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Pantestudines

Osteichthyes

Taxonomy
Pantestudines was named by Joyce et al. (2004). It is extant. It was considered monophyletic by Joyce et al. (2004).

It was assigned to Reptilia by Joyce et al. (2004), Li et al. (2018); and to Archelosauria by Jenkins et al. (2026).

Subtaxa
Eorhynchochelys, Pappochelys, Testudinata
Sister taxa
Testudinata
Entered
by J. Tennant (authorized by P. Mannion) on 2012-12-19; modified by E. Vlachos on 2018-08-22

Synonymy list
YearName and author
2004Pantestudines Joyce et al.
2018Pantestudines Li et al.
2026Pantestudines Jenkins et al.

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RankNameAuthor
kingdomAnimalia()
—Bilateria
—EubilateriaAx 1987
—Deuterostomia
phylumChordataHaeckel 1874
subphylumVertebrata
superclassGnathostomata
classOsteichthyes
subclassSarcopterygii()
subclassDipnotetrapodomorpha(Nelson 2006)
subclassTetrapodomorpha()
—Tetrapoda
—Reptiliomorpha
RankNameAuthor
—Anthracosauria
subclassAmphibiosauriaKuhn 1967
—Cotylosauria()
—Amniota
—Sauropsida
classReptilia
subclassEureptilia()
—Romeriida
—Diapsida()
—Archosauromorpha(Huene 1946)
—ArchelosauriaCrawford 2015
—Pantestudines
—Pantestudines

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.

Unr. Pantestudines Joyce et al. 2004
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G. †Eorhynchochelys Li et al. 2018
+
†Eorhynchochelys sinensis Li et al. 2018
G. †Pappochelys Schoch and Sues 2015
+
†Pappochelys rosinae Schoch and Sues 2015
Unr. Testudinata Oppel 1811 [turtle]
+
G. †Amblypus Hitchcock 1858
+
†Amblypus dextratus Hitchcock 1858
G. †Ancyropus Hitchcock 1845
+
†Ancyropus heteroclitus Hitchcock 1841
+
Invalid names: Sauroidichnites jacksoni Hitchcock 1841 [synonym]
Fm. †Australochelyidae Gaffney and Kitching 1994
+
G. †Waluchelys Sterli et al. 2020
+
†Waluchelys cavitesta Sterli et al. 2020
G. †Australochelys Gaffney and Kitching 1994
+
†Australochelys africanus Gaffney and Kitching 1994
G. †Chengyuchelys Young and Chow 1953
+
†Chengyuchelys baenoides Young and Chow 1953
†Chengyuchelys latimarginalis Young and Chow 1953
G. †Emydhipus Fuentes Vidarte et al. 2003
Or. †Eunotosauria Watson 1917
+
Invalid names: Eunotosauridae Nopcsa 1923 [empty], Eunotosauroidea Nopcsa 1928 [empty]
G. †Helcura Hitchcock 1848
+
†Helcura anguinea Hitchcock 1858
†Helcura caudata Hitchcock 1858
†Helcura littoralis Hitchcock 1848
†Helcura surgens Hitchcock 1858
Unr. †Helochelydridae Nopcsa 1928
+
G. †Aragochersis Perez-Garcia et al. 2019
+
†Aragochersis lignitesta Perez-Garcia et al. 2019
G. †Helochelydra Nopcsa 1928
+
†Helochelydra anglica Lydekker 1889
†Helochelydra bakewelli Mantell 1833
†Helochelydra nopcsai Lapparent de Broin and Murelaga 1999
G. †Helochelys von Meyer 1854
+
†Helochelys danubina von Meyer 1854
G. †Naomichelys Hay 1908
+
G. †Plastremys Owen 1881
+
†Plastremys lata Owen 1881
†Plastremys rutteri Andrews 1920
Subfm. †Solemydinae Tong et al. 2023
+
G. †Solemys Lapparent de Broin and Murelaga 1996
+
†Solemys gaudryi Matheron 1869
†Solemys vermiculata Lapparent de Broin and Murelaga 1996
G. †Trachydermochelys Seeley 1869
+
G. †Tretosternon Owen 1842
Invalid names: Solemydidae Lapparent de Broin and Murelaga 1996 [objective synonym]
G. †Kallokibotion Nopcsa 1923
+
†Kallokibotion bajazidi Nopcsa 1923
+
Invalid names: Kallokibotion magnificum Nopcsa 1923 [synonym], Thalassodromeus sebesensis Grellet-Tinner and Codrea 2015 [synonym]
Unr. †Mesochelydia Joyce 2017
+
G. †Condorchelys Sterli 2008
+
†Condorchelys antiqua Sterli 2008
G. †Eileanchelys Anquetin et al. 2009
+
†Eileanchelys waldmani Anquetin et al. 2009
G. †Heckerochelys Sukhanov 2006
+
†Heckerochelys romani Sukhanov 2006
G. †Indochelys Datta et al. 2000
+
†Indochelys spatulata Datta et al. 2000
G. †Kayentachelys Gaffney et al. 1987
+
†Kayentachelys aprix Gaffney et al. 1987
Or. †Odontochelydia Karl 2012
+
Invalid names: Odontochelyidae Li et al. 2008 [empty]
G. †Palaeochersis Rougier et al. 1995
+
†Palaeochersis talampayensis Rougier et al. 1995
Unr. †Paracryptodira Gaffney 1975
+
G. †Arundelemys Lipka et al. 2006
+
†Arundelemys dardeni Lipka et al. 2006
Fm. †Baenidae Cope 1882
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Subfm. †Baeninae Cope 1882
+
Invalid names: Baenodd Gaffney 1972 [empty], Hayemdodd Gaffney 1972 [empty]
Unr. †Baenodda Gaffney 1972
+
