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Testudinidae

Reptilia - Testudines - Testudinidae

Taxonomy
Testudinidae was named by Batsch (1788) [plastron and carapace]. It is extant. It was considered monophyletic by Joyce et al. (2004) and Vlachos (2018).

It was assigned to Testudinata by Gray (1869); to Cryptodira by Lydekker (1889); to Cryptodira by Hay (1902); to Cryptodira by Andrews (1906); to Cryptodiri by Jaekel (1911) and Perrier (1928); to Emydoidae by Hay (1930); to Chelonia by Owen (1842), Leidy (1873) and Stirton (1953); to Trionychoidea by Estes (1964); to Testudinata by Estes and Berberian (1970); to Testudines by Böhme (2008); to Pantestudinidae by Joyce et al. (2004) and Vlachos (2018); to Cryptodira by Cope (1871), Cope (1875), Gadow (1898), Lambe (1908), Kuhn (1946), Thurmond and Jones (1981), Meylan and Auffenberg (1986), Meylan and Sterrer (2000), Lapparent de Broin (2002), Hutchison et al. (2004), Staesche et al. (2007), Vlachos and Tsoukala (2014), Vlachos et al. (2014), Vlachos et al. (2015), Vlachos and Tsoukala (2016), Vlachos and Delfino (2016), Turvey et al. (2017) and de Lapparent de Broin et al. (2020); to Testudines by Frank and Ramus (1996), Manzano et al. (2009), Zacarias et al. (2013), Redkozubov et al. (2020), Villa et al. (2020) and Franz et al. (2020); and to Testudinoidea by Baur (1893), Kuhn (1966), Gaffney (1975), Antunes and de Broin (1977), Carroll (1988), Takahashi et al. (2003), Hervet (2003), Karl (2007), Pérez-García et al. (2013), van Dijk et al. (2014), Garcia et al. (2016), Pérez-García et al. (2017), de la Fuente et al. (2018), Vlachos (2021) and Perez-Garcia et al. (2021).

Synonyms
Synonymy list
YearName and author
1788Testudinidae Batsch
1830Testudinites sellovii Weiss p. 293
1835Testudo sellovii Fitzinger
1842Testudinidae Owen p. 160
1868Emys sombrerensis Leidy p. 180
1869Testudinidae Gray p. 166
1871Testudinidae Cope p. 236
1873Testudinidae Leidy p. 339
1875Testudinidae Cope p. 16
1875Testudo undata Cope p. 994
1875Testudo klettiana Cope p. 995
1875Testudo undata Cope p. 995
1884Testudo quadratus Cope
1886Caryoderma Cope
1887Testudo paranensis Scalabrini
1889Caryoderma Cope p. 662
1889Testudinidae Lydekker p. 71
1889Testudo formosa Moreno
1889Testudo sellouwi Moreno p. 29
1892Testudo pertenius Cope
1893Testudinidae Baur p. 675
1893Testudo laticaudata Cope
1893Testudo pertenuis Cope
1898Testudinidae Gadow p. 20
1898Xerobates undata Gilbert
1902Gopherus atascosae Hay
1902Testudo atascosae Hay
1902Testudinidae Hay p. 449
1902Testudo klettiana Hay p. 451
1902Testudo laticaudata Hay p. 451
1902Testudo pertenuis Hay p. 451
1902Testudo quadrata Hay p. 451
1902Testudo undata Hay p. 452
1903Stylemys calaverensis Sinclair
1906Testudinidae Andrews p. 277
1906Testudo peragrans Hay p. 15
1906Testudo exornata Lambe p. 187 figs. Pl. 3:1-3
1908Testudinidae Lambe
1908Testudo exornata Lambe
1911Testudinidae Jaekel p. 186
1916Gopherus praecedens Hay
1916Testudo luciae Hay
1916Testudo ocalana Hay
1916Testudo hayi Sellards
1917Testudo equicomes Hay p. 41
1923Testudo annae Hay p. 113
1923Testudo francisi Hay p. 116
1923Chelys patagonica Wieland
1928Testudinidae Perrier p. 3113
1929Testudo louisekressmani Wark
1930Testudinidae Hay p. 99
1930Gopherus atascosae Hay p. 101
1930Gopherus praecedens Hay p. 101
1930Stylemys calaverensis Hay p. 101
1930Testudo annae Hay p. 102
