Iguanidae

The Fijian crested iguana Brachylophus vitiensis, photographed by Joseph Burgess.

Belongs within: Lepidosauria.

The Iguanidae are a family of lizards, the iguanas and their relatives, found primarily in the Americas with the genus Brachylophus found in Fiji. Many iguanids are primarily herbivorous.

Characters (from Grismer 2002): Caudal vertebrae with two pairs of transverse processes; valves in descending colon; multicusped, compressed and flared posterior marginal teeth; medially oriented supratemporal bone on parietal.

<==Iguanidae
    |--Iguana DS86
    |    |--I. delicatissima GWN71
    |    `--I. iguana DS86
    |--Ophyoessoides aculeatus DS86
    |--Pristiguana R88
    |--Conolophus DS86
    |    |--C. pallidus USDI77
    |    `--C. subcristate MH96
    |--Enyalioides DS86
    |    |--E. cofanorum DS86
    |    `--E. laticeps DS86
    |--Polychrus DS86
    |    |--P. liogaster DS86
    |    `--P. marmoratus DS86
    |--Brachylophus DS90
    |    |--B. fasciatus DS90
    |    `--B. vitiensis DS90
    |--Leiocephalus H81
    |    |--L. eremitus Cope 1868 H81
    |    `--L. herminieri Dumeril & Bibron 1837 H81
    |--Sceloporus VH02
    |    |--S. cautus C72
    |    |--S. clarki F71
    |    |--S. graciosus F71
    |    |--S. grammicus VH02
    |    |--S. jarrovii VH02
    |    |--S. lineatulus F71
    |    |--S. magister F71
    |    |--S. occidentalis SRS02
    |    |    |--S. o. occidentalis C72
    |    |    `--S. o. biseriatus C72
    |    |--S. olivaceus JP79
    |    |--S. orcutti F71
    |    |--S. undulatus MH96
    |    |    |--S. u. undulatus M68
    |    |    |--S. u. consobrinus C72
    |    |    |--S. u. cowlesi C72
    |    |    |--S. u. elongatus C72
    |    |    |--S. u. erythrocheilus M68
    |    |    |--S. u. hyacinthinus C72
    |    |    `--S. u. tristichus C72
    |    |--S. virgatus C72
    |    `--S. woodi ZL74
    |--Cyclura RFN02
    |    |--C. collei (Gray 1845) H81
    |    |--C. cornuta H81
    |    |    |--C. c. cornuta H81
    |    |    `--C. c. onchiopsis (Cope 1885) [incl. C. nigerrima Cope 1886] H81
    |    |--C. macleayi GWN71
    |    |--C. mattea Miller 1918 H81
    |    |--C. pinguis USDI77
    |    `--C. portoricensis Barbour 1919 H81
    |--Uta F71
    |    |--U. antiquus F71
    |    |--U. nolascensis F71
    |    |--U. palmeri F71
    |    `--U. stansburiana F71
    |         |--U. s. stansburiana F71
    |         |--U. s. elegans F71
    |         `--U. s. nevadensis F71
    |--Streptosaurus mearnsi F71
    |--Urosaurus F71
    |    |--U. graciosus F71
    |    |--U. microscutatus F71
    |    `--U. ornatus F71
    |--Callisaurus draconoides F71
    |--Holbrookia F71
    |    |--H. maculata F71
    |    `--H. texana F71
    |--Uma notata F71, JP79
    |--Sauromalus obesus GWN71
    |--Ctenosaura pectinata GWN71
    |--Dipsosaurus dorsalis GWN71
    |--Basiliscus GWN71
    |    |--B. basiliscus
    |    `--B. vittatus GWN71
    `--Oplurus sebae GWN71

REFERENCES

[C72] Cole, C. J. 1972. Chromosome variation in North American fence lizards (genus Sceloporus; undulatus species group). Systematic Zoology 21 (4): 357-363.

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

[DS90] Dye, T., & D. W. Steadman. 1990. Polynesian ancestors and their animal world. American Scientist 78 (May-June): 207-215.

[F71] Ferguson, G. W. 1971. Variation and evolution of the push-up displays of the side-blotched lizard genus Uta (Iguanidae). Systematic Zoology 20 (1): 79-101.

[GWN71] Gorman, G. C., A. C. Wilson & M. Nakanishi. 1971. A biochemical approach towards the study of reptilian phylogeny: evolution of serum albumin and lactic dehydrogenase. Systematic Zoology 20 (2): 167-185.

Grismer, L. L. 2002. Amphibians and Reptiles of Baja California, including its Pacific islands, and the islands in the Sea of Cortés. University of California Press.