G. †Arvinachelys Lively 2015
Tr. †Baenini Cope 1882
Unr. †Eubaeninae Williams 1950
Unr. †Palatobaeninae Gaffney 1972
G. †Saxochelys Lyson et al. 2019
G. †Scabremys Sullivan et al. 2012
G. †Edowa Adrian et al. 2023
+
†Edowa zuniensis Adrian et al. 2023
G. †Eubaena Hay 1908
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†Eubaena cephalica Hay 1904
†Eubaena hatcheri Hay 1901
G. †Gehennachelys Adrian et al. 2023
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†Gehennachelys maini Adrian et al. 2019
G. †Hayemys Gaffney 1972
+
†Hayemys latifrons Hay 1908
G. †Lakotemys Joyce et al. 2020
+
†Lakotemys australodakotensis Joyce et al. 2020
G. †Neurankylus Lambe 1902
+
†Neurankylus baueri Gilmore 1916
+
Invalid names: Neurankylus hutchisoni Lively 2016 [synonym], Neurankylus utahensis Lively 2016 [synonym]
†Neurankylus eximius Lambe 1902
+
Invalid names: Baena fluviatilis Parks 1933 [synonym], Charitemys captans Hay 1908 [synonym]
†Neurankylus lithographicus Larson et al. 2013
†Neurankylus notos Lichtig and Lucas 2016
†Neurankylus torrejonensis Lyson et al. 2016
Invalid names: Charitemys Hay 1908 [synonym]
G. †Peckemys Lyson and Joyce 2009
G. †Probaena Hay 1903
G. †Protobaena Joyce and Lyson 2015
+
†Protobaena wyomingensis Gilmore 1919
G. †Stygiochelys Gaffney and Hiatt 1971
G. †Thescelus Hay 1908
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†Thescelus insiliens Hay 1908
+
Invalid names: Baena longicauda Russell 1934 [synonym]
†Thescelus rapiens Hay 1908
G. †Trinitichelys Gaffney 1972
+
†Trinitichelys hiatti Gaffney 1972
Invalid names: Baena platyplastra Gilmore 1915 [nomen dubium], Hayemydinae Gaffney 1972 [empty], Thescelus hemispherica Gilmore 1935 [nomen dubium], Trinitichelyinae Gaffney 1972 [empty]
Superfm. †Baenoidea
+
Invalid names: Eubaenidae [empty], Neurankylidae [empty]
Fm. †Compsemydidae Pérez-García et al. 2015
+
G. †Berruchelus Pérez-García 2012
+
†Berruchelus russelli Pérez-García 2012
G. †Peltochelys Dollo 1884
+
G. †Tongemys Joyce et al. 2022
+
†Tongemys enigmatica Joyce et al. 2022
G. †Compsemys Leidy 1856
+
†Compsemys victa Leidy 1856
+
Invalid names: Compsemys parva Hay 1910 [synonym], Compsemys puercensis Gilmore 1919 [synonym], Compsemys torrejonensis Gilmore 1919 [synonym], Compsemys vafer Hay 1910 [synonym]
Invalid names: Emys obscurus Leidy 1856 [nomen dubium], Glyptops depressus Hay 1908 [nomen dubium]
Fm. †Pleurosternidae Cope 1868
+
G. †Ballerstedtia Karl et al. 2012
+
†Ballerstedtia typocardia Seeley 1869
G. †Desmemys Wegner 1911
G. †Dinochelys Gaffney 1979
+
†Dinochelys whitei Gaffney 1979
G. †Dorsetochelys Evans and Kemp 1976
+
+
Invalid names: Ballerstedtia bueckebergensis Karl et al. 2012 [synonym], Dorsetochelys delairi Evans and Kemp 1976 [synonym], Thalassemys ruetimeyeri Lydekker 1889 [synonym]
Invalid names: Dorsetochelys buzzops Bakker 1998 [nomen dubium]
G. †Glyptops Marsh 1890
+
†Glyptops ornatus Marsh 1890
+
Invalid names: Glyptops utahensis Gilmore 1916 [synonym]
Invalid names: Glyptops plicatulus Cope 1877 [nomen dubium]
G. †Pleurosternon Owen 1853
+
+
Invalid names: Megasternon koenigii Gray 1870 [synonym], Mesochelys durlstonensis Evans and Kemp 1975 [synonym], Pleurosternon concinnum Owen 1853 [synonym], Pleurosternon emarginatum Owen 1853 [synonym], Pleurosternon ovatum Owen 1853 [synonym], Pleurosternon oweni Seeley 1869 [synonym], Pleurosternon sedgwicki Seeley 1869 [synonym], Pleurosternon vansittarti Seeley 1869 [synonym], Pleurosternum portlandicum Lydekker 1889 [synonym]
†Pleurosternon moncayensis Pérez-Garcia et al. 2021
Invalid names: Digerrhum Cope 1870 [synonym], Mesochelys Evans and Kemp 1975 [synonym]
G. †Riodevemys Pérez-García et al. 2015
+
†Riodevemys inumbragigas Pérez-García et al. 2015
G. †Selenemys Pérez-García and Ortega 2011
+
†Selenemys lusitanica Pérez-García and Ortega 2011
G. †Toremys Pérez-García et al. 2015
+
†Toremys cassiopeia Pérez-García et al. 2015
G. †Uluops Carpenter and Bakker 1990
+
†Uluops uluops Carpenter and Bakker 1990
Invalid names: Desmemydinae Nopcsa 1928 [empty]
Invalid names: Desmemys bertelmanni Wegner 1911 [nomen dubium], Glyptops caelatus Hay 1908 [nomen dubium], Glyptops pervicax Hay 1908 [nomen dubium], Polythorax Cope 1876 [nomen dubium], Polythorax missuriensis Cope 1876 [nomen dubium], Probaena sculpta Hay 1903 [nomen dubium]
Unr. †Perichelydia Joyce 2017
+
Unr. †Meiolaniformes Sterli and de la Fuente 2013
