Anura

Hochstetter's frog Leiopelma hochstetteri, copyright David M. Green.

Belongs within: Lepospondyli.
Contains: Alytoidea, Pipoidea, Pelobatoidea, Neobatrachia.

The Anura, frogs and toads, are a diverse group of terrestrial to aquatic vertebrates that lack an external tail at maturity and are often adapted for a hopping mode of locomotion. Larvae, if present, are aquatic and legless with a strong, finned tail.

Synapomorphies (from Frost et al. 2006): Prefrontal bone absent; prearticular bone absent; palatine absent; nine or fewer vertebrae; atlas with a single centrum; first spinal nerve exiting from spinal nerve canal via intervertebral foramen; caudal vertebral segments fused into urostyle; hindlimbs significantly longer than forelimbs, including elongation of ankle bones; radius and ulna, tibia and fibula fused; hyobranchial elements fused into hyoid plate; keratinous jaw sheaths and keratodonts present on larval mouthparts; larva with single median spiracle; skin with large subcutaneous lymph spaces; two m. protractor lentis attached to lens.

<==Anura (see below for synonymy)
    |--Bombinanura [Archaeobatrachia, Discoglossanura, Lalagobatrachia, Mesobatrachia, Ranoidei, Sokolanura] FG06
    |    |--Alytoidea FB17
    |    `--Pipanura FB17
    |         |--Pipoidea PW11
    |         `--Acosmanura FG06
    |              |  i. s.: Eurycephalella alcinae FB17
    |              |--Pelobatoidea EJK08
    |              `--Neobatrachia PW11
    `--Leiopelmatoidea FB17
         |--Ascaphus Stejneger 1899 FG06 [Ascaphidae PW11]
         |    |--A. montanus FG06
         |    `--A. truei FG06
         `--Leiopelmatidae [Amphicoela, Liopelmatina] FG06
             |--Vieraella herbstii Reig 1961 G88, M93
             |--Notobatrachus degiustoi Reig 1955 G88, M93
             `--Leiopelma Fitzinger 1861 [=Liopelma Günther 1869] FG06
                  |  i. s.: L. waitomoensis Worthy 1987 WH02
                  |--+--L. auroraensis Worthy 1987 WH02
                  |  |--L. hochstetteri Fitzinger 1861 WH02
                  |  `--L. markhami Worthy 1987 WH02
                  `--+--L. pakeka Bell, Daugherty & Hay 1998 BDH98
                     `--+--L. archeyi Turbott 1942 BDH98, WH02
                        `--L. hamiltoni McCulloch 1919 BDH98

Anura incertae sedis:
  Phractops F13
    |--P. brevipalmatus F13
    `--P. brevipes F13
  Callula baleata M89
  Notodelphys ovifera H04
  Liaobatrachus FB17
    |--L. grabaui ZBH03
    `--L. zhaoi FB17
  Enneabatrachus hechti R00
  Eobatrachus agilis Marsh 1887 (n. d.) R00
  Comobatrachus aenigmatis Hecht & Estes 1960 (n. d.) R00
  Paradiscoglossus americanus Estes & Sanchíz 1982 R00
  Theatonius lancensis Fox 1976 R00
  Gobiatidae R00
    |--Gobiatoides parvus Roček & Nessov 1993 R00
    `--Gobiates Špinar & Tatarinov 1986 R00
         |--G. asiaticus Roček & Nessov 1993 R00
         |--G. bogatchovi Roček & Nessov 1993 R00
         |--G. dzhyrakudukensis Roček & Nessov 1993 R00
         |--G. fritschi Roček & Nessov 1993 R00
         |--G. furcatus Roček & Nessov 1993 R00
         |--G. kermeentsavi Špinar & Tatarinov 1986 R00
         |--G. kizylkumensis Roček & Nessov 1993 R00
         |--G. leptocolaptus (Borsuk-Białynicka 1978) R00
         |--G. sosedkoi (Nessov 1981) [=Eopelobates sosedkoi] R00
         |--G. spinari Roček & Nessov 1993 R00
         `--G. tatarinovi Roček & Nessov 1993 R00
  Altanulia alifanovi Gubin 1993 R00
  Palaeophrynos gessneri Tschudi 1839 RR00
  Batrachus RR00
    |--B. lacustris Pomel 1853 RR00
    |--B. lemanensis Pomel 1853 RR00
    `--B. naiadum Pomel 1853 RR00
  Lutetiobatrachus gracilis RR00
  Liventsovkia jucunda RR00
  Ranavus scarabellii RR00

