Hesperotestudo Temporal range: Early Miocene–Late Pleistocene PreꞒ Ꞓ O S D C P T J K Pg N

Shell of Hesperotestudo orthopygia

Skull of Hesperotestudo osborniana
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Reptilia
Order:	Testudines
Suborder:	Cryptodira
Superfamily:	Testudinoidea
Family:	Testudinidae
Genus:	†Hesperotestudo Williams, 1950
Type species
Hesperotestudo osborniana (Hay, 1904)
Species
See text

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 Hesperotestudo 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 Hesperotestudo 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 roughly co-incident with the arrival of the first humans in North America. There is apparently a site in Florida where one individual may have been killed that some suggested were evidence of butchering, although others suggested that the turtle was neither cooked nor does a ledge that was found near it date at the same time as it.^[3]^[5]^[6]

Taxonomy

Species list is based on Vlachos, 2018^[2]

† Hesperotestudo Williams 1950^[7]
- †Hesperotestudo bermudae Meylan and Sterrer 2000^[8] Bermuda, Middle Pleistocene c. 310,000 years before present (YBP) - shell length c. 50 centimetres (1.6 ft)^[3]^[9]
- †Hesperotestudo crassiscutata (Leidy 1889)^[10] 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)^[11]
- †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

^ ^a ^b 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.
^ ^a ^b ^c 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. S2CID 214641639.
^ ^a ^b ^c ^d 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.
^ Dunbar, James S.; Webb, S. David (1996). "17. Bone and Ivory Tools from Submerged Paleoindian Sites in Florida". In Anderson, David G.; Sassaman, Kenneth E. (eds.). The Paleoindian and Early Archaic Southeast. University Alabama Press. pp. 331–353.
^ Grayson, Donald K.; Meltzer, David J. (2003). "A requiem for North American overkill". Journal of Archaeological Science. 30 (5): 585–593. doi:10.1016/S0305-4403(02)00205-4.
^ 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.

Tortoise family

Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Subclass: Anapsida
Order: Testudines
Suborder: Cryptodira
Superfamily: Testudinoidea
Family: Testudinidae

Genera

Species of the tortoise family

Agrionemys

Russian tortoise

Chaco tortoise
Red-footed tortoise
Yellow-footed tortoise
†Chelonoidis alburyorum
†Chelonoidis cubensis
†Chelonoidis lutzae
†Chelonoidis monensis
†Chelonoidis sellovii
†Chelonoidis sombrerensis

Galápagos tortoise

Chelonoidis complex

†Cheirogaster

†Cheirogaster bacharidisi
†Cheirogaster gymnesica
†Cheirogaster schafferi

Chersina

Angulate tortoise

†Hadrianus

†Hadrianus corsoni
†Hadrianus majusculus
†Hadrianus octonaria
†Hadrianus robustus
†Hadrianus schucherti
†Hadrianus utahensis
†Hadrianus vialovi

†Hesperotestudo

†Hesperotestudo percrassa

Pancake tortoise

Leopard tortoise

†Stylemys

†Stylemys botti
†Stylemys calaverensis
†Stylemys canetotiana
†Stylemys capax
†Stylemys conspecta
†Stylemys copei
†Stylemys emiliae
†Stylemys frizaciana
†Stylemys karakolensis
†Stylemys nebrascensis
†Stylemys neglectus
†Stylemys oregonensis
†Stylemys pygmea
†Stylemys uintensis
†Stylemys undabuna

Testudo

Phylogenetic arrangement of turtles based on turtles of the world 2017 update: Annotated checklist and atlas of taxonomy, synonymy, distribution, and conservation status. Key: †=extinct.

