, New York. pp. 255-257.

TCC [Turtle Conservation Coalition: Stanford C, Rhodin A, Dijk PP, Horne B, Blanck T, Goode E, Hudson R, Mittermeier R, Currylow A, Eisemberg C, et al., eds.]. 2018. Turtles in trouble: the world’s 25+ most endangered tortoises and freshwater turtles - 2018. Ojai (CA): IUCN SSC Tortoise and Freshwater Turtle Specialist Group, Turtle Conservancy, Turtle Survival Alliance, Turtle Conservation Fund, Chelonian Research Foundation, Conservation International, Wildlife Conservation Society, and Global Wildlife Conservation; 80 pp. Turtle Conservancy Coalition

Thelander, C.G. 1999. Life on the Edge: A Guide to California's Endangered Natural Resources. Wildlife. Biosystems Books Santa Cruz, California.

Tollis M, DeNardo DF, Cornelius JA, Dolby GA, Edwards T, Henen BT, Karl AE, Murphy RW, Kusumi K. 2017. The Agassiz’s desert tortoise genome provides a resource for the conservation of a threatened species. Jaiswal P, editor. PLoS ONE. 12(5):e0177708. doi:10.1371/journal.pone.0177708.

Tortoise and Freshwater Turtle Specialist Group 1996. Gopherus agassizii. The IUCN red list of threatened species [Internet]. Version 2018-2; e.T9400A12983037 [species assessed 1996 Aug 01; page accessed 2019 Feb 19].

UNEP. 2019. Species+: Gopherus agassizii [online database]. Nairobi, Kenya: UNEP [compiled by UNEP-WCMC, Cambridge, UK]. [accessed 2019 Feb 19]. 

U.S. Fish and Wildlife Service 1991. Endangered and threatened wildlife and plants: Finding on a petition to list the Sonoran desert tortoise as threatened or endangered 56 Fed. Reg. 29453 (June 27, 1991)

U.S. Fish and Wildlife Service 1994. Desert tortoise (Mojave population) recovery plan. U.S. Fish and Wildlife Service, Portland, OR: 73

U.S. Fish and Wildlife Service 2009. Spotlight Species 5-Year Action Plan (2010-2014)

U.S. Fish and Wildlife Service 2010. Desert tortoise recovery office. Reno, Nevada [Internet]. Mojave population of the desert tortoise (Gopherus agassizii). 5-year review: summary and evaluation. 9-30-10. [accessed 2022 Feb 01].

USFWS (U.S. Fish and Wildlife Service). 2015. Range-wide monitoring of the Mojave desert tortoise (Gopherus agassizii): 2013 and 2014. Annual Report. U.S. Department of the Interior, U.S. Fish and Wildlife Service, Reno, Nevada, USA.

U.S. Fish and Wildlife Service. 2021. Range-wide monitoring of the Mojave Desert Tortoise by the Desert Tortoise Recovery Office, Reno, NV [Internet]. Status of the Desert Tortoise and its critical habitat. 8-16-21 [accessed 2022 Feb 01].  PDF

Walde, A.D., D.K. Delaney, M.L. Harless, L.L. Pater 2007. Osteophagy by the desert tortoise (Gopherus agassizii). The Southwestern Naturalist. 52(1): 147-149.

Walde, A.D., M.L. Harless, D.K. Delaney, L.L. Pater 2006. Gopherus agassizii (Desert Tortoise). Diet. Herpetological Review 37(1)77-78.

Wallis, I.R., Henen, B.T. & Nagy, K.A. (1999). Egg size and annual egg production by female desert tortoises (Gopherus agassizii): the importance of food abundance, body size, and date of egg shelling. Journal of Herpetology 33, 394–408.

Woodbury, M. R. Hardy 1948. Studies of the desert tortoise, Gopherus agassizii. Ecological Monographs, 19(2): 145-200.

Woodman, A.P. and S.M. Juarez. 1988. Juvenile desert tortoises utilized as primary prey of nesting common ravens near Kramer, California. Paper presented at the 13th Annual Meeting and Symposium of the Desert Tortoise Council held March 26-27, 1988, Laughlin, Nevada.

Van Devender, T.R., K.B. Moodie 1977. The desert tortoise in the Late Pleistocene with comments about its earlier history. Desert Tortoise Council Proceedings of 1977 Symposium. Las Vegas, Nevada March 1977.

Van Devender, T. 2002. Cenozoic environments and the evolution of the gopher tortoises (genus Gopherus) In: T. Van Devender (Ed.) The Sonoran Desert Tortoise. University of Arizona Press. Tucson, Arizona. pp. 29-51

Van Devender, T. 2002. The Sonoran Desert Tortoise. University of Arizona Press. Tucson, Arizona.

Williams, T. 1981 Experimental analysis of the gait and frequency of locomotion of the tortoise, with a simple mathematical description. Journal of Physiology 310:307-320.

Zylstra, E.R., R.J. Steidl 2007. Habitat use by Sonoran desert tortoises. Journal of Wildlife Management 73(5):747-754.

Additional Internet Resources

Close-up of Shell Scutes

Desert tortoise shell

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