Emydidae

European pond turtle Emys orbicularis, copyright Nicola Maraspini.

Belongs within: Centrocryptodira.

The Emydidae are a family of mostly aquatic tortoises found primarily in the Americas. The only representatives native elsewhere are species of the genus Emys found in the western Palaearctic region. However, the red-eared slider Trachemys scripta has, through the pet trade, become an invasive species in many parts of the world.

Synapomorphies (from Gaffney & Meylan 1988): Batagurine process absent, no pterygoid-basioccipital contact; angular usually reaching Meckel's cartilage; double articulation of fifth and sixth cervical; chromosome number 2n = 50.

<==Emydidae [Panemydidae]
    |  i. s.: Glyptemys CP03
    |           |--G. insculpta (Le Conte 1830) JPG04
    |           `--G. muhlenbergii (Schoepf 1801) JPG04
    |         Pseudograptemys Hutchison 1996 H96
    |           `--*P. inornata (Loomis 1904) [=Chrysemys inornata, Graptemys inornata; incl. G. cordifera Clark 1937] H96
    |--Emydinae GM88
    |    |--Clemmys Ritgen 1828 GM88, F91
    |    |    |--C. caspica CS77
    |    |    |--C. guttata (Schneider 1792) JPG04
    |    |    |--C. insculpta W-CL06
    |    |    `--C. muhlenbergi MB93
    |    `--+--+--Emydoidea GM88
    |       |  `--Terrapene GM88
    |       |       |--T. carolina (Linnaeus 1758) JPG04
    |       |       |--T. coahuila Schmidt & Owens 1944 JPG04
    |       |       |--T. nelsoni Stejneger 1925 JPG04
    |       |       `--T. ornata (Agassiz 1857) JPG04
    |       `--Emys Dumeriel 1806 GM88, F91
    |            |--E. blandingii (Holbrook 1838) JPG04
    |            |--E. cibollensis Cope 1877 C77
    |            |--E. euthnetus C77
    |            |--E. gravis C77
    |            |--E. lativertebralis Cope 1876 C77
    |            |--E. marmorata Baird & Girard 1852 JPG04
    |            |--E. nebrascensis Leidy 1851 H96
    |            |--E. orbicularis (Linnaeus 1758) K08, L58 [=Testudo orbicularis L58]
    |            |--E. testudineus C77
    |            `--E. vyomingensis C77
    `--Deirochelyinae [Deirochelyoidae] GM88
         |--Malaclemys terrapin (Schoepf 1793) GM88, JPG04
         |--Graptemys GM88
         |    |--G. barbouri Carr & Marchand 1942 JPG04
         |    |--G. caglei Haynes & McKown 1974 JPG04
         |    |--G. ernsti Lovich & McCoy 1992 JPG04
         |    |--G. flavimaculata Cagle 1954 JPG04
         |    |--G. geographica (LeSueur 1817) JPG04
         |    |--G. gibbonsi Lovich & McCoy 1992 JPG04
         |    |--G. kohni CP03
         |    |--G. nigrinoda Cagle 1954 JPG04
         |    |--G. oculifera (Baur 1890) JPG04
         |    |--G. ouachitensis Cagle 1953 JPG04
         |    |--G. pseudogeographica (Gray 1831) JPG04
         |    |--G. pulchra Baur 1893 JPG04
         |    `--G. versa Stejneger 1925 JPG04
         `--+--Chrysemys Gray 1844 GM88, H96
            |    |--C. antiqua (Clark 1937) [=Trachemys antiqua] H96
            |    |--C. bicarinata (Bell 1849) B93
            |    |--C. picta (Schneider 1783) [=Testudo picta] JPG04
            |    |--C. scripta B81
            |    |--C. testudiniformis (Owen 1844) B93
            |    `--C. timida Hay 1908 H96
            `--+--Deirochelys reticularia (Latreille 1801) GM88, JPG04
               `--+--Pseudemys GM88
                  |    |--P. alabamensis Baur 1893 JPG04
                  |    |--P. concinna (Le Conte 1830) JPG04 [=Chrysemys concinna R93]
                  |    |--P. floridana (Le Conte 1830) JPG04
                  |    |--P. nelsoni Carr 1938 JPG04
                  |    |--P. rubriventris (Le Conte 1830) JPG04
                  |    |--P. scripta B81
                  |    |--P. texana Baur 1893 JPG04
                  |    `--P. troosti D56
                  `--Trachemys GM88
                       |--T. adiutrix (Vanzolini 1995) JPG04
                       |--T. callirostris (Gray 1855) JPG04
                       |--T. decorata (Barbour & Carr 1940) JPG04
                       |--T. decussata (Gray 1831) JPG04
                       |--T. dorbigni (Dumeril & Bibron 1835) JPG04
                       |--T. emolli (Legler 1990) JPG03
                       |--T. gaigeae (Hartweg 1939) JPG04
                       |--T. grayi CP03
                       |--T. nebulosa (Van Denburgh 1895) JPG04
                       |--T. ornata (Gray 1831) JPG04
                       |--T. scripta (Schoepf 1792) JPG04
                       |    |--T. s. scripta W-CL06
                       |    |--T. s. elegans W-CL06
                       |    `--T. s. trootsi W-CL06
                       |--T. stejnegeri (Schmidt 1928) JPG04
                       |--T. taylori (Legler 1960) JPG04
                       |--T. terrapen (Bonnaterre 1789) JPG04
                       |--T. venusta (Gray 1855) JPG04
                       `--T. yaquia (Legler & Webb 1970) JPG04