1930Testudo equicomes Hay p. 103
1930Testudo exornata Hay p. 103
1930Testudo francisi Hay p. 103
1930Testudo hayi Hay p. 103
1930Testudo klettiana Hay p. 103
1930Testudo laticaudata Hay p. 104
1930Testudo luciae Hay p. 104
1930Testudo ocalana Hay p. 104
1930Testudo peragrans Hay p. 104
1930Testudo pertenuis Hay p. 104
1930Testudo quadratus Hay p. 104
1930Testudo undata Hay p. 105
1935Testudo castrensis Bergounioux
1946Testudo quadratus Gilmore p. 309
1946Testudinidae Kuhn p. 58
1950Gopherus hayi Williams
1950Gopherus ocalana Williams p. 70
1953Testudinidae Stirton p. 612
1955Testudo semenensis Bergounioux
1957Geochelone luciae Ray
1957Geochelone ocalana Ray
1957Gopherus hayi Ray
1962Geochelone (Hesperotestudo) equicomes Auffenberg p. 630
1963Geochelone (Caudochelys) hayi Auffenberg p. 78
1963Geochelone (Hesperotestudo) equicomes Auffenberg p. 91
1963Geochelone (Caudochelys) laticaudata Auffenberg p. 93
1963Stylemys calaverensis Auffenberg p. 95
1964Stylemys calaverensis Auffenberg p. 323
1964Testudinidae Estes p. 99
1966Testudinidae Kuhn p. 26
1967Geochelone sombrerensis Auffenberg
1970Testudinidae Estes and Berberian p. 5
1974Chelonoidis sellowi Auffenberg
1974Chelonoidis sombrerensis Auffenberg
1975Testudinidae Gaffney p. 429
1977Testudinidae Antunes and de Broin
1977Hadrianus castrensis de Broin
1981Testudinidae Thurmond and Jones p. 123
1982Gopherus praecedens Bramble p. 854
1986Testudinidae Meylan and Auffenberg p. 280
1988Testudinidae Carroll
1996Testudinidae Frank and Ramus
2000Testudinidae Meylan and Sterrer
2002Testudinidae Lapparent de Broin
2003Testudinidae Hervet
2003Testudinidae Takahashi et al.
2004Testudinidae Hutchison et al. p. 42
2004Testudinidae Joyce et al.
2007Testudinidae Karl p. 56
2007Testudinidae Staesche et al.
2008Testudinidae Böhme p. 168
2009Testudinidae Manzano et al.
2013Testudinidae Pérez-García et al.
2013Testudinidae Zacarias et al.
2014Testudinidae Vlachos and Tsoukala
2014Testudinidae van Dijk et al.
2015Chelonoidis sombrerensis Rhodin et al.
2015Testudinidae Vlachos et al.
2016Testudinidae Garcia et al.
2016Testudinidae Vlachos and Delfino
2016Testudinidae Vlachos and Tsoukala
2017Testudinidae Pérez-García et al.
2017Testudinidae Turvey et al.
2018Testudinidae Vlachos
2018Testudinidae de la Fuente et al.
2020Testudinidae Franz et al.
2020Testudinidae Redkozubov et al.
2020Testudinidae Villa et al.
2020Testudinidae de Lapparent de Broin et al. p. 412
2021Testudinidae Perez-Garcia et al.
2021Testudinidae Vlachos

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RankNameAuthor
kingdomAnimalia()
Bilateria
EubilateriaAx 1987
Deuterostomia
phylumChordataHaeckel 1874
subphylumVertebrata
superclassGnathostomata
Osteichthyes()
subclassSarcopterygii()
subclassDipnotetrapodomorpha(Nelson 2006)
subclassTetrapodomorpha()
Tetrapoda
Reptiliomorpha
RankNameAuthor
Anthracosauria
subclassAmphibiosauriaKuhn 1967
Cotylosauria()
Amniota
Sauropsida
classReptilia
Testudinata(Oppel 1811)
orderTestudinesBatsch 1788
suborderCryptodira
Pantestudinoidea
superfamilyTestudinoidea
familyTestudinidaeBatsch 1788
familyTestudinidaeBatsch 1788

If no rank is listed, the taxon is considered an unranked clade in modern classifications. Ranks may be repeated or presented in the wrong order because authors working on different parts of the classification may disagree about how to rank taxa.