Anura [Acaudata, Anoura, Anuri, Anuria, Discoglossoidei, Ecaudata, Ecaudati, Heteromorpha, Leiopelmatanura, Miura, Pygomolgaei, Raniformia]

*Type species of generic name indicated

REFERENCES

[BDH98] Bell, B. D., C. H. Daugherty & J. M. Hay. 1998. Leiopelma pakeka, n. sp. (Anura: Leiopelmatidae), a cryptic species of frog from Maud Island, New Zealand, and a reassessment of the conservation status of L. hamiltoni from Stephens Island. Journal of the Royal Society of New Zealand 28 (1): 39–54.

[EJK08] Evans, S. E., M. E. H. Jones & D. W. Krause. 2008. A giant frog with South American affinities from the Late Cretaceous of Madagascar. Proceedings of the National Academy of Sciences of the USA 105 (8): 2951–2956.

[FB17] Feng, Y.-J., D. C. Blackburn, D. Liang, D. M. Hillis, D. B. Wake, D. C. Cannatella & P. Zhang. 2017. Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous–Paleogene boundary. Proceedings of the National Academy of Sciences of the USA 114 (29): E5864–E5870.

[FG06] Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, A. Haas, C. F. B. Haddad, R. O. de Sá, A. Channing, M. Wilkinson, S. C. Donnellan, C. J. Raxworthy, J. A. Campbell, B. L. Blotto, P. Moler, R. C. Drewes, R. A. Nussbaum, J. D. Lynch, D. M. Green & W. C. Wheeler. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1–370.

[F13] Fry, D. B. 1913. On a Varanus and a frog from Burnett River, Queensland, and a revision of the variations in Limnodynastes dorsalis, Gray. Records of the Australian Museum 10 (2): 17–34, pls 1–3.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[M93] Milner, A. R. 1993. Amphibian-grade Tetrapoda. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 665–679. Chapman & Hall: London.

[M89] Modigliani, E. 1889. Materiali per la fauna erpetologica dell’isola Nias. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a, 7: 113–124.

[PW11] Pyron, R. A., & J. J. Wiens. 2011. A large-scale phylogeny of Amphibia including over 2800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution 61: 543–583.

[R00] Roček, Z. 2000. Mesozoic anurans. In: Heatwole, H., & R. L. Carroll (eds) Amphibian Biology vol. 4. Palaeontology. The evolutionary history of amphibians pp. 1295–1331. Surrey Beatty & Sons.

[RR00] Roček, Z., & J.-C. Rage 2000. Tertiary Anura of Europe, Africa, Asia, North America, and Australia. In: Heatwole, H., & R. L. Carroll (eds) Amphibian Biology vol. 4. Palaeontology. The evolutionary history of amphibians pp. 1332–1387. Surrey Beatty & Sons.

[WH02] Worthy, T. H., & R. N. Holdaway. 2002. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).

[ZBH03] Zhou, Z., P. M. Barrett & J. Hilton. 2003. An exceptionally preserved Lower Cretaceous ecosystem. Nature 421: 807–814.

Lepospondyli

Reconstruction of Odonterpeton triangulare by Smokeybjb.

Belongs within: Batrachosauria.
Contains: Nectridea, Albanerpetontidae, Gymnophiona, Anura, Caudata.

The Lepospondyli are a clade of amphibian-grade tetrapods known from the Lower Carboniferous to the Upper Permian though the living amphibians may also be nested here. The basal members of the clade were the lizard- or salamander-like microsaurs, all small animals with relatively short legs. The 'microsaurs' were paraphyletic to more derived subgroups such as the Nectridea or Lysorophia. The Lysorophia were small, serpentine animals with an extremely elongate trunk and reduced, poorly ossified limbs and girdles (Ruta et al. 2003).

