After Carroll, 1988, and Maylan & Starrer, 2000, with Recent species from Uetz, Etzold & Chenna, 1995-2000 [“The Reptile Database”]
<==o Testudinidae Batsch, 1788 (non Gray, 1825) (tortoises; maakilpikonnat) |?-o †Achilemys Hay, 1908 | `-- †A. cassouleti Claude & Tong, 2004 |?- †Floridemys [Bystra] Mioc. NA. |?-o †Hadrianus Cope, 1872 [Testudo??] Eoc. NA. | `-- †H. majusculus |?- †Impregnochelys L. Mioc. EAf. |?- †Kansuchelys U. Mioc. or ?Olig. EAs. |?- †Sinohadrianus Eoc. EAs. |?- †Dithyrosternon valdense Pictet & Humbert, 1855-57 |?- Acinixys planicauda `--+--o Manouria ?Gray, 1854 [Geochelone (Manouria)] | |-- M. emys (Schlegel & Müller, in Temmink, 1844) [taxonomic history] | |-- M. phayrei (Blyth, 1854) [taxonomic history] | `-- M. impressa (Günther, 1882) [taxonomic history] `--+--o †Stylemys Leidy, 1851; Olig.-Mioc. NA. As. ?Eu. | |-- †S. nebrascensis Leidy, 1851 | `-- †S. amphitorax Cope, 1873 `--+--o Xerobatinae Gaffney & Maylan, 1988 | |--+-- †“Bamz turtoise” | | `?-o †Hesperotestudo Williams, 1950 | | |-- †H. bermudae Maylan & Starrer, 2000 Pleist. NA. | | |-- †H. osborniana (Hay, 1904) | | |--o †(Hesperotestudo) | | | |-- †H. (H.) alleni Auffenberg | | | `-- †H. (H.) turgida (Cope) | | `--o †(Caudochelys) | | |-- †H. (C.) ducateli (Collins & Lynn) | | |-- †H. (C.) tedwhitei (Williams) | | `-- †H. (C.) hayi (Sellards) | `--o “Gopherus” Rafinesque, 1832 sensu lato (kofferikilpikonnat) | |-- †Oligopherus laticuneus (Cope, 1873) Hutchison, 1996 [taxonomic history] | `--+--o Xerobates Agassiz, 1857 | | |-- X. agassizi Cooper, 1863 [taxonomic history] (Desert Tortoise) | | `-- X. berlandieri Agassiz, 1857 [taxonomic history] [taxonomic history] (Texas Tortoise) | `--o Gopherus Rafinesque, 1832 | |-- †G. vagus Hay, 1908 [taxonomic history] | |-- †G. pansus Hay, 1908 [taxonomic history] | |-- †G. brevisternus Loomis, 1909 [taxonomic history] | |-- †G. canyonensis Johnston, 1937 [taxonomic history] | |-- †G. donlaloi Reynoso & Montellano-Ballesteros, 2004 [taxonomic history] | |-- G. flavomarginatus Legner, 1959 [taxonomic history] | `-- G. polyphemus (Daudin, 1802) [taxonomic history] (Gopher Tortoise; kofferikilpikonna) `--o Testunidinae? Batsch, 1788 |?- †“Ergilemys” bruneti Broin, 1977 |?-o †Ergilemys Chkhikvadze, 1972; Eoc.-Olig. EAs. | |-- †E. insolitus (Gilmore, 1931) [Testudo insolitus Gilmore, 1931] | `-- †E. insolitus saikanensis Chkhikvadze, 1972 |?-o †Cheirogaster Bergounioux, 1935 | `-- †C. perpiniana (Depéret, 1885) [Testudo perpiniana Depéret, 1885; Ergilemys perpiniana (Depéret) Chkhikvadze, 1972] |--o Indotestudo [Geochelone: (Indotestudo)] | |-- I. elongata (Blyth, 1853) | `-- I. travancorica |--o “Testudo” Linnaeus, 1758 sensu lato | |?