*Type species of generic name indicated

REFERENCES

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

[B81] Brooks, D. R. 1981. Raw similarity measures of shared parasites: an empirical tool for determining host phylogenetic relationships? Systematic Zoology 30 (2): 203–207.

[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.

[CS77] Cramp, S., & K. E. L. Simmons (eds) 1977. Handbook of the Birds of Europe, the Middle East and North Africa: The Birds of the Western Palaearctic vol. 1. Ostrich to Ducks. Oxford University Press: Oxford.

[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.

[F91] Freess, W. B. 1991. Beiträge zur Kenntnis von Fauna und Flora des marinen Mitteloligozäns bei Leipzig. Altenburger Naturwissenschaftliche Forschungen 6: 3–74.

[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.

[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.

[K08] Korsós, Z. 2008. History of the herpetological collection of the Hungarian Natural History Museum. Annales Historico-Naturales Musei Nationalis Hungarici 100: 37–93.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[MB93] Malecki, R. A., B. Blossey, S. D. Hight, D. Schroeder, L. T. Kok & J. R. Coulson. 1993. Biological control of purple loosestrife. BioScience 43 (10): 680–686.

[R93] Rieppel, O. 1993. Patterns of diversity in the reptilian skull. In: Hanken, J., & B. K. Hall (eds) 1993. The Skull vol. 2. Patterns of Structural and Systematic Diversity pp. 344–390. The University of Chicago Press.

[W-CL06] Watkins-Colwell, G. J., T. A. Leenders, B. T. Roach, D. J. Drew, G. Dancho & J. Yuckienuz. 2006. New distribution records for amphibians and reptiles in Connecticut, with notes on the status of an introduced species. Bulletin of the Peabody Museum of Natural History 47 (1–2): 47–62.

Last updated: 8 December 2021.

Testudinidae

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

Belongs within: Testudinoidae.
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
    |  i. s.: Hadrianus Cope 1872 CP03, C76
    |           `--H. corsonii Leidy 1871 C76
    |--Manouria GM88
    |    |--M. emys CP03
    |    `--M. impressa CP03
    `--+--+--Hesperotestudo GM88
       |  `--Testudininae GM88
       `--Xerobatinae GM88
            |--Stylemys GM88
            `--+--Xerobates GM88
               `--Gopherus GM88
                    |--G. agassizii CP03
                    |--G. berlandieri CP03
                    `--G. polyphemus CP03

REFERENCES

[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.