Fm. Testudinidae Batsch 1788
show all | hide all
G. †Achilemys Hay 1908
G. Aldabrachelys Loveridge and Williams 1957
+
Aldabrachelys gigantea Schweigger 1812 [Aldabra giant tortoise]
+
Invalid names: Testudo dussumieri Gray 1831 [synonym]
Invalid names: Dipsochelys Bour 1982 [synonym]
G. †Anhuichelys Yeh 1979
+
Anhuichelys doumuensis Tong et al. 2016
+
Invalid names: Anhuichelys xinzhouensis Chen 1983 [synonym]
G. Astrochelys Gray 1873
+
Astrochelys radiata Shaw 1802 [Radiated tortoise]
Astrochelys rogerbouri Kehlmaier et al. 2023
Astrochelys yniphora Vaillant 1885 [Ploughshare tortoise]
G. †Barnesia Karl 2018
G. Cachuga Gray 1855
+
Cachuga lineata Gray 1831
Cachuga tectum Gray 1831
G. †Cautleya Lydekker 1885
G. Chersina Gray 1831
+
Chersina angulata Schweigger 1812 [South African bowsprit tortoise]
G. †Cylindraspis Fitzinger 1835
+
Cylindraspis indica Schneider 1783
Cylindraspis inepta Gunther 1873
Cylindraspis peltastes Dumeril and Bibron 1835
G. Damonia Gray 1869
+
Damonia hamiltoni Gray 1831
G. †Eochersina Karl 2018
G. †Eochersis Karl 2018
+
Eochersis eiflaensis Karl 2018
G. †Floridemys Williams 1950
+
Floridemys hurdi Weems and George 2013
Floridemys nana Hay 1916
Floridemys nancei Weems 2023
Invalid names: Bystra Hay 1916 [replaced]
G. †Fontainechelon Pérez-García et al. 2016
+
Fontainechelon cassouleti Claude and Tong 2004
Tr. Geochelonini Swingland and Klemens 1989
+
G. Chelonoidis Fitzinger 1835
+
Chelonoidis abingdonii Gunther 1877 [Pinta giant tortoise]
Chelonoidis alburyorum Franz and Franz 2009
+
Chelonoidis alburyorum alburyorum Franz and Franz 2009
Chelonoidis alburyorum keegani Franz et al. 2020
Chelonoidis alburyorum sementis Franz et al. 2020
Chelonoidis australis Moreno 1889
+
Invalid names: Testudo praestans Rovereto 1914 [synonym]
Chelonoidis becki Rothschild 1901 [Volcan wolf giant tortoise]
Chelonoidis carbonarius Spix 1824 [Red-footed tortoise]
+
Invalid names: Geochelone atlantica López-Jurado et al. 1998 [synonym]
Chelonoidis chathamensis Van Denburgh 1907 [San Cristobal giant tortoise]
Chelonoidis chilensis Gray 1870 [Chaco tortoise]
Chelonoidis cubensis Leidy 1868
Chelonoidis darwini Van Denburgh 1907 [Santiago giant tortoise]
Chelonoidis denticulata Linnaeus 1766 [Yellow-footed tortoise]
Chelonoidis dominicensis Albury et al. 2018
Chelonoidis duncanensis Garman 1996 [Pinzon giant tortoise]
Chelonoidis gallardoi Rovereto 1914
Chelonoidis gersoni Viñola-López and Almonte 2022
Chelonoidis gringorum Simpson 1942
Chelonoidis hesterna Auffenberg 1971
Chelonoidis hoodensis Van Denburgh 1907 [Espanola giant tortoise]
Chelonoidis lutzae Zacarias et al. 2013
Chelonoidis marcanoi Turvey et al. 2017
Chelonoidis meridiana Vlachos et al. 2022
Chelonoidis monensis Williams 1952
Chelonoidis nigra Quoy and Gaimard 1824 [Floreana giant tortoise]
Chelonoidis petersi Freiberg 1973
Chelonoidis petrocellii Agnolin 2021
Chelonoidis phantastica Van Denburgh 1907 [Fernandina giant tortoise]
Chelonoidis porteri Rothschild 1903 [Santa Cruz giant tortoise]
Chelonoidis vicina Gunther 1875 [Southern Isabela giant tortoise]
G. †Titanochelon Pérez-García and Vlachos 2014
+
Titanochelon bacharidisi Vlachos et al. 2014
Titanochelon bolivari Hernandez-Pacheco 1917
+
Invalid names: Testudo richardi Bergounioux 1938 [synonym]
Titanochelon ginsburgi Broin 1977
Titanochelon kayadibiensis Karl et al. 2021
Titanochelon perpiniana Depéret 1885
Titanochelon schafferi Szalai 1931
Titanochelon schleichi Pappa et al. 2023
Titanochelon vitodurana Biedermann 1862
+