+
G. †Chubutemys Gaffney et al. 2007
+
†Chubutemys copelloi Gaffney et al. 2007
Fm. †Meiolaniidae Lydekker 1887
+
G. †Gaffneylania Sterli et al. 2015
G. †Meiolania Owen 1886
G. †Ninjemys Gaffney 1992
G. †Niolamia Ameghino 1899
+
Invalid names: Crossochelys Simpson 1937 [synonym]
G. †Warkalania Gaffney et al. 1992
Invalid names: Gaffneylania Sterli et al. 2015 [valid], Meiolania Owen 1886 [valid], Miolaniidae [synonym], Miolaniidae [synonym], Ninjemys Gaffney 1992 [valid], Niolamia Ameghino 1899 [valid], Warkalania Gaffney et al. 1992 [valid]
G. †Otwayemys Gaffney et al. 1998
+
†Otwayemys cunicularius Gaffney et al. 1998
G. †Patagoniaemys Sterli and de la Fuente 2011
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†Patagoniaemys aeschyli Agnolin et al. 2026
†Patagoniaemys gasparinae Sterli and de la Fuente 2011
G. †Peligrochelys Sterli and de la Fuente 2013
+
†Peligrochelys walshae Sterli and de la Fuente 2013
G. †Trapalcochelys Sterli et al. 2013
+
†Trapalcochelys sulcata Sterli et al. 2013
Unr. †Sichuanchelyidae Tong et al. 2012
+
G. †Laurasichersis Pérez-García 2020
+
†Laurasichersis relicta Pérez-García 2020
G. †Mongolochelys Khosatzky 1997
+
†Mongolochelys efremovi Khosatzky 1997
G. †Sichuanchelys Ye and Pi 1997
+
†Sichuanchelys chowi Ye and Pi 1997
†Sichuanchelys palatodentata Joyce et al. 2016
G. †Spoochelys Smith and Kear 2013
+
†Spoochelys ormondea Smith and Kear 2013
G. †Platycheloides Haughton 1928
+
G. †Ypomonetikochelys Szczygielski and Dróżdż 2026
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†Ypomonetikochelys euryaspis Szczygielski and Dróżdż 2026
Invalid names: Triassochelyidae Abel 1919 [synonym]
G. †Proganochelys Baur 1887
+
+
Invalid names: Psammochelys keuperina Quenstedt 1889 [objective synonym], Stegochelys dux Jaekel 1914 [synonym]
†Proganochelys tenertesta Joyce et al. 2009
Invalid names: Psammochelys Quenstedt 1889 [synonym]
Fm. †Proterochersidae Nopcsa 1928
+
G. †Chinlechelys Joyce et al. 2009
G. †Cryptochersis Szczygielski and Dróżdż 2026
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†Cryptochersis paraxene Szczygielski and Dróżdż 2026
G. †Keuperotesta Szczygielski and Sulej 2016
G. †Proterochersis Fraas 1913
+
†Proterochersis limendorsa Szczygielski and Sulej 2016
†Proterochersis porebensis Szczygielski and Sulej 2016
†Proterochersis robusta Fraas 1913
+
Invalid names: Chelytherium obscurum Meyer 1863 [synonym], Chelytherium obscurum Meyer 1863 [synonym], Murrhardtia staeschei Karl and Tichy 2000 [synonym], Murrhardtia staeschei Karl and Tichy 2000 [synonym], Proterochersis intermedia Fraas 1913 [synonym], Proterochersis intermedia Fraas 1913 [synonym]
†Proterochersis robusta Fraas 1913
+
Invalid names: Chelytherium obscurum Meyer 1863 [synonym], Chelytherium obscurum Meyer 1863 [synonym], Murrhardtia staeschei Karl and Tichy 2000 [synonym], Murrhardtia staeschei Karl and Tichy 2000 [synonym], Proterochersis intermedia Fraas 1913 [synonym], Proterochersis intermedia Fraas 1913 [synonym]
Invalid names: Chelytherium Meyer 1863 [synonym], Murrhardtia Karl and Tichy 2000 [synonym]
G. †Thaichelys Szczygielski et al. 2025
+
†Thaichelys ruchae de Broin 1984
G. †Saltosauropus Bernier et al. 1984
+
†Saltosauropus latus Bernier et al. 1984
G. †Siamochelys Tong et al. 2002
+
†Siamochelys peninsularis Tong et al. 2002
Or. Testudines Batsch 1788 [turtle]
+
Subor. †Amphichelydia Lydekker 1889
+
Superfm. †Pleurosternoidea Hay 1930
+
Fm. †Kallokibotiidae Nopcsa 1923
Invalid names: Apertotemporalidae [empty]
Superfm. †Proganochelyoidea Baur 1887
+
Invalid names: Archaeochelydiidae Bergounioux 1938 [empty]
G. †Asturichelys Bergounioux 1957
G. †Chelonipus von Lilienstern 1939
+
†Chelonipus cuneiformis von Lilienstern 1939
†Chelonipus plieningeri Haubold 1971
†Chelonipus torquatus von Lilienstern 1939
Subor. Cryptodira Cope 1868
+
Fm. Chelonemydidae Rütimeyer 1872
+
Invalid names: Propleuridae Cope 1870 [synonym]
Superfm. Chelonioidea Baur 1893 [sea turtle]
+
Fm. Cheloniidae Bonaparte 1832
G. †Cimochelys Zangerl 1960
Fm. Dermochelyidae Lydekker 1889 [leatherback]
+
Invalid names: Psephophoridae Goette 1899 [synonym], Sphargidae Gray 1825 [synonym]
Unr. †Dermochelyoidae Williams 1950
Fm. †Desmatochelyidae Williston 1894
Fm. †Euclastidae France de Lapparent et al. 2025
G. †Kimurachelys Brinkman et al. 2015
G. †Leviathanochelys Castillo-Visa et al. 2022