The earliest clear relatives of the modern amphibians are known from the Early Triassic. Chinlestegophis jenkinsi, a probably burrowing amphibian from the Upper Triassic of Colorado, is a potential relative of caecilians, sharing with them a wide cultriform process on the skull, laterally placed eyes and double tooth rows on the mandible. The Early Jurassic Eocaecilia micropoda was even more caecilian-like as indicated by features like the loss of the tympanic ear and presence of the pseudodentary (Pardo, Small & Huttenlocker 2017). The salamanders and frogs are united in a clade Batrachia, exclusive of the caecilians, by a number of shared features such as loss of the postfrontal, surangular and splenial bones and loss of dermal scales (Frost et al. 2006). Triadobatrachus massinoti, from the early Triassic of Madagascar, was similar to a modern frog in overall appearance but retained a higher number of vertebrae, including a short tail, and was not adapted for jumping.

Lepospondyli [Hapsidopareiontidae, Holospondyli, Microbrachomorpha, Microsauria, Tuditanomorpha]
    |  i. s.: Trachystegos megalodon Carroll 1966 M93
    |--+--Trihecaton [Trihecatontidae] DS20
    |  |    `--T. howardinus Vaughn 1972 M93
    |  `--+--Nectridea DS20
    |     `--+--Microbrachidae M93
    |        |    |--Paramicrobrachis fritschi Kuhn 1959 M93
    |        |    `--Microbrachis Fritsch 1876 DS20, D07
    |        |         |--M. obtusatum D07
    |        |         `--M. pelikani Fritsch 1876 RCQ03
    |        `--Hyloplesion [Hyloplesiontidae] DS20
    |             `--H. longicostatum Fritsch 1876 RCQ03
    `--+--+--Pantylidae RCQ03
       |  |    |--Pantylus cordatus Cope 1871 DS20, RCQ03
       |  |    `--Stegotretus agyrus Berman, Eberth & Brinkman 1988 AR08, RCQ03
       |  `--Tuditanidae RCQ03
       |       |  i. s.: Boii crassidens (Fritsch 1876) M93
       |       |--Asaphestera intermedia (Dawson 1894) DS20, RCQ03
       |       `--+--Crinodon DS20
       |          `--Tuditanus punctulatus Cope 1874 DS20, RCQ03
       `--+--Saxonerpeton geinitzi (Credner 1890) AR08, RCQ03
          `--+--+--+--Rhynchonkos [Goniorhynchidae, Rhynchonkidae] DS20
             |  |  |    `--R. stovalli (Olson 1970) RCQ03
             |  |  `--+--Dvellecanus carrolli PS17
             |  |     `--Aletrimyti gaskillae PS17
             |  `--+--+--Huskerpeton englehorni PS17
             |     |  `--Gymnarthridae PS17
             |     |       |--Hylerpeton dawsoni T85
             |     |       |--Leiocephalikon problematicum C00, T85
             |     |       |--Elfridia Thayer 1985 T85
             |     |       |    `--*E. bulbidens Thayer 1985 T85
             |     |       |--Euryodus DS20
             |     |       |    |--E. dalyae AR08
             |     |       |    `--E. primus Olson 1939 RCQ03
             |     |       `--Cardiocephalus DS20
             |     |            |--C. dalyae C00
             |     |            |--C. peabodyi AR08
             |     |            `--C. sternbergi Broili 1904 RCQ03
             |     `--+--Nannaroter mckinzei PS17
             |        `--Ostodolepidae PS17
             |             |--Micraroter erythrogeios Daly 1973 DS20, RCQ03
             |             |--Pelodosotis elongatum Carroll & Gaskill 1978 DS20, RCQ03
             |             `--Ostodolepis brevispinatus Williston 1913 M93
             `--+--+--Odonterpeton [Odonterpetontidae] DS20
                |  |    `--O. triangulare Moodie 1909 RCQ03
                |  `--Sparodus Fritsch 1876 DS20, RCQ03
                |       `--S. validens RCQ03
                `--+--+--Hapsidopareion lepton Daly 1973 DS20, RCQ03
                   |  `--Llistrofus Carroll & Gaskill 1978 DS20, RCQ03
                   |       `--L. pricei Carroll & Gaskill 1978 M93
                   `--+--Brachystelechidae PS17
                      |    |--Batropetes fritschia (Geinitz & Deichmüller 1882) DS20, RCQ03
                      |    `--+--Carrolla Langston & Olson 1986 RCQ03
                      |       |    `--C. craddocki RCQ03
                      |       `--Quasicaecilia Carroll 1990 RCQ03
                      |            `--Q. texana Carroll 1990 M93
                      `--+--Lysorophia RCQ03
                         |    |--Molgophidae G88
                         |    |    |--Molgophis G88
                         |    |    `--Megamolgophis G88
                         |    `--Cocytinidae [Lysorophidae] AR08
                         |         |--Cocytinus G88
                         |         |--Lysorophus tricarinatus C07
                         |         |--Pleuroptyx clavatus C00
                         |         `--Brachydectes DS20
                         |              |--B. elongatus Wellstead 1991 RCQ03
                         |              `--B. newberryi Cope 1868 RCQ03
                         `--+--Albanerpetontidae DS20
                            `--+--+--Rileymillerus cosgriffi PSH17
                               |  `--+--Chinlestegophis Pardo, Small & Huttenlocker 2017 PSH17
                               |     |    `--*C. jenkinsi Pardo, Small & Huttenlocker 2017 PSH17
                               |     `--Parabatrachia [Apoda] FG06
                               |          |--Eocaecilia micropodia Jenkins & Walsh 1993 PS17, RCQ03
                               |          `--Gymnophiona DS20
                               `--Batrachia FB17
                                    |--Salientia RCQ03
                                    |    |  i. s.: Czatkobatrachus polonicus RR00
                                    |    |--Anura PSH17
                                    |    |--Prosalirus [Prosaliridae] C01
                                    |    |    `--P. bitis Shubin & Jenkins 1995 R00
                                    |    `--Triadobatrachus Kuhn 1962 AR08, RR00 (see below for synonymy)
                                    |         `--T. massinoti (Piveteau 1936) RCQ03
                                    |--Pangerpeton DS20
                                    |--Caudata DS20
                                    `--+--Belyanerpeton DS20
                                       `--Karauridae M00
                                            |--Karaurus sharovi Ivakhnenko 1978 DS20, RCQ03
                                            `--Kokartus honorarius M00

