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Gopherus morafkai MURPHY, BERRY, EDWARDS, LEVITON, LATHROP & RIEDLE, 2011

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Higher TaxaTestudinidae, Testudinoidea, Testudines (turtles)
Subspecies 
Common NamesE: Morafka’s Desert Tortoise 
SynonymGopherus morafkai MURPHY, BERRY, EDWARDS, LEVITON, LATHROP & RIEDLE 2011
Xerobates agassizii COOPER 1861 (partim)
Testudo agassizii — COPE 1875
Gopherus agassizii — STEJNEGER 1893
Scaptochelys agassizii — BRAMBLE 1982
Gopherus morafkai — TTWG 2014
Gopherus morafkai — TTWG 2021 
DistributionUSA (Arizona), Mexico (Sonora)

Type locality: Tucson (approximate location 32° 7' N, 110° 56' W, elevation 948 m), Pima County, Arizona  
Reproductionoviparous. 
TypesHolotype: CAS 33867 (California Academy of Sciences) ; juvenile collected on 9 July 1912 by H. Brown and preserved in ethanol. Paratypes. ROM (Royal Ontario Museum) 47501, formerly CAS 13165, an immature tortoise collected by H. Brown from 20 miles (32 km) west of Tucson, (presumably the Roskruge Mountains, Pima County) Arizona, USA (approximate location 32° 7' N, 111° 18' W, where tortoises occur today), on 9 March 1908, received at CAS alive on 23 March 1908, and died 8 July 1908; CAS 34263, a juvenile collected by J.R. Slevin in the Catalina Mountains (Santa Catalina Mountains), foothills at west end of mountains, Pima County, Arizona, USA on 15 May 1912 (approximate location 32° 21’ N, 110° 57’ W). Specimens are preserved in ethanol. 
DiagnosisDiagnosis: All of the species of Gopherus and their hybrids can be easily diagnosed using molecular data. Morphologically, G. morafkai can be separated from both G. flavomarginatus and G. polyphemus in having relatively smaller front feet. Whereas the distance from the bases of the first to fourth claws is the same on all feet in G. morafkai, in the latter two species the distance from the bases of the first and third claws on the forelimb is about the same as the distance between the bases of the first and fourth claws on the hindlimb (Auffenberg and Franz 1978). The diagnosis of living specimens of G. morafkai, G. berlandieri and G. agassizii can be impossible in captive tortoises because of extensive hybridization (Edwards et al. 2010) and because of abnormalities in shell, head and limb integument from poor nutrition (Donoghue 2006). However, in non‐hybrid individuals, G. morafkai can be separated from G. berlandieri in having a rounded snout when viewed from above as opposed to a wedge‐shaped snout in G. berlandieri (Auffenberg and Franz 1978). Further, in G. morafkai the gular projections do not normally diverge, and it has a single axillary scale preceding each bridge, yet in G. berlandieri the gular projections often diverge and the axillary scales are often paired. Morphologically, G. morafkai can be separated from G. agassizii in having a relatively narrower shell, shorter gular scutes, shorter projections of the anal scutes and in having a flatter, pear‐shaped carapace (Table 1). Ecologically, whereas G. agassizii predominantly occurs in valleys and alluvial fan topography, G. morafkai prefers slopes and rocky hillsides (Riedle et al. 2008), including animals of the isolated population in northwestern Arizona (McLuckie et al. 1999). 
CommentSynonymy: after MURPHY et al. 2011. Note that Xerobates lepidocephalus has been synonymized with G. agassizii by CRUMLY & GRISMER 1994, a position that is also followed by the TTWG 2014 checklist but not Murphy et al. 2011 who synonymize Xerobates lepidocephalus with morafkai and more recently with G. evgoodei (Edwards et al. 2016). 
EtymologyThe new species is a patronym for the late Professor David Joseph Morafka in recognition of his many contributions to the biology and conservation of the species of Gopherus and his unsurpassed way of facilitating research, even among researchers with very different perspectives. 