Invalid names: Testudo picteti Biedermann 1862 [synonym]
Subfm. Gopherinae Swingland and Klemens 1989
+
Invalid names: Gopherinii Swingland and Klemens 1989 [empty], Manourinii Swingland and Klemens 1989 [empty]
G. Gopherus Rafinesque 1832 [gopher tortoise]
+
Subg. †Gopherus (Gopherus) Rafinesque 1832
Gopherus agassizii Cooper 1863 [Mojave desert tortoise]
Gopherus auffenbergi Mooser 1972
+
Invalid names: Gopherus pargensis Mooser 1980 [synonym]
Gopherus berlandieri Agassiz 1857
Gopherus brevisternus Loomis 1909
Gopherus depressus Brattstrom 1961
+
Invalid names: Gopherus dehiscus Des Lauriers 1965 [synonym]
Gopherus donlaloi Reynoso and Montellano-Ballesteros 2004
Gopherus edae Hay 1906
+
Invalid names: Testudo hollandi Hay 1906 [synonym], Testudo pansa Hay 1908 [synonym], Testudo vaga Hay 1908 [synonym]
Gopherus hexagonatus Cope 1893
+
Invalid names: Bysmachelys canyonensis Johnston 1937 [synonym], Gopherus huecoensis Strain 1966 [synonym], Testudo campester Hay 1908 [synonym]
Gopherus laticuneus Cope 1873
Gopherus mohavensis Merriam 1919
Gopherus morafkai Murphy et al. 2011 [Sonoran desert tortoise]
Gopherus polyphemus Daudin 1802 [Florida gopher tortoise]
Invalid names: Bysmachelys Johnston 1937 [synonym], Gopherus (Oligopherus) Hutchison 1996 [invalid subgroup], Scaptochelys Bramble 1982 [synonym], Testudo milleri Brattstrom 1961 [nomen dubium], Xerobates Agassiz 1857 [synonym]
G. Homopus Duméril and Bibron 1834 [cape tortoise]
+
Homopus areolatus Thunberg 1787 [Parrot-beaked tortoise]
Homopus boulengeri Duerden 1906 [Karoo dwarf tortoise]
Homopus comptoni Owen and Bell 1849
Homopus femoralis Boulenger 1888 [Greater dwarf tortoise]
Homopus fenestratus Cooper and Broadley 1990
Homopus scutella Meyer 1845
Homopus signatus Gmelin 1789 [Speckled tortoise]
Homopus solus Branch 2007 [Nama tortoise]
G. †Juvemys Hervet 2003
+
Juvemys labarrerei Hervet 2003
G. †Kansuchelys Yeh 1963
+
Kansuchelys ovalis Yeh 1963
G. Malacochersus Lindholm 1929
+
Malacochersus tornieri Siebenrock 1903 [African pancake tortoise]
G. Manouria Gray 1852
+
Manouria emys Schlegel and Muller 1844 [Asian brown tortoise]
+
Invalid names: Geochelone emys Schlegel and Muller 1840 [synonym]
Manouria impressa Gunther 1882 [Impressed tortoise]
+
Invalid names: Geochelone impressa Gunther 1822 [synonym]
Manouria oyamai Takahashi et al. 2003
Manouria yushensis Yeh 1963
G. †Megalochelys Falconer and Cautley 1837
+
Megalochelys sivalensis Falconer and Cautley 1837
+
Invalid names: Cautleya annuliger Lydekker 1885 [synonym], Colossochelys atlas Falconer and Cautley 1844 [objective synonym], Testudo punjabiensis Lydekker 1889 [synonym]
G. †Miotestudo Deraniyagala 1967
+
Miotestudo ibba Deraniyagala 1967
G. †Nicoria Gray 1855
+
Nicoria tricarinata Blyth 1856
G. †Palaeoemys Schleich 1994
+
Palaeoemys hessiaca Schleich 1994
Palaeoemys occitana Hervet 2003
G. †Pelorochelon Pérez-García et al. 2016
+
Pelorochelon eocaenica Hummel 1935
Pelorochelon soriana Pérez-García et al. 2016
Invalid names: Asturichelys multicostatus Bergounioux 1957 [nomen dubium], Cheirogaster steinbacheri Karl 1996 [nomen dubium]
Subfm. Platysterninae Gray 1869
+
G. Platysternon Gray 1831 [big-headed turtle]
+
Platysternon megacephalum Danilov and Parham 2008
G. †Protestudo Chkhikvadze 1970
+
Protestudo bessarabica Riabinin 1918
G. Psammobates Fitzinger 1835
+
Psammobates antiquorum Broadley 1997
Psammobates geometricus Linneaus 1758 [Geometric tortoise]
Psammobates oculifer Kuhl 1820 [Serrated tortoise]