G. †Nichollsemys Brinkman et al. 2006
Fm. †Protostegidae Cope 1873
Fm. †Toxochelyidae Baur 1895
Invalid names: Dollochelys rogovichi Averianov 2002 [nomen dubium], Pandermochelys Joyce 2004 [empty], Pliochelys Portis 1890 [nomen vanum], Progonosaurus Portis 1890 [nomen vanum], Turgaiscapha kushmurunica Averianov 2002 [nomen dubium]
G. †Chitracephalus Dollo 1884
+
†Chitracephalus dumonii Dollo 1885
+
Invalid names: Salasemys pulcherrima Fuentes et al. 2003 [synonym]
Invalid names: Salasemys Fuentes et al. 2003 [synonym]
Superfm. †Dermochelyoidea
+
G. †Pygmaeochelys Laube 1896
Invalid names: Chelone regularis Fritsch 1905 [nomen dubium], Pygmaeochelys michelobana Laube 1896 [nomen dubium]
Unr. Durocryptodira Danilov and Parham 2008
+
G. †Toxochelys Cope 1873
Unr. Eucryptodira Gaffney 1975
+
G. †Borealochelys Brinkman and Tarduno 2005
Superfm. Chelydroidea Baur 1893
G. †Enaliochelys Seeley 1869
G. †Hoyasemys Pérez-García et al. 2012
Superfm. †Macrobaenoidea Sukhanov 1964
Fm. †Sandownidae Tong and Meylan 2013
Fm. †Sinemydidae Yeh 1963
G. †Tienfucheloides Nessov 1978
Fm. †Xinjiangchelyidae Nessov 1990
Superfm. †Xinjiangchelyoidea Nessov 1990
Invalid names: Adocoidea Cope 1870 [empty], Angolachelonia Mateus et al. 2009 [empty], Centrocryptodira Gaffney and Meylan 1988 [empty], Peltochelyidae Seeley 1880 [empty], Plesiochelyoidea Baur 1888 [empty], Polycryptodira Gaffney and Meylan 1988 [empty]
G. †Galvechelone Pérez-García and Murelaga 2012
+
†Galvechelone lopezmartinezae Pérez-García and Murelaga 2012
Fm. Kinosternoidea Hay 1892 [mud turtle]
+
G. Sternotherus Gray 1825 [musk turtle]
Invalid names: Sternotherus Gray 1825 [valid]
Fm. †Nanhsiungchelyidae Yeh 1966
+
G. †Anomalochelys Hirayama et al. 2001
G. †Basilemys Hay 1902
G. †Bulganemys Sukhanov and Narmandakh 2006
G. †Hanbogdemys Sukhanov and Narmandakh 2006
G. †Jiangxichelys Tong and Mo 2010
G. †Kharakhutulia Sukhanov et al. 2008
G. †Nanhsiungchelys Yeh 1966
G. †Xianyuechelys Ke et al. 2024
G. †Yuchelys Tong et al. 2012
G. †Zangerlia Mlynarski 1972
Invalid names: Bulganemys Sukhanov and Narmandakh 2000 [nomen nudum], Charitonyx Chkhikvadze 1980 [nomen dubium], Charitonyx tajanikolaevae Chkhikvadze 1980 [nomen dubium], Hanbogdemysa Sukhanov 2000 [nomen nudum]
Unr. Panchelonioidea Joyce et al. 2004
+
Unr. Pancheloniidae Joyce et al. 2004
Invalid names: Sphenophyllum colombianum Huertas 2003 [nomen vanum]
Unr. †Pantestudinoidea Joyce et al. 2004
+
Superfm. Testudinoidea Gray 1825
G. †Polysternum Bergounioux 1935
+
Fm. Carettochelyidae Gill 1889
G. †Kappachelys Hirayama et al. 2013
Unr. †Pantrionychidae Joyce et al. 2004
Fm. Trionychidae Gray 1825 [softshell turtle]
+
G. Apalone Rafinesque 1832 [softshell turtle]
G. Aspideretes Hay 1904
G. Aspidonectes Wagler 1830
Subfm. †Chitrainae Jasinski et al. 2022
G. †Conchochelys Hay 1905
G. †Crassithecachelys Chkhikvadze 2000
Subfm. Cyclanorbinae Hummel 1929
G. †Eugenechelys Chkhikvadze 2000
G. †Hummelichelys Chkhikvadze 2000
Subfm. †Kuhnemydinae Jasinski et al. 2022
G. †Striatochelys Massonne et al. 2023
G. †Temnotrionyx Hay 1908
Subfm. Trionychinae Gray 1825 [softshell turtle]
G. †Zangerlichelys Chkhikvadze 2000
Invalid names: Altaytrionyx Chkhikvadze 2008 [nomen dubium], Altaytrionyx burtschaki Chkhikvadze 2008 [nomen dubium], Altaytrionyx devjatkini Chkhikvadze 2008 [nomen dubium], Altaytrionyx phiruzae Chkhikvadze 2008 [nomen dubium], Amyda menneri Chkhikvadze 1988 [nomen dubium], Aspideretes jaxarticus Riabinin 1938 [nomen nudum], Aspideretes muyuensis Lei and Ye 1985 [nomen dubium], Aspilus Portis 1885 [nomen dubium], Aspilus cortesii Portis 1885 [nomen dubium], Chitra minor Jaekel 1911 [nomen dubium], Lissemydinae Williams 1950 [empty], Paleotrionyx jimenezfuentesi Chkhikvadze 1995 [nomen dubium], Plastomenus gabunii Chkhikvadze 1984 [nomen dubium], Plastomenus jaxarticus Riabinin 1938 [nomen nudum], Plastomenus mlynarskii Chkhikvadze 1970 [nomen dubium], Platypeltis subcircularis Chow and Yeh 1957 [nomen dubium], Procyclanorbis Portis 1901 [nomen dubium], Rafetini Chkhikvadze 2000 [empty], Rafetus karkhualexandri Chkhikvadze 2000 [nomen nudum], Rafetus yexiangkuii Chkhikvadze 1999 [nomen dubium], Trionyx acutiformis Bergounioux 1935 [nomen dubium], Trionyx amansii Gray 1831 [nomen dubium], Trionyx anthracotheriorum Portis 1883 [nomen dubium], Trionyx aquitanicus Delfortrie 