[H81] Honegger, R. E. 1981. List of amphibians and reptiles either known or thought to have become extinct since 1600. Biological Conservation 19: 141-158.

[JP79] Jackson, J. F. & J. A. Pounds. 1979. Comments on assessing the dedifferentiating effect of gene flow. Systematic Zoology 28 (1): 78-85.

[M68] Maslin, T. P. 1968. Taxonomic problems in parthenogenetic vertebrates. Systematic Zoology 17 (3): 219-231.

[MH96] Miller, S. A., & J. P. Harley. 1996. Zoology (3^rd ed.) Wm. C. Brown Publishers: Dubuque (Iowa).

[RFN02] Raxworthy, C. J., M. R. J. Forstner & R. A. Nussbaum. 2002. Chameleon radiation by oceanic dispersal. Nature 415: 784-787.

[R88] Rieppel, O. 1988. The classification of the Squamata. In The Phylogeny and Classification of the Tetrapods, vol. 1. Amphibians, Reptiles, Birds (M. J. Benton, ed.) pp. 261-293. Clarendon Press: Oxford.

[SRS02] Shanbhag, B. A., R. S. Radder & S. K. Saidapur. 2002. Members of opposite sex mutually regulate gonadal recrudescence in the lizard Calotes versicolor (Agamidae). Journal of Biosciences 27 (5): 529-537.

[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.

[VH02] Vidal, N., & S. B. Hedges. 2002. Higher-level relationships of snakes inferred from four nuclear and mitochondrial genes. Comptes Rendus Biologies 325: 977-985.

[ZL74] Zimmerman, J. R. & J. A. Ludwig. 1974. Multiple-discriminant analysis of geographical variation in the aquatic beetle, Rhantus gutticollis (Say) (Dytiscidae). Systematic Zoology 24 (1): 63-71.

Cryptodira

Carapace of Pleurosternon bullocki, from Milner (2004).

Belongs within: Testudinata.
Contains: Centrocryptodira.

The Cryptodira are the larger of the two major clades of living testudinates. Living cryptodires are characterised by the ability to draw the head directly backwards within the shell, but this feature is not yet present in stem members of the clade. Kayentachelys aprix is one such stem cryptodire, known from the Early Jurassic of Arizona.

Synapomorphies (from Gaffney & Meylan 1988): Main adductor tendon with cartilaginous nodule articulating on trochlear surface; processus pterygoideus externus with flat vertical plane oriented at right angles to plane of pterygoid; ventral process of prefrontal contacting vomer.

<==Cryptodira
    |--Kayentachelys [Kayentachelydia, Kayentachelyidae] GM88
    |    `--K. aprix GM88
    `--Selmacryptodira [Baenoidea] GM88
         |--Pleurosternidae GM88
         |    |--Glyptops [Glyptopsidae] GM88
         |    `--Pleurosternon [incl. Mesochelys] GM88
         |         `--P. bullocki N85
         `--Daiocryptodira GM88
              |--Eucryptodira GM88
              |    |--Centrocryptodira GM88
              |    `--Plesiochelyidae GM88
              |         |  i. s.: Parachelys GM88
              |         |         Eurysternum GM88
              |         |--Portlandemys GM88
              |         `--+--Plesiochelys GM88
              |            `--Thalassemys [Thalassemydidae] GM88
              |                 `--T. moseri GM88
              `--Baenidae GM88
                   |--Trinitichelys GM88
                   `--Baeninae GM88
                        |  i. s.: Neurankylus GM88
                        |--Hayemys GM88
                        `--Baenodd GM88
                             |--Plesiobaena GM88
                             `--Baenini GM88
                                  |--Baenina GM88
                                  |    |--Chisternon GM88
                                  |    `--Baena arenosa GM88, C77
                                  `--Eubaenina GM88
                                       |--Eubaena cephalica GM88
                                       `--+--Palatobaena GM88
                                          `--Stygiochelys Gaffney & Hiatt 1971 GM88, D07
                                               `--S. estesi D07

*Type species of generic name indicated

REFERENCES

[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.

[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.

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

Last updated: 11 August 2017.

Testudinata

Amazon river turtle Podocnemis expansa, from here.

Belongs within: Lepidosauromorpha.
Contains: Cryptodira, Chelidae.

The Testudinata are the turtles and tortoises, a clade of reptiles unmistakeable through the enclosure of the body in a bony shell. The shell incorporates the ribcage of the animal, and the shoulder girdle has become enclosed within the ribcage. The relationships of testudinates to other reptiles remain rather uncertain due to their highly modified anatomy. The most commonly supported views are a relationship to the anapsid procolophonians or to the diapsid sauropterygians.

