Pair of male gopher tortoises Gopherus polyphemus duking it out, photographed by Daniel Parker.

Belongs within: Pantestuguria.
Contains: Testudininae.

The Testudinidae are the land tortoises, the most entirely terrestrial group of the Testudinata.

Synapomorphies (from Gaffney & Meylan 1988): Two or fewer phalanges in manus and pes; only four digits in pes; quadrate usually enclosing stapes; axillary and inguinal glands absent; slightly wide fissura ethmoidalis; cloacal bursae absent; coracoid blade wide; foramen nervi trigemini with maxillary aperture; trochanters of femur coalesced.

<==Testudinidae [Pantestudinidae]
    |  i. s.: Hadrianus Cope 1872CP03, C77
    |           |--H. corsonii Leidy 1871C77
    |           `--H. majusculus Hay 1904B93
    |         Cylindraspis Fitzinger 1835FB-P07
    |           |--C. indica (Schneider 1783)FB-P07 [=Geochelone indicaD81]
    |           |--C. inepta (Günther 1873)FS01, H81 [=Geochelone ineptaD81]
    |           |--C. peltastes (Duméril & Bibron 1835)FS01, H81 [=Geochelone peltastesD81]
    |           |--C. triserrata (Günther 1873)FS01, H81 [=Geochelone triserrataD81]
    |           `--C. vosmaeri (Fitzinger 1826)FB-P07 [=Geochelone vosmaeriD81]
    |  `--Hesperotestudo Williams 1950GM88, H96
    |       |--H. brontops (Marsh 1890) [=Testudo brontops]H96
    |       `--H. turgida (Cope 1870)H96
    `--+--Manouria Gray 1852FB-P07
       |    |--M. emys (Schlegel & Müller 1844) [=Testudo emys]JPG04
       |    |    |--M. e. emysFB-P07
       |    |    `--M. e. phayreiFB-P07
       |    `--M. impressa (Günther 1882)JPG04
            |--Stylemys Leidy 1851GM88, H96
            |    |--S. amphithorax (Cope 1873) (see below for synonymy)H96
            |    |--S. nebrascensis Leidy 1851 (see below for synonymy)H96
            |    `--‘Gopherus’ neglectus Brattstrom 1961H96
               `--Gopherus Rafinesque 1832FB-P07, H96 [incl. Bysmachelys Johnston 1937H96]
                    |  i. s.: G. agassizii (Cooper 1863)JPG04
                    |         G. berlandieri (Agassiz 1857)JPG04
                    |         G. flavomarginatus Legler 1959JPG04
                    |--*G. (Gopherus) polyphemus (Daudin 1801)H96, FB-P07
                    `--G. (Oligopherus Hutchison 1996)H96
                         `--G. (*O.) laticuneus (Cope 1873) (see below for synonymy)H96

Gopherus (*Oligopherus) laticuneus (Cope 1873) [=Testudo laticunea, Go. laticunea, Scaptochelys laticunea; incl. Testudo praeextans Lambe 1913, Gopherus praeextans, T. quadratus Cope 1884, T. thomsoni Hay 1908, Geochelone thompsoni (l. c.)]H96

Stylemys amphithorax (Cope 1873) [=Testudo amphithorax; incl. T. ligonius Cope 1873, Stylemys ligonia, T. (Hesperotestudo) ligonia]H96

Stylemys nebrascensis Leidy 1851 [incl. Emys culbertsonii Leidy 1851, E. hemispherica Leidy 1851, Testudo lata Leidy 1851, E. oweni Leidy 1851]H96

*Type species of generic name indicated


[B93] Benton, M. J. 1993. Reptilia. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 681–715. Chapman & Hall: London.

[CP03] Claude, J., E. Paradis, H. Tong & J.-C. Auffray. 2003. A geometric morphometric assessment of the effects of environment and cladogenesis on the evolution of the turtle shell. Biological Journal of the Linnean Society 79: 485–501.

[C77] Cope, E. D. 1877. Report upon the extinct Vertebrata obtained in New Mexico by parties of the expedition of 1874. Geographical Surveys West of the One Hundredth Meridian 4(2): i–iv, 1–370.

[D81] Day, D. 1981. The Doomsday Book of Animals: A unique natural history of three hundred vanished species. Ebury Press: London.

[FS01] Flannery, T., & P. Schouten. 2001. A Gap in Nature: Discovering the World’s Extinct Animals. Text Publishing: Melbourne.

[GM88] Gaffney, E. S., & P. A. Meylan. 1988. A phylogeny of turtles. In: Benton, M. J. (ed.) The Phylogeny and Classification of the Tetrapods vol. 1. Amphibians, Reptiles, Birds pp. 157–219. Clarendon Press: Oxford.

[H81] Honegger, R. E. 1981. List of amphibians and reptiles either known or thought to have become extinct since 1600. Biological Conservation 19: 141–158.

[H96] Hutchison, J. H. 1996. Testudines. In: Prothero, D. R., & R. J. Emry (eds) The Terrestrial Eocene–Oligocene Transition in North America pp. 337–353. Cambridge University Press.

[JPG04] Joyce, W. G., J. F. Parham & J. A. Gauthier. 2004. Developing a protocol for the conversion of rank-based taxon names to phylogenetically defined clade names, as exemplified by turtles. Journal of Paleontology 78 (5): 989–1013.

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