Psammobates tentorius Bell 1828 [Tent tortoise]
G. Pyxis Bell 1827
+
Pyxis arachnoides Bell 1827 [Common spider tortoise]
Pyxis planicauda Grandidier 1867 [Flat-shelled spider tortoise]
G. †Sinohadrianus Ping 1929
+
Sinohadrianus ezoensis Shikama 1953
G. †Solitudo Valenti et al. 2022
+
Solitudo gymnesica Bate 1914
Solitudo robusta Leith-Adams 1877
+
Invalid names: Testudo robustissima Tagliaferro 1913 [synonym], Testudo spratti Leith-Adams 1877 [synonym]
Solitudo sicula Valenti et al. 2022
Subfm. Testudininae Gray 1825
+
G. Agrionemys Khosatzky and Mlynarsky 1966
+
Agrionemys horsfieldii Gray 1844 [Central Asia tortoise]
G. Centrochelys Gray 1872 [spurred tortoise]
+
Centrochelys marocana Gmira et al. 2013
Centrochelys punica Arambourg 1979
Centrochelys sulcata Miller 1779 [African spurred tortoise]
Centrochelys vulcanica Lopez-Jurado and Mateo 1993
G. †Cheirogaster Bergounioux 1935
+
Cheirogaster gigas Bravard 1844
Cheirogaster maurini Bergounioux 1935
Invalid names: Taraschelon Pérez-García 2016 [synonym]
G. †Ergilemys Ckhikvadze 1972
+
Ergilemys bruneti Broin 1977
Ergilemys insolitus Matthew and Granger 1923
Ergilemys saikanensis Ckhikvadze 1972
Unr. †Geochelona Vlachos and Rabi 2018
+
G. †Alatochelon Pérez-García et al. 2020
+
Alatochelon myrteum Pérez-García et al. 2020
G. Geochelone Fitzinger 1835
+
Subg. Geochelone (Chelonoidis) Fitzinger 1835
Geochelone carbonaria Spix 1824 [red-footed tortoise]
Geochelone chilensis Gray 1870 [Chaco tortoise]
Geochelone costaricensis Segura 1944
Geochelone demissa Gilmore 1931
Geochelone elegans Schoepff 1794 [Indian star tortoise]
Geochelone elephantopus Harlan 1827 [Galapagos giant tortoise]
Geochelone gigantea Schweigger 1812 [Aldabra tortoise]
Geochelone kalganensis Gilmore 1931
Geochelone laetoliensis Meylan and Auffenberg 1987
Geochelone nigra Quoy and Gaimard 1824 [Galapagos tortoise]
Geochelone oskarkuhni Mlynarski 1969
Geochelone pardalis Bell 1828 [Leopard tortoise]
Geochelone platynota Blyth 1863 [Burmese star tortoise]
Geochelone radiata Shaw 1802 [Radiated tortoise]
Geochelone stromeri Meylan and Auffenberg 1986
Geochelone sulcata Miller 1779 [African spurred tortoise]
Geochelone travancorica Boulenger 1907
Geochelone yniphora Vaillant 1885 [angonoka]
Invalid names: Geochelone (Asterochelys) Gray 1873 [invalid subgroup], Geochelone (Geochelone) Fitzinger 1835 [invalid subgroup]
G. †Gigantochersina Chkhikvadze 1989
+
Gigantochersina ammon Andrews 1903
+
Invalid names: Testudo beadnelli Andrews 1906 [synonym]
G. †Impregnochelys Meylan and Auffenberg 1986
+
Impregnochelys pachytectis Meylan and Auffenberg 1986
G. Indotestudo Lindholm 1929
+
Indotestudo elongata Blyth 1853 [Elongate tortoise]
+
Invalid names: Geochelone elongata Blyth 1853 [synonym]
Indotestudo forstenii Schlegel and Muller 1844 [Travancore tortoise]
Indotestudo kaiseni Gilmore 1931
Indotestudo takasago Matsumoto 1929
Indotestudo travancorica Boulenger 1907
G. Kinixys Bell 1827 [hingeback tortoise]
+
Kinixys belliana Gray 1830 [Bells hinge-back tortoise]
Kinixys erosa Schweigger 1812 [forest hingeback tortoise]
Kinixys homeana Bell 1827 [Home's hingeback tortoise]
Kinixys lobatsiana Power 1927 [Lobatse hinge-back tortoise]
Kinixys natalensis Hewitt 1935 [Natal hinge-back tortoise]
Kinixys nogueyi Lataste 1886 [Western hinge-back tortoise]
Kinixys zombensis Hewitt 1931 [Southeastern hinge-back tortoise]
G. †Mesochersus de Lapparent de Broin 2003
+
Mesochersus orangeus de Lapparent de Broin 2003
G. †Namibchersus de Lapparent de Broin 2003
+