1869 [nomen dubium], Trionyx aspidiformis Laube 1900 [nomen dubium], Trionyx australiensis De Vis 1894 [nomen dubium], Trionyx austriacus Peters 1859 [nomen dubium], Trionyx borkenensis Gramann 1956 [nomen dubium], Trionyx bowerbanki Lydekker 1889 [nomen dubium], Trionyx bruxelliensis Winkler 1869 [nomen dubium], Trionyx burdigalensis Bergounioux 1935 [nomen dubium], Trionyx capellinii bulgaricus Khosatzky et al. 1983 [nomen dubium], Trionyx chaubeti Bergounioux 1935 [nomen dubium], Trionyx ciryi Bergounioux 1935 [nomen dubium], Trionyx cliftii Fitzinger 1835 [nomen nudum], Trionyx croaticus Koch 1915 [nomen dubium], Trionyx danovi Chkhikvadze 1989 [nomen dubium], Trionyx dodunii Gray 1831 [nomen nudum], Trionyx elongatus Liebus 1930 [nomen dubium], Trionyx erquelinnensis Dollo 1909 [nomen nudum], Trionyx hilberi Hoernes 1892 [nomen dubium], Trionyx ishiharaensis Miura and Uyama 1987 [nomen dubium], Trionyx italicus Schauroth 1865 [nomen dubium], Trionyx jakhimovitchae Chkhikvadze 1989 [nomen dubium], Trionyx kazusensis Otsuka 1969 [nomen dubium], Trionyx khosatzkyi Chkhikvadze 1983 [nomen dubium], Trionyx laurillardii Gray 1831 [nomen dubium], Trionyx lockardi Gray 1831 [nomen nudum], Trionyx lorioli Portis 1882 [nomen dubium], Trionyx manouri Gray 1831 [nomen dubium], Trionyx marini Hernández Sampelayo and Bataller 1944 [nomen dubium], Trionyx moldaviensis Khosatzky 1986 [nomen dubium], Trionyx mourieri Bergounioux 1935 [nomen dubium], Trionyx munzenbergensis Hummel 1927 [nomen nudum], Trionyx oligocenica Negri 1892 [nomen nudum], Trionyx oweni Reinach 1900 [nomen dubium], Trionyx parisiensis Gray 1831 [nomen dubium], Trionyx partschii Peters 1855 [nomen dubium], Trionyx pedemontana Portis 1879 [nomen dubium], Trionyx petersi trifailensis Teppner 1914 [nomen dubium], Trionyx pliocaenicus Reinach 1903 [nomen dubium], Trionyx pompignanensis Bergounioux 1933 [nomen dubium], Trionyx pontanus Laube 1895 [nomen dubium], Trionyx portisi Ristori 1891 [nomen dubium], Trionyx preschenensis Laube 1900 [nomen dubium], Trionyx propinquus Ristori 1891 [nomen dubium], Trionyx pseudovindobonensis Szalai 1934 [nomen dubium], Trionyx pustulatus Owen and Bell 1849 [nomen dubium], Trionyx ragusensis De Gregorio 1883 [nomen dubium], Trionyx reticulatus Rieppel 1979 [nomen dubium], Trionyx rocchettiana Portis 1882 [nomen dubium], Trionyx roncensis Harrassowitz 1919 [nomen dubium], Trionyx rotundiformis Bergounioux 1933 [nomen dubium], Trionyx sculptus Gilmore 1931 [nomen dubium], Trionyx senckenbergianus Reinach 1903 [nomen dubium], Trionyx senensis Ristori 1891 [nomen dubium], Trionyx siegeri Heritsch 1909 [nomen dubium], Trionyx sophiae Heritsch 1909 [nomen dubium], Trionyx stormsi Delheid 1899 [nomen dubium], Trionyx ubeensis Chitani 1925 [nomen dubium], Trionyx valdensis Portis 1882 [nomen dubium], Trionyx zakhidovi Khosatzky 1966 [nomen dubium], Zaisanonyx Chkhikvadze 2008 [nomen dubium]
Superfm. Trionychoidea Fitzinger 1826
+
Fm. †Adocidae Cope 1869
G. †Basilochelys Tong et al. 2009
Fm. Kinosternoidea Hay 1892 [mud turtle]
+
G. Sternotherus Gray 1825 [musk turtle]
Invalid names: Sternotherus Gray 1825 [valid]
G. †Sinaspideretes Young and Chow 1953
+
Invalid names: Yehguia Danilov and Parham 2006 [synonym]
Invalid names: Adocusia Danilov and Parham 2006 [empty], Carettochelyoidea Romer 1956 [empty], Chersidae Zittel 1890 [empty], Cryptoderinea Vaillant 1894 [empty], Dermatochelydidae [empty], Glyptopidae [empty], Glyptopsidae Marsh 1890 [empty], Indochelyidae Datta et al. 2000 [empty], Kallokibotioninae Nopsca 1923 [empty], Kayentachelyidae Gaffney et al. 1987 [empty], Kinosternia Gaffney and Meylan 1988 [empty], Kinosternoidae Holroyd and Hutchison 2002 [empty], Meiolaniidae Lydekker 1887 [empty], Pantrionychia Joyce et al. 2004 [empty], Platysternoidea Baur 1893 [empty], Selmacryptodira Gaffney et al. [empty], Tritosternidae Lydekker 1889 [empty]
G. †Entoplastrites Huene 1956
+
Fm. †Haichemydidae Sukhanov and Narmandakh 2006
+
G. †Haichemys Sukhanov and Narmandakh 2006
G. †Larachelus Pérez-García and Murelaga 2012
+
†Larachelus morla Pérez-García and Murelaga 2012
G. †Odontochelys Li et al. 2008
+
†Odontochelys semitestacea Li et al. 2008
Unr. Pancryptodira Joyce et al. 2004
+
G. †Owadowia Szczygielski et al. 2017
+
†Owadowia borsukbialynickae Szczygielski et al. 2017
Fm. †Thalassemydidae Zittel 1889