Triadobatrachus Kuhn 1962 AR08, RR00 [=Protobatrachus Piveteau 1936 (preoc.) RR00; Proanura, Triadobatrachidae]

*Type species of generic name indicated

REFERENCES

[AR08] Anderson, J. S., R. R. Reisz, D. Scott, N. B. Fröbisch & S. S. Sumida. 2008. A stem batrachian from the Early Permian of Texas and the origin of frogs and salamanders. Nature 453: 515–518.

[C00] Carroll, R. L. 2000. Lepospondyls. In: Heatwole, H., & R. L. Carroll (eds) Amphibian Biology vol. 4. Palaeontology. The evolutionary history of amphibians pp. 1198–1269. Surrey Beatty & Sons.

[C01] Carroll, R. 2001. Chinese salamanders tell tales. Nature 410: 534–536.

[C07] Case, E. C. 1907. Revision of the Pelycosauria of North America. Carnegie Institute of Washington Publication 55: 1–176, 35 pls.

[DS20] Daza, J. D., E. L. Stanley, A. Bolet, A. M. Bauer, J. S. Arias, A. Čerňanský, J. J. Bevitt, P. Wagner & S. E. Evans. 2020. Enigmatic amphibians in mid-Cretaceous amber were chameleon-like ballistic feeders. Science 370: 687–691.

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

[FB17] Feng, Y.-J., D. C. Blackburn, D. Liang, D. M. Hillis, D. B. Wake, D. C. Cannatella & P. Zhang. 2017. Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous–Paleogene boundary. Proceedings of the National Academy of Sciences of the USA 114 (29): E5864–E5870.

[FG06] Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, A. Haas, C. F. B. Haddad, R. O. de Sá, A. Channing, M. Wilkinson, S. C. Donnellan, C. J. Raxworthy, J. A. Campbell, B. L. Blotto, P. Moler, R. C. Drewes, R. A. Nussbaum, J. D. Lynch, D. M. Green & W. C. Wheeler. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1–370.

[G88] Gray, J. 1988. Evolution of the freshwater ecosystem: the fossil record. Palaeogeography, Palaeoclimatology, Palaeoecology 62: 1–214.

[M93] Milner, A. R. 1993. Amphibian-grade Tetrapoda. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 665–679. Chapman & Hall: London.

[M00] Milner, A. R. 2000. Mesozoic and Tertiary Caudata and Albanerpetontidae. In: Heatwole, H., & R. L. Carroll (eds) Amphibian Biology vol. 4. Palaeontology. The evolutionary history of amphibians pp. 1412–1444. Surrey Beatty & Sons.