[C76] Cope, E. D. 1876. 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.

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

Kinosternidae

Kinosternon cruentatum, photographed by Zippo Zimmermann.

Belongs within: Trionychoidea.

The Kinosternidae of North and South America include the musk turtles, so called because of their production of a foul-smelling substance from glands under the tail when disturbed.

Synapomorphies (from Gaffney & Meylan 1988): Ten pairs of peripherals; abdominal scales absent; no ribs connecting ninth and tenth vertebrae to eight costal; frontal not reaching orbit; maxilla contacting quadratojugal; ilio-ischial notch in acetabulum; incised anterior musk duct.

<==Kinosternidae
    |  i. s.: Sternotherus WA98
    |           |--+--+--S. carinatus WA98
    |           |  |  `--S. depressus WA98
    |           |  `--S. minor WA98
    |           |       |--S. m. minor WA98
    |           |       `--S. m. peltifer WA98
    |           `--S. odoratus WA98
    |         Xenochelys formosa WA98
    |--Staurotypinae GM88
    |    |--Staurotypus GM88
    |    `--Claudius GM88
    `--Kinosternon GM88
         |--K. baurii WA98
         |--K. flavescens WA98
         |--K. scorpioides DS86
         `--K. subrubrum WA98
              |--K. s. subrubrum WA98
              |--K. s. hippocrepis WA98
              `--K. s. steindachneri WA98

REFERENCES

[DS86] Dixon, J. R. & P. Soini. 1986. The Reptiles of the Upper Amazon Basin, Iquitos Region, Peru. Milwaukee Public Museum: Milwaukee.

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

[WA98] Walker, D., & J. C. Avise. 1998. Principles of phylogeography as illustrated by freshwater and terrestrial turtles in the southeastern United States. Annual Review of Ecology and Systematics 29: 23-58.

Geoemyda

Geoemyda spengleri, photographed by Jan Ševčík.

Belongs within: Testudinoidae.

Geoemyda (leaf turtles) is a genus of freshwater turtles found in eastern Asia.

Characters (from Rooij 1915): Head covered with smooth skin, undivided. Alveolar surfaces narrow, without median ridge; choanae between the eyes. Neural plates hexagonal, short-sided behind or alternately tetragonal and octagonal. Plastron united to the carapace by suture. Digits with or without a short web; five claws on the fore limb, four on the hind limb. Tail very short.

<==Geoemyda
    |--G. punctularia Daudin 1802 M54
    |    |--G. p. punctularia [incl. G. bellii Gray 1863, G. dorsualis Spix 1824] M54
    |    |--G. p. diademata Mertens 1954 M54
    |    |--G. p. lunata Gray 1873 [incl. G. ventricosa Gray 1873] M54
    |    |--G. p. melanosterna Gray 1861 M54
    |    `--G. p. nasuta Boulenger 1902 M54
    |--G. pulcherrima [incl. G. callocephalus Gray 1863] M54
    |--G. spengleri CP03
    |    |--G. s. spengleri I92
    |    `--G. s. japonica Fan 1931 I92
    |--G. sylvatica CP03
    |--G. tricarinata [=Vicoria tricarinata] USDI77
    `--G. yuwonoi CP03

*Type species of generic name indicated

REFERENCES

[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.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[M54] Mertens, R. 1954. Zur Kenntnis der Schildkrötenfauna Venezuelas. Senckenbergiana Biologica 35: 3-7.

Rooij, N. de. 1915. The Reptiles of the Indo-Australian archipelago. E. J. Brill: Leiden.

[USDI77] USDI (United States Department of the Interior). 1977. Endangered and threatened wildlife and plants - republication of list of species. Federal Register 42: 36420-36431.

Testudininae

Radiated tortoises Asterochelys radiata photographed in Madagascar by czwg.

Belongs within: Testudinidae.
Contains: Geochelone.