References
  • Averill-Murray, Roy C. 2019. Reproductive Ecology and Life History of Female Sonoran Desert Tortoises (Gopherus morafkai). Sonoran Herpetologist 32 (2): 37 - get paper here
  • Averill-Murray, Roy C.; Christen H. Fleming, and J. Daren Riedle 2020. Reptile Home Ranges Revisited: a Case Study of Space Use of Sonoran Desert Tortoises (Gopherus morafkai). Herpetological Conservation and Biology, 15 (2): - get paper here
  • Averill-Murray, Roy C.; Terry E. Christopher, and Brian T. Henen 2018. Reproductive Ecology and Life History of Female Sonoran Desert Tortoises (Gopherus morafkai). Herpetological Monographs Dec 2018, Vol. 32, No. 1: 34-50. - get paper here
  • Babb, Randall D., Thomas C. Brennan, David D. Kandiyeli, Christina M. Akins and Thomas R. Jones. 2013. Geographic Distribution: Gopherus morafkai (Sonoran Desert tortoise). Herpetological Review 44 (4): 623 - get paper here
  • Bezy, Robert L. and Charles J. Cole 2014. Amphibians and Reptiles of the Madrean Archipelago of Arizona and New Mexico. American Museum Novitates (3810): 1-24 - get paper here
  • Bramble DM 1982. Scaptochelys: generic revision and evolution of gopher tortoises. Copeia 1982: 852–867, doi: 10.2307/1444097 - get paper here
  • Bridges, Andy; Heather L. Bateman, Audrey K. Owens, Cristina A. Jones and William Miller 2016. Microhabitat Selection of Juvenile Sonoran Desert Tortoises (Gopherus morafkai) in Central Arizona. Chelonian Conservation and Biology 15 (2): 219-230 - get paper here
  • Chessman, B. C. 2024. The conundrum of turtle and tortoise basking: A critical review. Journal of Zoology - get paper here
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  • Edwards T, Karl AE, Vaughn M, Rosen PC, Torres CM, Murphy RW 2016. The desert tortoise trichotomy: Mexico hosts a third, new sister-species of tortoise in the Gopherus morafkai–G. agassizii group. ZooKeys 562: 131-158. doi: 10.3897/zookeys.562.6124 - get paper here
  • Edwards, Taylor 2014. The Mexican Tortoise project. The Tortoise 1 (3): 128-141 - get paper here
  • Edwards, Taylor; Mercy Vaughn, Philip C. Rosen, Cristina Meléndez Torres, Alice E. Karl, Melanie Culver and Robert W. Murphy 2016. Shaping species with ephemeral boundaries: the distribution and genetic structure of desert tortoise (Gopherus morafkai) in the Sonoran Desert region. Journal of Biogeography43 (3): 484–497, DOI: 10.1111/jbi.12664 - get paper here
  • Enderson, Erik F.; Thomas R. Van Devender, Robert L. Bezy 2014. Amphibians and reptiles of Yécora, Sonora and the Madrean Tropical Zone of the Sierra Madre Occidental in northwestern Mexico. Check List 10 (4): 913-926 - get paper here
  • Gray, K.M., and R. Steidl. 2014. Effects of buffelgrass invasion on density and condition of Sonoran Desert Tortoises (Gopherus morafkai). Sonoran Herpetologist 27 (4):91-95. - get paper here
  • Lemos-Espinal JA, Smith GR, Rorabaugh JC 2019. A conservation checklist of the amphibians and reptiles of Sonora, Mexico, with updated species lists. ZooKeys 829: 131-160 - get paper here
  • Lovich, Jeffrey E.; Roy C. Averill-Murray, Mickey Agha, Joshua R. Ennen, and Meaghan Austin 2017. Variation in Annual Clutch Phenology of Sonoran Desert Tortoises (Gopherus morafkai) in Central Arizona. Herpetologica Dec 2017, Vol. 73, No. 4: 313-322. - get paper here
  • MANNING, JEFFREY A.; TAYLOR EDWARDS, JOHN CLEMONS, DANIEL J. LEAVITT, CAREN S. GOLDBERG & MELANIE CULVER. 2022. Scat as a source of DNA for population monitoring. Ecology and Evolution 12(11): 1–7. - get paper here