[PSH7] Pardo, J. D., B. J. Small & A. K. Huttenlocker. 2017. Stem caecilian from the Triassic of Colorado sheds light on the origins of Lissamphibia. Proceedings of the National Academy of Sciences of the USA 114 (27): E5389–E5395.

[PS17] Pardo, J. D., M. Szostakiwskyj, P. E. Ahlberg & J. S. Anderson. 2017. Hidden morphological diversity among early tetrapods. Nature 546: 642–645.

[R00] Roček, Z. 2000. Mesozoic anurans. In: Heatwole, H., & R. L. Carroll (eds) Amphibian Biology vol. 4. Palaeontology. The evolutionary history of amphibians pp. 1295–1331. Surrey Beatty & Sons.

[RR00] Roček, Z., & J-C. Rage. 2000. Proanuran stages (Triadobatrachus, Czatkobatrachus). In: Heatwole, H., & R. L. Carroll (eds) Amphibian Biology vol. 4. Palaeontology. The evolutionary history of amphibians pp. 1283–1294. Surrey Beatty & Sons.

[RCQ03] Ruta, M., M. I. Coates & D. L. J. Quicke. 2003. Early tetrapod relationships revisited. Biological Reviews 78: 251–345.

[T85] Thayer, D. W. 1985. New Pennsylvanian lepospondyl amphibians from the Swisshelm Mountains, Arizona. Journal of Paleontology 59 (3): 684–700.

Last updated: 9 February 2022.

Centrocryptodira

Common snapping turtle Chelydra serpentina, photographed by M. J. Baker.

Belongs within: Pancryptodira.
Contains: Pantrionychia, Emydidae, Pantestuguria, Kinosternidae, Dermochelyoidae, Cheloniidae.

The clade Centrocryptodira unites the crown cryptodiran tortoises with the extinct Meiolaniidae, the horned turtles. The Meiolaniidae were often large tortoises with horn- or knob-like protrusions on the rear of the skull and an armoured tail surrounded by bony rings. Remains are known from South America and Australasia, and they survived in the latter region until the Holocene, only becoming extinct within the last 2000 years or so. Among the major lineages of the crown group, the Chelydridae include the snapping turtles (Chelydra serpentina and Macroclemys temminckii) of North and South America, and the big-headed turtle (Platysternon megacephalum) of southeastern Asia. Members of this family have large heads with strong jaws, and limbs that cannot be fully retracted into the shell (Animal Diversity Web). The Kinosternoidea are freshwater turtles found in North and South America, characterised by the loss of the extragular and pectoral scales of the plastron and the absence of scales on the skull roof. Within the Kinosternoidea, the Dermatemydidae are represented by a singly living species, the hickatee Dermatemys mawii of Central America. The Chelonioidea include the living sea turtles, characterised by the modification of the legs into flippers with elongation of the third and fourth digits and flattening of the carpals and tarsals (Animal Diversity Web).

Characters (from Gaffney & Meylan 1988): Formed vertebral articulations; posterior section of canalis caroticus internus with relatively thick floor; plastral buttresses not reaching costal bones.