- †“Testudo” promarginata Reinach, 1900 | |--+--o Agrionemys Khozatsky & Mlynarski, 1966 [Testudo Linnaeus, 1758, in partim; Testudinella Gray, 1870 (preoccupied)] (nelivarvaskilpikonnat) | | | |-- †A. turgaica (Riabinin, 1926) [“Testudo” turgaica Riabinin, 1926] | | | |-- †A. djetyogus | | | |-- †A. honanensis | | | |-- †A. kegenika | | | |-- †A. ranovi | | | |-- †A. tekessa | | | |-- †A. tunhuanensis | | | |-- A. horsfieldii (Gray, 1844) [Testudo horsfieldii Gray, 1844] (nelivarvaskilpikonna)] | | | |-- A. kazachstanica Chkhikvadze, 1988 | | | `-- A. rustamovi Chkhikvadze et al., 1990 | | `--o †Protestudo Chkhikvadze, 1970 [Testudo Linnaeus, 1758, in partim] [jr. synon. of Agrionemys??] | | |-- †P. alba | | |-- †P. bessarabica (Riabinin, 1915) [“Testudo” bessarabica Riabinin, 1915] | | |-- †P. derevskii | | |-- †P. hipparionum | | |-- †P. illiberalis | | |-- †P. karabastuzica | | |-- †P. lavrovi | | |-- †P. moldavica | | |-- †P. oskarkhuni | | `-- †P. punjabensis | |--+-- †Paleotestudo canetotiana (Lartet, 1851) | | |-- †“Testudo” antiqua Bronn, 1831 | | `--o Eurotestudo Lapparent de Broin, Bour, Parham & Perälä, 2006 [Testudo Linnaeus, 1758, in partim; Chersine Merrem, 1820; Medaestia Wussow, 1916] Plioc.-R. Eu. | | |-- †E. pyrenaica (Depéret & Donnezan, 1890) | | |-- †E. globosa (Portis, 1890) | | |-- †E. lunellensis (Almera & Bofill, 1903) | | |-- †E. szalai Mlynarski, 1955 | | |--o E. hermanni (Gmelin, 1789) [Testudo hermanni Gmelin, 1789] (espanjankilpikonna) | | | |-- E. h. hermanni | | | `-- E. h. robertmertensi | | |-- E. boettgeri (Mojsisovics, 1889) [Testudo boettgeri Mojsisovics, 1889] | | `-- E. hercegovinensis (Werner, 1899) [Testudo hercegovinensis Werner, 1899] | `--o Testudo Linnaeus, 1758 [Chersine Merrem, 1820; Medaestia Wussow, 1916] | |?- †T. semenensis Bergounioux, 1955 | |?- †T. kalganesis | |?- †T. bosborica | |?- †T. eldarica | |?- †T. bulcarica Amiranashvili, 2000 | |-- (Pseudotestudo) kleinmanni | `--o (Testudo) | |-- †T. (T.) kenitrensis Gmira, 1993 | |-- †T. (T.) marmorum Gaudry, 1862 | |-- T. (T.) antakyensis Perälä, 1996 | |--+-- T. (T.) marginata Schoepff, 1793 (mustakilpikonna) | | `-- †T. (T.) weissingeri Bour, 1995 | `--o T. (T.) graeca (kreikankilpikonna) | |-- T. (T.) g. graeca | |-- T. (T.) g. ibera | |-- T. (T.) g. terrestris | `-- T. (T.) g. zarudnui |--o “Geoemydines” | |?- †Namibchersus | |?- †Astrochelys yniphora | |-- Chersina angulata (Schweigger, 1812) Gray, 1831 [Testudo angulata Schweigger, 1812; Testudo bellii Gray, 1828; Chersina angulata Gray, 1831; Neotestudo angulata Hewitt, 1931; Goniochersus angulatus Mertens, Müller & Rust, 1934; Chersine angulata Hewitt, 1937; Goniocherus angulatus Mertens & Wermuth, 1955] | |--o Homopus Duméril & Bibron, 1835 | | |-- H. areolatus Thunberg, 1787 | | |-- H. boulengeri Duerden, 1906 | | |-- H. femoralis Boulenger, 1888 | | |-- †H. fenestratus Cooper & Broadley, 1990 | | `-- H. signatus | |--o Kinixys Bell, 1827 (kellokilpikonnat) | | |?