  • Murphy, Robert W.;Kristin H. Berry, Taylor Edwards, Alan E. Leviton, Amy Lathrop, J. Daren Riedle 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. ZooKeys 113: 39–71; doi: 10.3897/zookeys.113.1353 - get paper here
  • Nevárez-de los Reyes; Manuel, David Lazcano, Javier Banda-Leal and Ian Recchio 2014. Notes on Mexican Herpetofauna 22: Herpetofauna of the Continental Portion of the Municipality of Hermosillo, Sonora, Mexico. Bull. Chicago Herp. Soc. 49(8):105-115 - get paper here
  • Peralta-García A, Valdez-Villavicencio JH, Fucsko LA, Hollingsworth BD, Johnson JD, Mata-Silva V, Rocha A, DeSantis DL, Porras LW, and Wilson LD. 2023. The herpetofauna of the Baja California Peninsula and its adjacent islands, Mexico: composition, distribution, and conservation status. Amphibian & Reptile Conservation 17(1&2): 57–142
  • Skubowius, Bernd 2012. Echsen, Schildkröten, Amphibien und Spinnentiere – die vielfältige Welt der „creeping crawlers“ in Arizona. Draco 13 (50): 70-78 - get paper here
  • Sullivan, B.K., A.K. Owens, K.O. Sullivan & E.A. Sullivan 2016. Spatial Ecology of Sonoran Desert Tortoises (Gopherus morafkai): I. Fidelity in Home Range, Refuge Use and Foraging Behavior Journal of Herpetology 50 (3): 509-519. - get paper here
  • Sullivan, B.K., and E.A. Sullivan. 2015. Winter basking by hatchling Sonoran Desert Tortoises, Gopherus morafkai. Sonoran Herpetologist 28 (1):10-11. - get paper here
  • Sullivan, Brian K. & Thomas M. Cahill 2019. Seasonal Timing of Consumption of Calcium-Rich Caliche in the Sonoran Desert Tortoise (Gopherus morafkai) in Central Arizona. Chelonian Conservation and Biology Jun 2019 Vol. 18, No. 1: 98-101 - get paper here
  • Sullivan, Brian K. and Elizabeth A. Sullivan 2018. Late Spring Mining of Minerals by Female Sonoran Desert Tortoises. Sonoran Herpetologist 31 (3): 56-57 - get paper here
  • Sullivan, Brian K. and Elizabeth Sulivan. 2014. Gopherus morafkai (Sonoran desert tortoise) drinking behavior. Herpetological Review 45 (3): 483-484 - get paper here
  • Sullivan, Brian K. and Elizabeth Sullivan. 2016. Gopherus morafkai (Sonoran Desert Tortoise) Drinking behavior. Herpetological Review 47 (1): 123-124 - get paper here
  • Sullivan, Brian K.; Roy Averill-Murray, Keith O. Sullivan, Justin R. Sullivan, Elizabeth A. Sullivan, and J. Daren Riedle 2014. Winter Activity of the Sonoran Desert Tortoise (Gopherus morafkai) in Central Arizona. Chelonian Conservation and Biology Jul 2014, Vol. 13, No. 1: 114-119. - get paper here
  • TTWG [Peter Paul van Dijk, John B. Iverson, Anders G.J. Rhodin, H. Bradley Shaffer, and Roger Bour] 2014. Turtles of the World, 7th Edition: Annotated Checklist of Taxonomy, Synonymy, Distribution with Maps, and Conservation Status. 000.v7. Chelonian Research Monographs (ISSN 1088-7105) No. 5, doi:10.3854/crm.5.000.checklist.v7.2014 - get paper here
  • TTWG; Rhodin, A.G.J., Iverson, J.B., Bour, R., Fritz, U., Georges, A., Shaffer, H.B., and van Dijk, P.P. 2021. Turtles of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status (9th Ed.). In: Rhodin, A.G.J., Iverson, J.B., van Dijk, P.P., Stanford, C.B., Goode, E.V., Buhlmann, K.A., and Mittermeier, R.A. (Eds.). Chelonian Research Monographs 8:1–472. doi:10.3854/crm.8.checklist.atlas.v9.2021. - get paper here
  • Turner, D. S., Van Devender, T. R., Hale, S. F., Zach, R., Martínez, R., Van Devender, R. W., ... & Paholski, C. 2022. Amphibians and reptiles of Rancho Las Playitas area, Sonora, Mexico. Sonoran Herpetologist, 35, 50-59 - get paper here
  • Valverde, J. 2011. Die Schildkröten Mexikos. Reptilia (Münster) 16 (91): 16-23 - get paper here
 
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