Centrocryptodira
    |--Meiolaniidae [Meiolanoidea, Miolaniidae] GM88
    |    |--Niolamia argentina Ameghino 1899 B93
    |    |--Ninjemys oweni WW10
    |    |--Warkalania carinaminor WW10
    |    `--Meiolania Owen 1886 D07
    |         |--M. argentina Smith Woodward 1901 F71
    |         |--M. brevicollis WW10
    |         |--M. damelipi White, Worthy et al. 2010 WW10
    |         |--M. mackayi Anderson 1925 F71
    |         |--M. oweni Woodward 1888 B93
    |         `--M. platyceps Owen 1886 WW10
    `--Cryptodira (see below for synonymy) JPG04
         |  i. s.: Emarginachelys GM88
         |--Pantrionychia AS09
         `--+--Cryptoderinea [Pancryptoderinea] JPG04
            |    |--Pantestudinoidea JPG04
            |    |    |--Lindholmemys JPG04
            |    |    |--Mongolemys JPG04
            |    |    `--Testudinoidea [Emychelydia, Panemychelydia] GM88
            |    |         |--Emydidae GM88
            |    |         `--Pantestuguria AS09
            |    `--Panchelydridae JPG04
            |         |--Sinemydidae B93
            |         |    |--Sinemys lens Wiman 1930 B93
            |         |    `--Protochelydra zangerli Erickson 1973 GM88, B93
            |         `--Chelydridae [Chelydrinae, Chelydroidea] GM88
            |              |  i. s.: Devisia methodes B28
            |              |--Chelydra serpentina (Linnaeus 1758) GM88, JPG04 [=Testudo serpentina JPG04]
            |              |    |--C. s. serpentina N10
            |              |    |--C. s. acutirostris N10
            |              |    |--C. s. osceola N10
            |              |    `--C. s. rossignonii N10
            |              `--Platysternini GM88
            |                   |--Macroclemys GM88 [=Macrochelys (l. c.) N10, Macroclemmys (l. c.) N10]
            |                   |    `--M. temminckii (Troost 1835) [=Chelonura temminckii] JPG04
            |                   `--Platysternina [Panplatysternon] GM88
            |                        |--Chelydropsis GM88
            |                        `--Platysternon [Platysternidae] AS09
            |                             `--P. megacephalum Gray 1831 JPG04
            `--+--Pankinosternoidea JPG04
               |    |--Agomphus JPG04
               |    `--Kinosternoidea [Kinosternoidae] JPG04
               |         |--Pankinosternidae [Kinosternia] JPG04
               |         |    |--Hoplochelys GM88
               |         |    `--Kinosternidae GM88
               |         `--Dermatemydidae [Pandermatemys] GM88
               |              |--Baptemys Leidy 1873 GM88, C77
               |              |    |--*B. wyomingensis Leidy 1870 JPG04
               |              |    `--B. tricarinata Hay 1908 B93
               |              `--Dermatemys AS09
               |                   |--D. costilatus Cope 1875 C77
               |                   |--D. mawii Gray 1847 JPG04
               |                   `--D. wyomingensis C77
               `--Chelonioidea [Euchelonioidea, Panchelonioidea] JPG04
                    |  i. s.: Corsochelys GM88
                    |         Rhinochelys GM88
                    |--Pandermochelys JPG04
                    |    |  i. s.: Santanachelys gaffneyi JPG04
                    |    |--Allopleuron Baur 1888 GM88, F91
                    |    |--Dermochelyoidae GM88
                    |    `--Desmatochelyidae B93
                    |         |--Desmatochelys lowi Williston 1898 GM88, B93
                    |         `--Notochelone costata (Owen 1882) B93
                    `--Pancheloniidae JPG04
                         |  i. s.: Euclastes JPG04
                         |--Toxochelys [Toxochelyinae] GM88
                         |    `--T. latiremis Cope 1873 B93
                         `--+--Ctenochelys [Lophochelyinae] GM88
                            |    |--C. acris Zangerl 1953 B93
                            |    `--C. tenuitesta Zangerl 1953 B93
                            `--+--Osteopyginae [Osteopygidae] TH02
                               |    |--Osteopygis emarginatus Cope 1868 B93
                               |    |--Rhetechelys GM88
                               |    `--Erquelinnesia TH02
                               |         |--E. gosseleti (Dollo 1886) B93
                               |         `--E. planimenta TH02
                               `--+--Cheloniidae AS09
                                  `--Eochelyinae TH02
                                       |--Argillochelys Lydekker 1889 TH02
                                       |--Puppigerus Cope 1871 TH02
                                       |    `--P. camperi TH02
                                       `--Eochelone Dollo 1903 GM88, TH02
                                            `--E. brabantica TH02

Cryptodira [Chelomacryptodira, Chersemyda, Panchersemyda, Pantrionychoidea, Polycryptodira, Procoelocryptodira, Trionychoidea] JPG04

*Type species of generic name indicated

REFERENCES

[AS09] Alfaro, M. E., F. Santini, C. Brock, H. Alamillo, A. Dornburg, D. L. Rabosky, G. Carnevale & L. J. Harmon. 2009. Nine exceptional radiations plus high turnover explain species diversity in jawed vertebrates. Proceedings of the National Academy of Sciences of the USA 106 (32): 13410–13414.

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

[B28] Betrem, J. G. 1928. Monographie der Indo-Australischen Scoliiden mit zoogeographischen Betrachtungen. H. Veenman & Zonen: Wageningen.

[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.

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

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[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.

[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.

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

[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.

Last updated: 30 January 2022.

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.