The Testudininae contains the land tortoises most closely related to the type genus, Testudo, with many species having been referred in the past to either that genus or Geochelone. Among the most famous members of the group are the giant tortoises of the Galapagos (Chelonoidis) and Seychelles islands (Aldabrachelys). The genus Testudo includes species found in the Mediterranean region and western Asia, including Hermann's tortoise T. hermanni which is commonly kept in captivity around Europe. Kinixys, the hinged tortoises, is a genus of African tortoises that possess a hinge in the rear part of the shell allowing the rear legs to be covered after retraction.

See also: Tortoise resurrection.

Synapomorphies (from Gaffney & Meylan 1988): Process of surangular interdigitates with dentary, lower jaw relatively gracile; mental glands absent; medial centrale large, separating distal end of radius from contact with medial proximal carpals.

<==Testudininae
    |--+--+--Impregnochelys GM88
    |  |  `--Kinixys GM88
    |  |       |--K. belliana CP03
    |  |       |--K. erosa CP03
    |  |       `--K. homeana CP03
    |  `--+--Indotestudo GM88
    |     |    |--I. elongata CP03
    |     |    |--I. forsteni CP03
    |     |    `--I. travancorica CP03
    |     `--Testudinini GM88
    |          |--+--+--Chersina angulata Gray 1866 GM88, G66
    |          |  |  `--+--Acinixys GM88
    |          |  |     `--Pyxis arachnoides GM88, CP03
    |          |  `--+--Homopus areolatus GM88, CP03
    |          |     `--Psammobates GM88
    |          |          |--P. geometricus CP03
    |          |          |--P. oculifer CP03
    |          |          `--P. tentorius CP03
    |          `--Testudinina GM88
    |               |--Malacochersus tornieri GM88, CP03
    |               `--Testudo GM88
    |                    |--T. antakyensis Perälä 1996 TK03
    |                    |--T. denticulata Gray 1866 G66
    |                    |--T. graeca Linnaeus 1758 TK03
    |                    |    |--T. g. graeca TK03
    |                    |    |--T. g. anamurensis Weissinger 1987 TK03
    |                    |    |--T. g. armeniaca Chkhikvadze & Bakradze 1991 TK03
    |                    |    |--T. g. ibera Pallas 1814 TK03
    |                    |    `--T. g. terrestris Forsskål 1775 TK03
    |                    |--T. hermanni CP03
    |                    |    |--T. h. hermanni M55
    |                    |    `--T. h. robertmertensi M55
    |                    |--T. horsfieldii TK03
    |                    |--T. kleinmanni TK03
    |                    |--T. klettiana Cope 1875 C77
    |                    |--T. loveridgii LW08
    |                    |--T. marginata CP03
    |                    |--T. perses Perälä 2002 TK03
    |                    |--T. tabulata Gray 1866 G66
    |                    |--T. undata Cope 1875 C77
    |                    `--T. weissingeri CP03
    `--Megalochelyini [Megalochelyina] GM88
         |--+--Asterochely radiata GM88, CP03
         |  `--+--Megalochelys GM88
         |     `--Aldabrachelys Loveridge & Williams 1957 GM88, N10 [incl. Dipsochelys Bour 1982] N10
         |          |--‘Dipsochelys’ abrupta (Grandidier 1868) [=Geochelone abrupta] N10
         |          |--‘Dipsochelys’ arnoldi Bour 1982 N10
         |          |--‘Geochelone’ daudinii Dumeril & Bibron 1835 [=Dipsochelys daudinii] N10
         |          |--‘Geochelone’ dussumieri Gray 1831 [=Dipsochelys dussumieri] N10
         |          |--A. elephantina CP03
         |          |--A. grandidieri (Vaillant 1885) M94, N10 (see below for synonymy)
         |          |--‘Geochelone’ hololissa Gunther 1877 [=Dipsochelys hololissa] N10
         |          `--A. robusta M94
         `--+--Chelonoidis GM88
            |    |--C. carbonaria CP03
            |    |--C. chilensis CP03
            |    |--C. denticulata CP03
            |    `--C. nigra CP03
            `--Geochelone GM88

Aldabrachelys grandidieri (Vaillant 1885) M94, N10 [=Geochelone grandidieri N10, Dipsochelys grandidieri N10, Testudo grandidieri M94]

*Type species of generic name indicated

REFERENCES

[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.