Testudines

Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Clade: Diapsida
Order: Testudines

Genus

Chelonioidea
(Sea turtles)

Cheloniidae	†Allopleuron Caretta †Carolinochelys Chelonia †Eochelone Eretmochelys †Gigantatypus †Glarichelys †Itilochelys Lepidochelys †Mexichelys †Miocaretta Natator †Pacifichelys †Syllomus †Tasbacka
Dermochelyidae	†Arabemys †Corsochelys †Cosmochelys Dermochelys †Eosphargis †Mesodermochelys †Psephophorus
	†Euclastes †Peritresius †Procolpochelys †Protosphargis †Puppigerus

Kinosternoidea

Dermatemydidae	Dermatemys
Kinosternidae	Claudius †Hoplochelys Kinosternon Staurotypus Sternotherus

Testudinoidea

Emydidae	†Acherontemys Chrysemys Clemmys Deirochelys Emys Actinemys Emydoidea Glyptemys Graptemys Malaclemys Pseudemys Terrapene Trachemys †Wilburemys
Geoemydidae	Batagur †Banhxeochelys Cuora Cyclemys Geoclemys Geoemyda Hardella Heosemys Leucocephalon Malayemys Mauremys Melanochelys Morenia Notochelys Orlitia Pangshura Rhinoclemmys Sacalia Siebenrockiella Vijayachelys
Platysternidae	Platysternon
Testudinidae	Aldabrachelys Astrochelys Centrochelys Chelonoidis Chersina Cylindraspis †Cymatholcus †Floridemys Geochelone Gopherus †Hadrianus †Hesperotestudo Homopus Indotestudo Kinixys Malacochersus Manouria †Megalochelys †Oligopherus Psammobates Pyxis †Solitudo Stigmochelys †Stylemys Testudo

Trionychia

Carettochelyidae	†Allaeochelys †Anosteira Carettochelys
Trionychidae	Amyda Apalone †Axestemys Chitra Cyclanorbis Cycloderma †Drazinderetes Dogania †Gilmoremys †Hutchemys †Khunnuchelys Lissemys Nilssonia Palea †Palaeoamyda Pelochelys Pelodiscus Rafetus Trionyx
	†Basilochelys †Sinaspideretes

Chelydridae	Chelydra †Chelydrops †Chelydropsis †Emarginachelys †Macrocephalochelys Macrochelys †Planiplastron †Protochelydra
†Nanhsiungchelyidae	†Anomalochelys †Basilemys †Jiangxichelys
†Protostegidae	†Alienochelys †Archelon †Atlantochelys †Bouliachelys †Calcarichelys †Cratochelone †Desmatochelys †Iserosaurus †Notochelone †Ocepechelon †Pneumatoarthrus †Protostega †Rhinochelys †Santanachelys †Terlinguachelys
	†Adocus †Argillochelys †Bashuchelys †Ctenochelys †Prionochelys †Toxochelys

Pleurodira

†Araripemydidae	†Araripemys
†Bothremydidae	†Araiochelys †Arenila †Azabbaremys †Bothremys †Cearachelys †Chedighaii †Chupacabrachelys †Eotaphrosphys †Foxemys †Galianemys †Ilatardia †Inaechelys †Itapecuruemys †Jainemys †Kinkonychelys †Kurmademys †Labrostochelys †Nigeremys †Phosphatochelys †Polysternon †Puentemys †Rosasia †Rhothonemys †Sankuchemys †Taphrosphys †Ummulisani †Zolhafah
Chelidae	Acanthochelys Chelodina Chelus Elseya Elusor Emydura Hydromedusa †Lomalatachelys Mesoclemmys Myuchelys Phrynops Platemys †Prochelidella Pseudemydura Rheodytes Rhinemys †Yaminuechelys
Pelomedusidae	Pelomedusa Pelusios
Podocnemididae	†Albertwoodemys †Bauruemys †Brontochelys †Caninemys †Carbonemys †Cerrejonemys †Cordichelys Erymnochelys †Lapparentemys †Latentemys Peltocephalus Podocnemis †Stupendemys
†Sahonachelyidae	†Sahonachelys †Sokatra

	†Caribemys †Caririemys †Tacuarembemys

Phylogenetic arrangement of turtles based on Turtles of the World 2017 Update: Annotated Checklist and Atlas of Taxonomy, Synonymy, Distribution, and Conservation Status. † = extinct.

See also List of Testudines families