+
G. †Changisaurus Young 1959
G. †Phyllemys Schmidt 1944
G. †Sontiochelys Stache 1905
G. †Yaxartemys Riabinin 1948
Invalid names: Acichelyidae Lydekker 1889 [synonym], Emys beaugrandi Sauvage 1872 [nomen dubium], Trionyx primoevus Bergounioux 1937 [nomen dubium]
Unr. †Panpleurodira Danilov and Parham 2008
+
G. †Caribemys Fuente and Iturralde-Vinet 2001
Unr. †Dortokidae Lapparent de Broin and Murelaga 1996
+
G. †Dortoka Lapparent de Broin and Murelaga 1996
+
Invalid names: Ronella Gheerbrant et al. 1999 [synonym]
G. †Eodortoka Pérez-García et al. 2014
Invalid names: Muehlbachia Vremir and Codrea 2009 [nomen nudum], Muehlbachia nopcsai Vremir and Codrea 2009 [nomen nudum]
G. Pelomedusa Wagler 1830 [African helmeted turtle]
+
†Pelomedusa senutpickfordina de Lapparent de Broin 2008
Pelomedusa subrufa Bonaterre 1789 [Helmeted turtle]
+
G. †Notoemys Cattoi and Freiberg 1961
G. †Platychelys Wagner 1853
Subor. Pleurodira Cope 1865 [sideneck turtle]
+
Unr. †Bothremydidae Baur 1891
+
G. †Arenila Lapparent de Broin and Werner 1998
G. †Atolchelys Romano et al. 2014
G. †Azabbaremys Gaffney et al. 2001
Subfm. †Bothremydinae Gaffney 1975
Tr. †Bothremydini Gaffney et al. 2006
Unr. †Bothremydodda Gaffney et al. 2006
G. †Carteremys Williams 1953
Tr. †Cearachelyini Gaffney et al. 2006
G. †Cearachelys Gaffney et al. 2001
G. †Chedighaii Gaffney et al. 2006
G. †Elochelys Nopsca 1931
G. †Inaechelys de Araujo Carvalho et al. 2016
Tr. †Kurmademydini Gaffney et al. 2006
G. †Kurmademys Gaffney et al. 2001
G. †Labrostochelys Gaffney et al. 2006
G. †Nigeremys Broin 1977
+
Invalid names: Potamochelys Fitzinger 1843 [synonym]
G. †Palemys Gray 1870
+
Invalid names: Dacochelys Lydekker 1889 [synonym]
G. †Phosphatochelys Gaffney and Tong 2003
G. †Pleurochayah Adrian et al. 2021
G. †Rhothonemys Gaffney et al. 2006
G. †Rosasia Costa 1940
G. †Sankuchemys Gaffney et al. 2003
G. †Sindhochelys de Lapparent de Broin et al. 2021
Tr. †Taphrosphyini Gaffney et al. 2006
G. †Taphrosphys Cope 1869
+
Invalid names: Amblypeza Hay 1908 [synonym], Bantuchelys Dollo 1924 [synonym], Prochonias Cope 1869 [synonym]
G. †Ummulisani Gaffney et al. 2006
Invalid names: Kurmademydinae Gaffney et al. 2006 [empty], Podocnemis somaliensis Walker 1966 [invalid subgroup]
Fm. Chelidae Gray 1825
+
G. Acanthochelys Gray 1873
Unr. †Australochelida Joyce 2021
G. †Baalemys Sarda and Maniel 2024
G. †Birlimarr Megirian and Murray 1999
G. †Calvarichelys Oriozabala et al. 2025
Subfm. Chelinae Gray 1825
Subfm. Chelodininae Baur 1893
G. Chelus Duméril 1806
G. Elseya Gray 1867
+
Invalid names: Pelecomastes De Vis 1897 [synonym]
G. Emydura Bonaparte 1836
G. Hydromedusa Wagler 1830
Subfm. Hydromedusinae Baur 1893
G. †Mendozachelys de la Fuente et al. 2017
G. Myuchelys Thomson and Georges 2009
G. Phrynops Wagler 1830
+
Invalid names: Parahydraspis Wieland 1923 [synonym]
G. Platemys Wagner 1830
G. †Prochelidella Lapparent de Broin and de la Fuente 2001
G. Pseudemydura Siebenrock 1901
G. Rheodytes Legler and Cann 1980
G. †Rionegrochelys De la Fuente et al. 2017
G. †Yaminuechelys de la Fuente et al. 2001
Invalid names: Chelydidae Gray 1869 [synonym], Chelymys antiqua De Vis 1897 [nomen dubium], Chelymys arata De Vis 1897 [nomen dubium], Hydraspididae Cope 1868 [synonym], Hydraspis arenarius Rusconi 1934 [nomen dubium], Naiadochelys patagonica Staesche 1929 [nomen dubium], Pelecomastes ampla De Vis 1897 [nomen dubium], Platemys bergii Ambrosetti 1893 [nomen dubium], Platemys entreriana Ambrosetti 1893 [nomen dubium], Platemys holmbergii Ambrosetti 1893 [nomen dubium], Platemys mesopotamica Ambrosetti 1893 [nomen dubium], Platemys robusta Ambrosetti 1893 [nomen dubium], Platemys torrentium Burmeister 1885 [nomen dubium], Trionyx argentina Ameghino 1889 [nomen dubium]
Unr. Eupleurodira Gaffney and Meylan 1988
+
Invalid names: Bothremydinura Gaffney et al. 2006 [empty], Cheloides Gray 1825 [empty]
Unr. †Panchelidae Joyce et al. 2004
+
G. †Bonapartemys Lapparent de Broin and de la Fuente 2001
G. †Linderochelys de la Fuente et al. 2007
G. †Lomalatachelys Lapparent de Broin and de la Fuente 2001
G. †Palaeophrynops Lapparent de Broin and de la Fuente 2001
G. †Salamanchelys Bona 2006
Invalid names: Platemys guaranitica Ameghino 1889 [nomen dubium], Platemys patagonica Ameghino 1889 [nomen dubium]
Superfm. †Pelomedusoides Broin 1988
+
G. †Apodichelys Price 1954