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

[G66] Gray, J. E. 1866. Notes on some young specimens of tortoises (Testudo). Proceedings of the Zoological Society of London 1866: 305-306.

[LW08] Li, C., X.-C. Wu, O. Rieppel, L.-T. Wang & L.-J. Zhao. 2008. An ancestral turtle from the Late Triassic of southwestern China. Nature 456: 497-501.

[M94] MacPhee, R. D. E. 1994. Morphology, adaptations, and relationships of Plesiorycteropus, and a diagnosis of a new order of eutherian mammals. Bulletin of the American Museum of Natural History 220: 1-214.

[M55] Mertens, R. 1955. Die Amphibien und Reptilien der Insel Elba. Senckenbergiana Biologica 36: 287-296.

[N10] Naish, D. 2010. Tetrapod Zoology: Book One. CFZ Press: Bideford (UK).

[TK03] Türkozan, O., Y. Kumlutaş, H. Arikan, Ç. Ilgaz & A. Avcı. 2003. Morphological and serological comparison of Mediterranean spur-thighed tortoises, Testudo graeca, from the Aegean region and southeastern Turkey. Zoology in the Middle East 29: 41-50.

Dermochelyoidae

Leatherback turtle Dermochelys coriacea. Photo from here.

Belongs within: Centrocryptodira.

The clade Dermochelyoidae unites the living leatherback turtle Dermochelys coriacea and its close fossil relatives with the extinct family Protostegidae. Dermochelys coriacea is the largest living turtle and is readily distinguished from other living Testudines by the lack of a solid external shell.

Synapomorphies (from Gaffney & Meylan 1988): Jugal very large, ventral process extending posteriorly to form most of cheek margin; basisphenoid and occipital elements usually poorly ossified, with portions remaining cartilaginous; forelimbs very long in relation to hindlimbs and body; trochanter major of femur very long in relation to trochanter minor, projecting proximal to femur head; intertrochanteric fossa largely obliterated by fusion of trochanter bases.

<==Dermochelyoidae [Dermochelyoidea]
    |--Dermochelyidae GM88
    |    |--Eosphargis GM88
    |    `--+--Psephophorus GM88
    |       |    |--P. calvertensis Palmer 1909 K95
    |       |    |--P. eocaenus Andrews 1901 K95
    |       |    |--P. oregonensis Palmer 1940 K95
    |       |    |--P. polygonus v. Meyer 1847 K95
    |       |    |--P. pseudostracion (Gervais 1849) K95
    |       |    |--P. rupeliensis (v. Beneden 1883) K95
    |       |    |--P. scaldii (van Beneden 1871) K95
    |       |    `--P. terrypratchetti Köhler 1995 K95
    |       `--Dermochelys coriacea Linnaeus 1766 GM88, K95
    `--Protostegidae GM88
         |--Chelospharginae GM88
         |    |--Chelosphargis GM88
         |    `--Calcarichelys Zengerl 1953 GM88, D07
         |         `--C. gemma D07
         `--Protosteginae GM88
              |--Protostega gigas GM88, N85
              `--Archelon Wieland 1896 GM88, D07
                   `--A. ischyros D07

*Type species of generic name indicated

REFERENCES

[D07] Dixon, D. 2007. The Complete Illustrated Encyclopedia of Dinosaurs & Prehistoric Creatures. Hermes House: London.

[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.

[K95] Köhler, R. 1995. A new species of the fossil turtle Psephophorus (order Testudines) from the Eocene of the South Island, New Zealand. Journal of the Royal Society of New Zealand 25: 371–384.

[N85] Norman, D. 1985 (reprinted 2000). The Illustrated Encyclopedia of Dinosaurs. Salamander Books: London.

Last updated: 11 August 2017.