- †K. natalensis Hewitt | | |-- K. erosa (Schweigger, 1812) | | |-- K. homeana | | `--o K. belliana Gray, 1831 | | |-- K. b. belliana | | `-- K. b. nogueyi | |-- Malacochersus tornieri | |--o Psammobates | | |-- †P. antiquorum Broadley, 1997 | | |-- P. geometricus | | |-- P. oculifer | | `--o P. tentorius (Bell, 1828) | | |-- P. t. tentorius | | |-- P. t. trimeni | | `-- P. t. verroxii | `-- Pyxis arachnoides (madagaskarinkilpikonna) `--o “Geochelonini” |?- †“Geochelone” burchardii (Ahl, 1925) [Plioc. Canary Islands] |?-o †Megalochelys Falconer & Cautley, 1837 [Colossochelys Falconer & Cautley, 1844; Testudo Murchison, 1868; Geochelone Hooijer, 1971] | |-- †M. atlas (Falconer & Cautley, 1844) [Megalochelys sivalensis Falconer & Cautley, 1837 (nomen nudum); Colossochelys atlas Falconer & Cautley, 1844] | |?- †M. gadowi | |-- †M. cautleyi | |-- †M. margae | `-- †M. sondaari |?-o †Cylindraspis [Geochelone: Cylindraspis] (Mascarenes “racing” tortoises) | |-- †C. indica (Reunion Giant Tortoise) [†1840] | |-- †C. peltastes (Rodrigues Domed Tortoise) [†1800] | |-- †C. vosmaeri (Rodrigues Giant Saddleback Tortoise) [†1800] | |-- †C. inepta (Mauritius Giant Domed Tortoise) [†1735] | `-- †C. triserrata (Mauritius Giant Flat-shelled Tortoise) [†1735] |--o Chelonoidis Fitzinger, 1835 [Geochelone (Chelonoidis) Fitzinger, 1835; Elephantopus Gray 1874] (eteläamerikanmaakilpikonnat?) | |-- †C. alburyorum | |-- †C. cubensis | |-- C. carbonarius (Spix 1824) (red-footed tortoise) | |-- C. chilens (Gray, 1870a) (Chaco tortoise) | |-- C. denticulatus (Linnaeus, 1766) (yellow-footed tortoise) | |-- †C. lutzae | |-- †C. marcanoi | |-- †C. monensis | |-- †C. sombrerensis | `--o C. niger -species complex (Galápagos Giant Turtoise; galapagoksenjättilaiskilpikonnat) | |-- †C. abingdonii (Günther, 1877) [Testudo ephippium Günther, 1874 (nomen nudum)] (Pinta Island tortoise) [species of ”Lonesome George”] | |-- C. becki (Rothschild, 1901) (Volcán Wolf Galápagos tortoise) | |-- C. chathamensis (Van Denburgh, 1907) (San Cristobal Island Galápagos tortoise) | |-- C. darwini (Van Denburgh, 1907) (Santiago Island Galápagos tortoise) | |-- C. donfaustoi Poulakakis, Edwards, and Caccone in Poulakakis, Edwards, Chiari, Garrick, Russello, Benavides, Watkins-Colwell, Glaberman, Tapia, Gibbs, Cayot, & Caccone, 2015 (Eastern Santa Cruz tortoise) | |-- C. duncanensis (Pritchard, 1996) [Testudo duncanensis Garman, 1917 (n.n.); Geochelone nigra duncanensis Pritchard, 1996] (Pinzon Island Galápagos tortoise) | |-- C. guntheri (Baur, 1889) (Sierra Negra giant tortoise) | |-- C. hoodensis (Van Denburgh, 1907) (Española Island Galápagos tortoise) | |-- C. microphyes (Günther, 1875a) (Volcán Darwin giant tortoise, Darwin Volcano giant tortoise, Tagus Cove giant tortoise) | |-- †C. niger (Quoy & Gaimard, 1824b) [Testudo nigra Quoy & Gaimard 1824b; Testudo