Fm. †Araripemydidae Price 1973
G. †Caririemys Oliveira and Kellner 2007
Fm. †Euraxemydidae Gaffney et al. 2006
G. †Francemys Pérez-García 2019
G. †Itapecuruemys Batista et al. 2021
G. †Palaeaspis Gray 1870
Fm. †Peiropemydidae Gaffney et al. 2011
Fm. Pelomedusidae Cope 1868
+
Invalid names: Eusarkiidae Bergounioux 1952 [synonym], Peltocephalidae Gray 1870 [synonym], Sternothaeridae Cope 1868 [synonym]
Fm. Podocnemididae Cope 1868
Superfm. Podocnemidoidae Cope 1868
Superfm. †Podocnemidoidea Gaffney et al. 2006
Unr. †Sahonachelyidae Joyce et al. 2021
Invalid names: Chrysemys montolivensis Roman 1897 [invalid subgroup], Naiadochelys Hay 1908 [nomen dubium], Panpodocnemidae Joyce et al. 2004 [empty], Pelomedusera Gaffney et al. 2006 [empty], Taphrosphys dares Hay 1908 [nomen dubium]
Invalid names: Amphichelydidae Hay [empty], Megapleurodira Gaffney et al. 2006 [empty], Minipleurodira Gaffney et al. 2006 [empty], Notoemydidae Fernandez and de la Fuente 1994 [empty], Panpelomedusoides Joyce et al. 2004 [empty], Platychelira Gaffney et al. 2006 [empty]
Fm. †Sinochelyidae Chkhikvadze 1970
+
G. †Sinochelys Wiman 1930
+
†Sinochelys applanata Wiman 1930
+
Invalid names: Heishanemys imperfectus Bohlin 1953 [synonym], Peishanemys latipons Bohlin 1953 [synonym], Scutemys tecta Wiman 1930 [synonym]
Invalid names: Heishanemys Bohlin 1953 [synonym], Peishanemys Bohlin 1953 [synonym], Scutemys Wiman 1930 [synonym]
Invalid names: Peishanemydidae Nessov 1981 [synonym]
G. †Tacuarembemys Perea et al. 2014
+
†Tacuarembemys kusterae Perea et al. 2014
G. †Triassochelys Jaekel 1918
+
Invalid names: Stegochelys Jaekel 1914 [replaced]
Invalid names: Apertotemporalis Stromer 1934 [nomen dubium], Apholidemys Pomel 1847 [nomen dubium], Casichelydia Gaffney 1975 [empty], Chelone obovata Owen 1842 [nomen dubium], Chelone sopita Leidy 1865 [nomen vanum], Chelonia Latreille 1800 [synonym], Chelonii Latreille 1800 [synonym], Cryptrodiramorpha Lee 1995 [empty], Euclastochelys Bergounioux 1955 [nomen dubium], Euclastochelys interrupta Bergounioux 1955 [nomen dubium], Kallokibotionidae Gaffney and Meylan 1992 [empty], Meiolania damelipi White et al. 2010 [nomen dubium], Platychelone Dollo 1909 [nomen nudum], Platychelone emarginata Dollo 1909 [nomen nudum], Plesiochelydiae [empty], Podocnemis lata Ristori 1895 [invalid subgroup], Proganochelydia [empty], Senonemys suemegensis Bohn 1966 [nomen dubium], Sokotochelys Halstead 1979 [nomen dubium], Temnoclemmys cordiformis Bergounioux 1958 [nomen vanum], Trionyx fuchienensis Yeh 1974 [nomen dubium], Trionyx tshelkarensis Chkhikvadze 1973 [nomen nudum]
G. †Testudinites Weiss 1830
Fm. Testudofiexoolithidae Hirsch 1996
+
G. †Testudofiexoolithus Hirsch 1996
+
Fm. Testudoolithidae Hirsch 1996
+
G. †Testudoolithus Hirsch 1996
+
†Testudoolithus rigidus Hirsch 1996
G. †Tiantaioolithus Fang et al. 2003
+
†Tiantaioolithus jiangi Fang et al. 2003
Unr. †Thalassochelydia Anquetin et al. 2017
+
G. †Achelonia von Meyer 1860
+
†Achelonia formosa von Meyer 1860
Fm. †Eurysternidae Dollo 1886
+
G. †Chelonides Maack 1869
+
†Chelonides wittei Maack 1869
G. †Hydropelta Meyer 1850
+
†Hydropelta meyeri Tholliere 1851
G. †Idiochelys Meyer 1839
+
†Idiochelys fitzingeri Meyer 1839
+
Invalid names: Chelonemys ovata Jourdan 1862 [synonym], Chelonemys plana Jourdan 1862 [synonym], Idiochelys wagneri Meyer 1846 [synonym]
†Idiochelys wagneri Meyer 1846
G. †Palaeomedusa Meyer 1860
+
†Palaeomedusa testa von Meyer 1860
+
Invalid names: Eurysternum crassipes Wagner 1861 [objective synonym]
G. †Parachelys Meyer 1864
+
†Parachelys eichstaettensis von Meyer 1864
Invalid names: Plesiochelys minima Oertel 1915 [nomen dubium]
G. †Eurysternum Meyer 1839
+
†Eurysternum neuquinum Fernandez and de la Fuente 1988
†Eurysternum wagleri Meyer 1839
G. †Hylaeochelys Lydekker 1889
+
†Hylaeochelys belli Mantell 1844
+
Invalid names: Archaeochelys valdensis Lydekker 1889 [synonym], Hylaeochelys sollasi Nopcsa 1928 [synonym], Platemys dixoni Owen 1853 [synonym], Platemys mantelli Owen 1841 [synonym]
†Hylaeochelys kappa Pérez-García and Ortega 2014
+
Invalid names: Pleurosternon koeneni Grabbe 1884 [synonym]
Invalid names: Archaeochelys Lydekker 1889 [synonym]
G. †Jurassichelon Pérez-García 2015
+
†Jurassichelon moseri Bräm 1965
†Jurassichelon oleronensis Pérez-García 2015
G. †Neusticemys Fernandez and de la Fuente 1993
+