Namibchersus namaquensis Stromer 1926
G. †Paleotestudo Lartet 1851
+
Paleotestudo antiqua Bronn 1831
+
Invalid names: Testudo catalaunica Bataller 1926 [synonym]
Paleotestudo canetotiana Lartet 1851
Paleotestudo mellingi Peters 1868
Invalid names: Cheirogaster arrahonensis Bergounioux 1958 [nomen dubium], Paralichelys catalaunicus Bergounioux 1951 [nomen dubium]
G. Stigmochelys Gray 1873
+
Stigmochelys brachygularis Meylan and Auffenberg 1987
Stigmochelys crassa Andrews 1914
Stigmochelys pardalis Bell 1828 [Leopard tortoise]
G. †Stylemys Leidy 1851
+
Stylemys bottii de Stefano 1902
Stylemys capax Hay 1908
+
Invalid names: Stylemys conspecta Hay 1908 [synonym]
Stylemys frizaciana Lartet 1851
Stylemys gisellae Carbot-Chanona et al. 2022
Stylemys inusitata Hay 1906
+
Invalid names: Stylemys copei Koerner 1940 [synonym], Testudo primaeva Oelrich 1950 [synonym]
Stylemys karakolensis Riabinin 1927
Stylemys nebrascensis Leidy 1851
+
Invalid names: Emys culbertsonii Leidy 1852 [synonym], Emys hemispherica Leidy 1851 [synonym], Emys oweni Leidy 1851 [synonym], Gopherus neglectus Brattstrom 1961 [synonym], Testudo amphithorax Cope 1873 [synonym], Testudo emiliae Hay 1908 [synonym], Testudo lata Leidy 1851 [synonym], Testudo ligonius Cope 1873 [synonym]
Stylemys pygmea Lartet 1851
Invalid names: Stylemys oregonensis Leidy 1872 [nomen dubium], Testudo niobrarensis Leidy 1858 [nomen dubium]
Tr. Testudinini Batsch 1788
+
G. Acinixys Siebenrock 1902
G. Chersine Merrem 1820
+
Chersine khosatzkyi Redkozubov et al. 2020
Chersine lunellensis Almera and Bofill 1903
Invalid names: Eurotestudo Lapparent de Broin et al. 2006 [objective synonym]
G. Testudo Linnaeus 1758
+
Subg. Testudo (Chersine) Merrem 1820
Subg. †Testudo (Stylemys) Leidy 1851
Testudo amberiacensis Deperet 1894
Testudo amiatae Pantanelli 1893
Testudo antakyensis Perala 1996
Testudo antiqua Bronn 1831
Testudo atlas Falconer and Cautley 1844
Testudo bosporica Riabinin 1946
Testudo brevitesta Vlachos and Tsoukala 2016
Testudo brontops Marsh 1890
Testudo carinata Linnaeus 1758
Testudo cautleyi Lydekker 1889
Testudo cernovi Khosatzky 1948
Testudo changshanensis Wei et al. 1975
Testudo costarricensis Segura Paguaga 1944
Testudo craverii Portis 1879
Testudo denizoti Bergounioux 1935
Testudo despotti Szalai 1937
Testudo emys Schlegel and Müller 1840
Testudo escheri Pictet and Humbert 1856
Testudo eurysternum Gervais 1848
Testudo floweri Bodenheimer 1935
Testudo geometrica Linnaeus 1758
Testudo globosa Portis 1890
+
Invalid names: Testudo szalaii Mlynarski 1955 [synonym]
Testudo graeca Linnaeus 1758 [common tortoise]
Testudo hellenica Garcia et al. 2020
Testudo hermanni Gmelin 1789 [Hermann's tortoise]
Testudo hexagonata Cope 1893
Testudo hipparionum Wiman 1930
Testudo honanensis Wiman 1930
Testudo houzei Dollo 1909
Testudo hypercostata Wiman 1930
Testudo hypselonota Bourret 1941
Testudo ibera Pallas 1814 [Asia Minor spur-thighed tortoise]
Testudo indica Schneider 1783
Testudo kalksburgensis Toula 1896
Testudo kenitrensis Gmira 1993
Testudo kleinmanni Lortet 1883 [Egyptian tortoise]
Testudo lamanonii Gray 1831
Testudo larteti Pictet 1853
Testudo lohanica Perez-Garcia et al. 2022
Testudo lunanensis Yeh 1963
Testudo lutaria Linnaeus 1758
Testudo margae Hooijer 1948
Testudo marginata Schoepff 1792 [Marginated tortoise]
Testudo marmorum Gaudry 1862
Testudo nanus Gilmore 1931
Testudo neraudii Gray 1831
Testudo oughlamensis Gmira et al. 2013
Testudo pecorinii Abbazzi et al. 2008
Testudo promarginata Reinach 1900
Testudo pusilla Linnaeus 1758
