Hesperotestudo

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Superphylum: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Megaclassis: Osteichthyes
Cladus: Sarcopterygii
Cladus: Rhipidistia
Cladus: Tetrapodomorpha
Cladus: Eotetrapodiformes
Cladus: Elpistostegalia
Superclassis: Tetrapoda
Cladus: Reptiliomorpha
Cladus: Amniota
Classis: Reptilia
Cladus: Eureptilia
Cladus: Romeriida
Subclassis: Diapsida
Cladus: Sauria
Cladus: Archelosauria
Division: Pan-Testudines
Division: Testudinata
Ordo: Testudines
Subordo: Cryptodira
Superfamilia: Testudinoidea

Familia: Testudinidae
Genus: † Hesperotestudo
Species: † H. alleni – † H. angusticeps – † H. annae – † H. bermudae – † H. brontops – † H. campester – † H. equicomes – † H. impensa – † H. incisa – † H. johnstoni – † H. kalganensis – † H. niobrarensis – † H. orthopygia – † H. osborniana – † H. percrassa – † H. riggsi – † H. tumidus – † H. turgida – † H. wilsoni
Name

Hesperotestudo Williams, 1950
Type Species: Hesperotestudo osborniana (Hay, 1904), by original designation

References

Williams, E.E. 1950. "Testudo cubensis and the evolution of western hemisphere tortoises." Bulletin of the American Museum of Natural History 95: 1–36. Full text
Meylan, P.A., Sterrer, W. 2000. "Hesperotestudo (Testudines: Testudinidae) from the Pleistocene of Bermuda, with comments on the phylogenetic position of the genus." Zoological Journal of the Linnean Society 128 (1): 51–76. DOI: 10.1111/j.1096-3642.2000.tb00649.x
Turtle Extinctions Working Group (Rhodin, A.G.J., Thomson, S., Georgalis, G., Karl, H.-V., Danilov, I.G., Takahashi, A., de la Fuente, M.S., Bourque, J.R., Delfino, M., Bour, R., Iverson, J.B., Shaffer, H.B., and van Dijk, P.P.). 2015. Turtles and tortoises of the world during the rise and global spread of humanity: first checklist and review of extinct Pleistocene and Holocene chelonians. Chelonian Research Monographs. 5(8):000e.1–66. Download
Turtle Taxonomy Working Group (Rhodin, A.G.J., Iverson, J.B., Bour, R., Fritz, U., Georges, A., Shaffer, H.B. & van Dijk, P.P.). 2017. Turtles of the World: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status (8th Ed.). Chelonian Research Monographs 7: 1–292. ISBN 978-1-5323-5026-9. DOI: 10.3854/crm.7.checklist.atlas.v8.2017. Paywall. Full article (PDF). Reference page.

Hesperotestudo ("Western turtle") is an extinct genus of tortoise native to North and Central America (ranging as far south as Costa Rica[1]) from the Early Miocene to the Late Pleistocene.[2] Species of Hesperotesudo varied widely in size, with a large undescribed specimen from the Late Pleistocene of El Salvador reaching 150 cm (4.9 ft) in carapace length, larger than that of extant giant tortoises.[3] Historically considered a subgenus of Geochelone, it is now considered to be distantly related to that genus. Its relationships with other tortoises are uncertain.[2] The exposed areas of the bodies of Hesperotesudo species were extensively covered with large dermal ossicles, which in life were covered in keratin. It has been suggested that species of Hesperotestudo were relatively tolerant of cold weather.[4] Hesperotestudo became extinct at the end of the Pleistocene and beginning of the Holocene co-incident with the arrival of the first humans in North America, and sites have been found where Hesperotestudo were butchered.[3]
Taxonomy

Species list is based on Vlachos, 2018[2]