†Neusticemys neuquina Fernandez and de la Fuente 1988
Fm. †Plesiochelyidae Rütimeyer 1873
+
G. †Craspedochelys Rütimeyer 1873
+
†Craspedochelys jaccardi Pictet 1860
+
Invalid names: Plesiochelys choffati Sauvage 1898 [synonym]
†Craspedochelys passmorei Andrews 1921
†Craspedochelys picteti Rütimeyer 1873
+
Invalid names: Craspedochelys crassa Rütimeyer 1873 [synonym]
†Craspedochelys renzi Cadena et al. 2025
Invalid names: Emys dollfusii Lennier 1870 [nomen dubium]
G. †Globochelus de Lapparent de Broin et al. 2021
+
†Globochelus lennieri de Lapparent de Broin et al. 2021
G. †Plesiochelys Rütimeyer 1873
+
†Plesiochelys bigleri Puntener et al. 2017
†Plesiochelys etalloni Pictet and Humbert 1857
+
Invalid names: Plesiochelys langii Rütimeyer 1873 [synonym], Plesiochelys sanctaeverenae Rütimeyer 1873 [synonym], Plesiochelys solodurensis Rütimeyer 1873 [synonym], Stylemys hannoverana Maack 1869 [synonym], Stylemys lindenensis Maack 1869 [synonym]
†Plesiochelys normandicus Bergounioux 1937
Invalid names: Clemmys grayi Fitzinger 1835 [nomen oblitum], Emys hugi Gray 1831 [nomen oblitum], Emys hugii Gray 1831 [nomen oblitum], Emys jurensis Keferstein 1834 [nomen oblitum], Emys trionychoides Gray 1831 [nomen oblitum], Plesiochelys minor Portis 1878 [nomen dubium], Stegochelys Lydekker 1889 [synonym]
G. †Portlandemys Gaffney 1975
+
†Portlandemys gracilis Anquetin et al. 2015
†Portlandemys mcdowelli Gaffney 1975
G. †Tholemys Andrews 1921
G. †Yanduchelys Peng et al. 2005
+
†Yanduchelys delicatus Peng et al. 2005
Invalid names: Emys dutertrei Sauvage 1872 [nomen dubium], Hispaniachelys Slater et al. 2011 [nomen dubium], Hispaniachelys prebetica Slater et al. 2011 [nomen dubium], Stylemys lindenensis Maack 1869 [nomen dubium]
G. †Solnhofia Gaffney 1975
+
†Solnhofia brachyrhyncha Anquetin and Püntener 2020
†Solnhofia parsonsi Gaffney 1975
G. †Thalassemys Rütimeyer 1859
+
†Thalassemys bruntrutana Puntener et al. 2015
†Thalassemys carva Nessov 1984
†Thalassemys hugii Rütimeyer 1873
+
Invalid names: Eurysternum ignoratum Bräm 1965 [synonym], Thalassemys gresslyi Rütimeyer 1873 [synonym]
Invalid names: Parathalassemys Nessov and Krassevskaya 1984 [synonym]
G. †Tropidemys Rütimeyer 1873
+
†Pelobatochelys blakii Seeley 1875
†Tropidemys langii Rütimeyer 1873
+
Invalid names: Craspedochelys plana Rütimeyer 1873 [synonym], Tropidemys expansa Rütimeyer 1873 [synonym], Tropidemys gibba Rütimeyer 1873 [synonym]
†Tropidemys seebachi Portis 1878
Invalid names: Chelonia valanginiensis Pictet and Campiche 1862 [nomen dubium], Chelonides robusta Portis 1878 [nomen dubium]
Invalid names: Acichelys approximata Wagner 1856 [nomen dubium], Euryaspis radians Wagner 1861 [nomen dubium], Plesiochelys oblonga Oertel 1924 [nomen dubium], Plesiochelys pumilio Oertel 1924 [nomen dubium], Thalassemys heusseri Oertel 1924 [nomen dubium], Tropidemys morinica Sauvage 1880 [nomen dubium]
Subor. Thecophora Dollo
+
Invalid names: Chelonioidae Hay 1930 [empty], Chelydoidae Hay 1930 [empty], Emydoidae Hay 1930 [empty], Pleurosternoidae Hay 1930 [empty], Thalassemydoidae Hay 1930 [empty], Trionychoidae Hay 1930 [empty]
Invalid names: Aplax oberndorferi Meyer 1843 [nomen dubium], Athecae Cope 1871 [empty], Chelonipedidae Sarjeant and Langston 1994 [empty], Chelycarapookus Warren 1969 [nomen dubium], Chelycarapookus arcuatus Warren 1969 [nomen dubium], Chelys blakii Mackie 1863 [nomen dubium], Chengyuchelyidae Yeh 1990 [empty], Emys neumayri Seeley 1881 [nomen dubium], Emys paranensis Burmeister 1885 [nomen nudum], Heckerochelyidae Sukhanov 2006 [empty], Megasternon Gray 1842 [nomen nudum], Mongolochelyidae Sukhanov and Pozdnjakov [empty], Pappochelydia Danilov et al. 2014 [empty], Platemys sehuensis Ameghino 1889 [nomen dubium], Protochelys Lydekker 1889 [nomen dubium], Testudo elata Gervais 1877 [nomen dubium], Testudo stricklandi Phillips 1871 [nomen dubium], Tretosternon punctatum Owen 1842 [nomen dubium], Trionyx mammillaris Cope 1877 [nomen nudum]
Diagnosis
ReferenceDiagnosis
W. G. Joyce et al. 2004‘Pantestudines’ refers to the most inclusive clade containing Chelonia (orig. Testudo) mydas (Linnaeus, 1758), but not a single representative of Aves Linnaeus, 1758 (s. Gauthier and de Queiroz, 2001), Crocodylia Wermuth, 1953 (s. Benton and Clark, 1988), Lepidosauria Haeckel, 1866 (s. Gauthier et al., 1988a), or Mammalia Linnaeus, 1758 (s. Rowe, 1988)