Testudo scabra Linnaeus 1758
Testudo shansiensis Wiman 1930
Testudo sharanensis Yeh 1965
Testudo shensiensis Wiman 1930
Testudo shilouensis Hsu et al. 1973
Testudo sloanei Lydekker 1889
Testudo sphaerica Wiman 1930
Testudo steinheimensis Staesche 1931
Testudo tarakliensis Riabinin 1918
Testudo terrestris Forsskal 1775 [Middle Eastern spur-thighed tortoise]
Testudo ulanensis Gilmore 1931
Testudo yunnanensis Yeh 1963
Invalid names: Pseudotestudo Loveridge and Williams 1957 [synonym], Testudo marginata cretensis Bachmayer et al. 1975 [synonym]
Invalid names: Geoemydinei Theobald 1868 [empty]
Subfm. †Xerobatinae Agassiz 1857
+
G. †Hesperotestudo Williams 1950
+
Hesperotestudo bermudae Meylan and Sterrer 2000
+
Invalid names: Testudo sellardsi Hay 1916 [synonym]
Hesperotestudo ducateli Collins and Lynn 1936
Hesperotestudo harrisi Lichtig and Lucas 2022
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Invalid names: Geochelone nordensis Holman 1973 [synonym], Testudo rexroadensis Oelrich 1952 [synonym], Xerobates cyclopygius Cope 1878 [synonym]
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Invalid names: Caryoderma snovianum Cope 1886 [synonym], Testudo arenivaga Hay 1906 [synonym], Testudo farri Hay 1908 [synonym], Testudo impensa Hay 1908 [synonym]
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Invalid names: Geochelone alleni Auffenberg 1966 [synonym], Geochelone johnstoni Auffenberg 1962 [synonym], Geochelone mlynarskii Auffenberg 1988 [synonym], Geochelone oelrichi Holman 1972 [synonym], Testudo incisa Hay 1916 [synonym], Testudo riggsi Hibbard 1944 [synonym], Testudo wilsoni Milstead 1956 [synonym]
Hesperotestudo williamsi Auffenberg 1964
Invalid names: Caudochelys Auffenberg 1963 [synonym], Eupachemys Leidy 1877 [nomen dubium], Testudo tedwhitei Williams 1953 [nomen dubium]
Invalid names: Caryoderma Cope 1886 [nomen dubium], Chelys patagonica Wieland 1923 [nomen dubium], Emys sombrerensis Leidy 1868 [nomen dubium], Geochelone (Hesperotestudo) equicomes Hay 1917 [nomen dubium], Geocheloninae Swingland and Klemens 1989 [empty], Gopherus pertenius Cope 1892 [nomen dubium], Gopherus praecedens Hay 1916 [nomen dubium], Stylemys calaverensis Sinclair 1903 [nomen dubium], Testudininei de Broin and de la Fuente 1993 [empty], Testudinites sellovii Weiss 1830 [nomen dubium], Testudo annae Hay 1923 [nomen dubium], Testudo atascosae Hay 1902 [nomen dubium], Testudo castrensis Bergounioux 1935 [nomen dubium], Testudo cultrata Cope 1873 [nomen dubium], Testudo exornata Lambe 1906 [nomen dubium], Testudo formosa Moreno 1889 [nomen dubium], Testudo francisi Hay 1923 [nomen dubium], Testudo hayi Sellards 1916 [nomen dubium], Testudo klettiana Cope 1875 [nomen dubium], Testudo laticaudata Cope 1893 [nomen dubium], Testudo louisekressmani Wark 1929 [nomen dubium], Testudo luciae Hay 1916 [nomen dubium], Testudo ocalana Hay 1916 [nomen dubium], Testudo paranensis Scalabrini 1887 [nomen nudum], Testudo peragrans Hay 1906 [nomen dubium], Testudo quadratus Cope 1884 [nomen dubium], Testudo semenensis Bergounioux 1955 [nomen dubium], Testudo undata Cope 1875 [nomen dubium]
Diagnosis
ReferenceDiagnosis
E. Perrier 1928Tète complètement rétractile. Carapace recouverte par les plaques cornées typiques; 12 (rarement 11, par coalescence des gulaires) plaques cornées, en contact avec les marginales; pas de plaque intergulaire. Plastron formé par 9 plaques osseuses (un entoplastron) et recouvert par 12 plaques cornées; doigts distincts, articulés, 4 ou 5 ongles.