galapagoensis Baur, 1889] (Floreana Island Galápagos tortoise) | |-- C. phantasticus (Van Denburgh, 1907) (Fernandina Island Galápagos tortoise) | |-- C. porteri (Rothschild, 1903) (Santa Cruz Island Galápagos tortoise) | |-- C. vandenburghi (DeSola, 1930) (Volcán Alcedo giant tortoise, Alcedo Volcano giant tortoise) | `-- C. vicina (Günther, 1875a) (Isabela Island Galápagos tortoise) `--+--o Geochelone [Geochelone: (Geochelone)] | |?- †G. orthopygia (Cope, 1878) | |-- G. (G.) elegans | |-- G. (G.) pardalis (pantterikilpikonna) | |-- G. (G.) platynota | `-- G. (G.) sulcata `--+--o (Asterochelys) [Geochelone: (Asterochelys)] | |-- †A. abrupta | |-- A. radiata | `-- A. yniphora `--o Aldabrachelys [Geochelone: (Aldabrachelys)](aldebaranjättiläiskilpikonnat) |-- A. gigantea (aldebaranjättiläiskilpikonna) |-- †A. gouffei |-- †A. grandidieri `-- †A. sumeirei
Reference(s):
- Amiranashvili, N. G., 2000: Fossil and recent tortoises of Asia and Eastern Europe (Genera Testudo, Agrionemys, Protestudo)
–Journal of Vertebrate Paleontology: Vol. 20, #3, suppl. to #3, 26A. - Carroll, R. L., 1988: Vertebrate paleontology and evolution.
–W. H. Freeman and company, New York, 1988, 698 - Carroll, R. L., 1988: Appendix. 594-648.
in Carroll, R. L. 1988: Vertebrate paleontology and evolution.
–W. H. Freeman and company, New York, 1988, 698 - Holman, J. A., 1987: Herpetofauna of the Egelhoff site (Miocene: Barstovian) of North-Central Nebraska.
–Journal of Vertebrate Paleontology: Vol. 7, #2, pp. 109-120 - Lapparent de Broin, F. de, 2000: African chelonians from the Jurassic to the Present: Phases of development and preliminary catalogue of the fossil record.
–Palaeontologica Africana:Vol. 36, pp. 43-82 - de Lapparent de Broin, F., Bour, R., Parham, J. F. & Perälä, J., 2006: Eurotestudo, a new genus for the species Testudo hermanni Gmelin, 1789 (Chelonii, Testudinidae).
–Comptes Rendus de l'Académie des Sciencies, Paris: Palevol: Vol. 5, #6, pp. 803-811 [doi: 10.1016/j.crpv.2006.03.002] - de Lapparent de Broin, F., Bour, R. & Perälä, J., 2006: Morphological defenition of Eurotestudo (Testudinidae, Chelonii): first part.
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–Journal of Vertebrate Paleontology: Vol. 24, #4, pp. 822-837 - 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.)
in Rhodin, A.G.J., Iverson, J.B., van Dijk, P.P., Saumure, R.A., Buhlmann, K.A., Pritchard, P.C.H., & Mittermeier, R.A. (Eds.), 2017: Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group.
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–iNet: http://www.reptile-database.org/ - Zug, G. R., 2001: Turtles of the Lee Creek Mine (Pliocene: North Carolina). 203-218
in Ray, C. E. & Bohaska, D. J., (eds.) 2001: Geology an paleoecology of the Lee Creek Mine, North Carolina, III.
–Smithsonian Contributions to Paleobiology: Nr. 90, iii-365