† Hesperotestudo Williams 1950[5]
†Hesperotestudo bermudae Meylan and Sterrer 2000[6] Bermuda, Middle Pleistocene c. 310,000 years before present (YBP) - shell length c. 50 centimetres (1.6 ft)[3][7]
†Hesperotestudo crassiscutata (Leidy 1889)[8] Florida, Texas, Illinois, South Carolina, (possibly also El Salvador[1]) Middle-Late Pleistocene shell length c. 120–125 centimetres (3.94–4.10 ft)[3]
†Hesperotestudo ducateli (Collins and Lynn, 1936) Calvert Formation, Maryland, Middle Miocene (Langhian–Serravallian)
†Hesperotestudo gilbertii (Hay, 1899) Ogallala Formation, Kansas, Hemphillian, Late Miocene-Early Pliocene (Tortonian–Zanclean)
†Hesperotestudo orthopygia (Cope, 1878) (syn= Xerobates cyclopygius Cope, 1878 =Caryoderma snovianum Cope, 1886 = Testudo rexroadensis Oelrich, 1952 = Geochelone nordensis Holman, 1973) Kansas, California, Nebraska, Late Miocene-Pliocene shell length c. 120 centimetres (3.9 ft)[9]
†Hesperotestudo osborniana (Hay, 1905)(syn= Testudo arenivaga Hay, 1906 = Testudo farri Hay, 1908 = Testudo impensa Hay, 1908 = Testudo orthopygia angusticeps Matthew, 1924) Colorado, Nebraska, Montana, Early Miocene-Early Pliocene shell length up to 92 centimetres (3.02 ft)
†Hesperotestudo turgida (Cope, 1892) (syn = Testudo incisa Hay, 1916a = Testudo riggsi Hibbard, 1944 = Testudo wilsoni Milstead, 1956 = Geochelone johnstoni Auffenberg 1962 = Geochelone alleni Auffenberg, 1966 = Geochelone oelrichi Holman, 1972a = Geochelone mlynarskii Auffenberg, 1988) Florida, Oklahoma, Kansas, Texas, Nebraska, Late Miocene-Late Pleistocene shell length c. 22–25 centimetres (0.72–0.82 ft)
†Hesperotestudo williamsi (Auffenberg, 1964) Oakville Formation, Texas, Early Miocene (Burdigalian) shell length c. 33.4 centimetres (1.10 ft)

References

Carbot-Chanona, Gerardo; Rivera-Velázquez, Gustavo; Jiménez-Hidalgo, Eduardo; Reynoso, Víctor Hugo (2020-11-04). "The fossil record of turtles and tortoises (Testudines) of Mexico, Central America and the Caribbean Islands, with comments on its taxonomy and paleobiogeography: a bibliographic review". Revista Mexicana de Ciencias Geológicas. 37 (3): 269–283. doi:10.22201/cgeo.20072902e.2020.3.1581. ISSN 2007-2902.
Vlachos, Evangelos (2018-04-17). "A Review of the Fossil Record of North American Turtles of the Clade Pan-Testudinoidea". Bulletin of the Peabody Museum of Natural History. 59 (1): 3. doi:10.3374/014.059.0101. ISSN 0079-032X.
Rhodin, A.G.J.; Thomson, S.; Georgalis, G.; Karl, H.-V.; Danilov, I.G.; Takahashi, A.; de la Fuente, M.S.; Bourque, J.R.; Delfino M.; Bour, R.; Iverson, J.B.; Shaffer, H.B.; van Dijk, P.P.; et al. (Turtle Extinctions Working Group) (2015). "Turtles and tortoises of the world during the rise and global spread of humanity: first checklist and review of extinct Pleistocene and Holocene chelonians" (PDF). Chelonian Research Monographs. 5 (8): 000e.1–66. doi:10.3854/crm.5.000e.fossil.checklist.v1.2015. Archived (PDF) from the original on 21 September 2022.
Moll, Don; Brown, Lauren E. (2017). "Reinterpretation of the Climatic Adaptation of Giant Fossil Tortoises in North America". Herpetological Journal. 27 (3): 276–286.
Williams , E.E. 1950. Testudo cubensis and the evolution of Western Hemisphere tortoises. Bulletin of the American Museum of Natural History 95:1–36.
Meylan, P. A.; Sterrer, W. (January 2000). "Herperotestudo (Testudines: Testudinidae) from the Pleistocene of Bermuda, with comments on the phylogenetic position of the genus". Zoological Journal of the Linnean Society. 128 (1): 51–76. doi:10.1111/j.1096-3642.2000.tb00649.x.
Olson, S. L.; Meylan, P. A. (December 2009). "A Second Specimen of the Pleistocene Bermuda Tortoise, Hesperotestudo bermudae Meylan and Sterrer". Chelonian Conservation and Biology. 8 (2): 211–212. doi:10.2744/CCB-0766.1. S2CID 85595118. Retrieved 2012-04-12.
Leidy, J. 1889. Description of vertebrate remains from Peace Creek, Florida. Transactions of the Wagner Free Institute of Science of Philadelphia 2:19–31.
Biewer, Jacob; Sankey, Julia; Hutchison, Howard; Garber, Dennis (2016-03-08). "A fossil giant tortoise from the Mehrten Formation of Northern California". PaleoBios. 33. doi:10.5070/p9331030312. ISSN 2373-8189.