Measurements
No measurements are available
Composition: phosphaticsubp
Environment: terrestrialf
Locomotion: actively mobilec
Life habit: ground dwellingf
Diet: herbivoref
Diet 2: herbivoref
Reproduction: oviparousf
Comments: life habit and diet based on Ernst and Barbour 1989 and matches Uetz 2005 f
Created: 2005-08-26 07:16:55
Modified: 2005-09-09 16:16:46
Source: f = family, c = class, subp = subphylum
References: Hendy et al. 2009, Ernst and Barbour 1989, Carroll 1988

Age range

Maximum range based only on fossils: base of the Albian to the top of the Holocene or 113.00000 to 0.00000 Ma
Minimum age of oldest fossil (stem group age): 100.5 Ma

Collections (1196 total)

Oldest occurrences

Time interval Ma Country or state Original ID and collection number
Albian113.0 - 100.5Tunisia (Tataouine) Testudinidae indet. (116299)
Early/Lower Cenomanian99.6 - 93.5Morocco (Drâa-Tafilalet) Testudinidae indet. (164948)
Early/Lower Paleocene66.0 - 61.7China (Anhui) Anhuichelys siaoshihensis (132946 140377 179035)
Paleogene66.0 - 23.03Argentina (Chubut) Testudinidae indet. (208489)
Tertiary66.0 - 2.588China (Hubei) Testudo kalganensis (192670)
Cenozoic66.0 - 0.0Brazil (Acre) Testudinidae indet. (217645)
Middle Paleocene61.7 - 58.7China (Anhui) Anhuichelys doumuensis (179041 179042) Anhuichelys sp. (132945 179044) Anhuichelys sp., Anhuichelys tsienshanensis (179025 179038) Anhuichelys tsienshanensis (132942 179036 179037 179039 179040) Anhuichelys tsienshanensis, Anhuichelys doumuensis (132939)
Middle Paleocene61.7 - 58.7China (Hubei) Anhuichelys xinzhouensis (178873)
Ypresian56.0 - 47.8United Kingdom (England) Homopus comptoni (167274)
Ypresian56.0 - 47.8France (Champagne-Ardenne) Palaeoemys sp., Hadrianus sp. (73387)
Ypresian56.0 - 47.8France (Aude) Palaeoemys sp., Achilemys cassouleti (182641) Testudininei indet., Palaeoemys testudiniformis, Palaeoemys occitana, Juvemys labarrerei, Achilemys cassouleti (78236)
Ypresian56.0 - 47.8France (Marne) Hadrianus sp. (35539 35540 181653)
Ypresian - Lutetian56.0 - 41.3France (Occitanie) Testudinidae indet. (195457)
Eocene56.0 - 33.9Spain (Aragon) Testudo sp. (39017)
Eocene56.0 - 33.9Spain (Catalonia) Testudo sp. (38980 39015 39016)
Wasatchian55.8 - 50.3Canada (Northwest Territories) Geochelone sp. (15583)
MP 1055.8 - 48.6France (Picardie) Hadrianus sp. (68440)
MP 1148.6 - 40.4Germany (Hessen) Juvemys sp. (120036) Juvemys sp., Testudo sp. (22701)
MP 1348.6 - 40.4Germany (Eifel) Palaeoemys hessiaca, Barnesia eocaenica, Eochersis eiflaensis, Eochersina steinbacherae (24201)
Irdinmanhan48.6 - 37.2China (Nei Mongol) Testudo sp. (182111)