Cheloniidae

Loggerhead turtle Caretta caretta, photographed by Strobilomyces.

Belongs within: Procoelocryptodira.

The Cheloniidae (here used to include the Osteopygidae) is the total group containing living sea turtles with the exception of the leatherback Dermochelys coriacea.

Synapomorphies (from Gaffney & Meylan 1988): Secondary palate formed by broad maxilla with contributions from palatine, choanal passages separated by ventral plate of vomer forming prominent part of palate; foramen caroticum laterale and canalis caroticus lateralis much larger in diameter than foramen anterius canalis carotici interni and medial branch of canalis caroticus internus.

<==Cheloniidae
    |  i. s.: Tasbacka Nessov 1987 TH02
    |           |--*T. aldabergeni Nessov 1987 TH02
    |           `--T. ouleabdounensis Tong & Hirayama 2002 TH02
    |         Catapleura ruhoffi Weems 1988 TH02
    |--Osteopyginae [Osteopygidae] TH02
    |    |--Osteopygis emarginatus TH02
    |    |--Rhetechelys GM88
    |    `--Erquelinnesia TH02
    |         |--E. gosseleti TH02
    |         `--E. planimenta TH02
    `--+--Eochelyinae TH02
       |    |--Argillochelys Lydekker 1889 TH02
       |    |--Puppigerus Cope 1871 TH02
       |    |    `--P. camperi TH02
       |    `--Eochelone Dollo 1903 GM88, TH02
       |         `--E. brabantica TH02
       `--+--Carettini [Carettidae] GM88
          |    |--Procolpochelys GM88
          |    |--Lepidochelys olivacea GM88, TH02
          |    |--Caretta F13
          |    |    |--*C. caretta (Linnaeus 1758) I92
          |    |    |--C. olivacea F13
          |    |    `--C. remivaga Hay 1908 F13
          |    `--Colpochelys kempii [=Lepidochelys kempii] F13
          `--Chelonini GM88
               |--Eretmochelys imbricata (Linné 1766) GM88, I92
               `--Chelonia [incl. Natator McCulloch 1908] F13
                    |--‘Chelone’ cuneiceps F13
                    |--C. depressa Garman 1881 [incl. *Natator tessellatus McCulloch 1908] F13
                    |--C. imbricata [=Chelone imbricata, Testudo imbricata] R13
                    |--C. japonica F13
                    `--C. mydas (Linnaeus 1758) I92 [=Chelone mydas F13]
                         |--C. m. mydas BF88
                         |--C. m. agassizi M94
                         `--C. m. caranigra BF88

*Type species of generic name indicated

REFERENCES

[BF88] Bishop, M. J., & A. E. Friday. 1988. Estimating the interrelationships of tetrapod groups on the basis of molecular sequence data. In The Phylogeny and Classification of the Tetrapods, vol. 1. Amphibians, Reptiles, Birds (M. J. Benton, ed.) pp. 33-58. Clarendon Press: Oxford.

[F13] Fry, D. B. 1913. On the status of Chelonia depressa, Garman. Records of the Australian Museum 10 (7): 159-185, plates XIX-XXII.

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

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[M94] Moritz, C. 1994. Defining ‘evolutionarily significant units’ for conservation. Trends in Ecology and Evolution 9 (10): 373-375.

[R13] Roux, J. 1913. Les reptiles de la Nouvelle-Calédonie et des îles Loyalty. In Nova Caledonia: Forschungen in Neu-Caledonian und auf den Loyalty-Inseln. A. Zoologie vol. 1, Heft II (F. Sarasin & J. Roux, eds) pp. 79-160. C. W. Kreidels Verlag: Wiesbaden.

[TH02] Tong, H., & R. Hirayama. 2002. A new species of Tasbacka (Testudines: Cryptodira: Cheloniidae) from the Paleocene of the Ouled Abdoun phosphate basin, Morocco. Neues Jahrbuch für Geologie und Paläontologie Monatshefte 2002 (5): 277-294.