The earliest known turtles come from the Late Triassic (Middle Triassic if the fragment Priscochelys comes from a testudinate) with the basalmost testudinate Odontochelys lacking a developed carapace, though the plastron was already present (Li et al. 2008). As well as the presence of a carapace, testudinates other than Odontochelys are united by the absence of teeth on the maxilla, premaxilla and dentary, which are covered by a horny beak (Gaffney & Meylan 1988). The Late Triassic Proganochelys retained a series of flattened teeth on the roof of the mouth. More derived testudinates, the Casichelydia, are also united by the absence of the lacrimal bone and duct, the basipterygoid articulation being fused, presence of a single vomer, presence of a lateral and ventral wall to the middle ear, and having the paroccipital process tightly sutured to the braincase (Gaffney & Meylan 1988).

The turtle crown group (Testudines) is divided between two major lineages, the Cryptodira and Pleurodira, with the former being more diverse. The Pleurodira are mostly restricted to the Southern Hemisphere; members of this lineage are able to retract their head within the shell by bending the neck horizontally to one side.

<==Testudinata [Chelonia, Proganochelydia]
    |--Odontochelys Li, Wu et al. 2008 [Odontochelyidae] LW08
    |    `--*O. semitestacea Li, Wu et al. 2008 LW08
    `--+--Proganochelys Baur 1887 LW08, D07
       |    `--P. quenstedti GM88
       `--Casichelydia GM88
            |--Proterochersis LW08
            |--Palaeochersis LW08
            `--Testudines LW08
                 |--Cryptodira GM88
                 `--Pleurodira GM88
                      |  i. s.: Hydromeda tectifera H04
                      |         Araripemys Price 1973 D07
                      |           `--A. barretoi D07
                      |--Platychelys GM88
                      `--Eupleurodira GM88
                           |--Chelidae GM88
                           `--+--Pelomedusidae [Pelomedusinae] AS09
                              |    |--+--Taphrosphys GM88
                              |    |  `--Bothremys GM88
                              |    `--+--Pelomedusa subrufa GM88
                              |       `--Pelusios williamsi GM88, AS09
                              `--Podocnemididae [Podocneminae] AS09
                                     |--+--Shweboemys GM88
                                     |  `--Stereogenys GM88
                                     `--+--Erymnochelys GM88
                                        `--+--Peltocephalus GM88
                                           `--Podocnemis GM88
                                                |--P. expansa DS86
                                                |--P. sextuberculata DS86
                                                `--P. unifilis DS86

Testudinata incertae sedis:
  Plastomenus Cope 1872 C77
    |--P. communis Cope 1875 C77
    |--P. corrugatus Cope 1875 C77
    |--P. fractus Cope 1875 C77
    |--P. lachrymalis Cope 1874 C77
    |--P. multifoveatus Cope 1872 [incl. P. catenatus Cope 1875] C77
    |--P. oedemius C77
    |--P. serialis Cope 1876 C77
    |--P. thomasii C77
    `--P. trionychoides C77
  Scaptochelys agassizii AA87
  Anglocetus beatsoni Tarlo 1964 BM78
  Kallokibotion [Kallokibotiidae] PST99
    `--K. bajazidi Nopcsa 1923 PST99
  Cheirogaster gymnesica Bate 1914 RB05, BG-PQ-C02
  Ptychogaster RB05
  Solnhofia GM88
  Cylindraspis FS01
    |--C. indica FS01
    |--C. inepta FS01
    |--C. peltastes FS01
    |--C. triserrata FS01
    `--C. vosmaeri FS01
  Sandowina LW08
  Mongolemys LW08
  Australochelys LW08

*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.

[AA87] Avise, J. C., J. Arnold, R. M. Ball, E. Bermingham, T. Lamb, J. E. Neigel, C. A. Reeb & N. C. Saunders. 1987. Intraspecific phylogeography: the mitochondrial DNA bridge between population genetics and systematics. Annual Review of Ecology and Systematics 18: 489–522.

[BG-PQ-C02] Bailon, S., J. Garcia-Porta & J. Quintana-Cardona. 2002. Première découverte de Viperidae (Reptilia, Serpentes) dans les îles Baléares (Espagne): des vipères du Néogène de Minorque. Description d’une nouvelle espèce du Pliocène. Comptes Rendus Palevol 1: 227–234.

[BM78] Barnes, L. G., & E. Mitchell. 1978. Cetacea. In: Maglio, V. J., & H. B. S. Cooke (eds) Evolution of African Mammals pp. 582–602. Harvard University Press: Cambridge (Massachusetts).

[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.

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

[FS01] Flannery, T., & P. Schouten. 2001. A Gap in Nature: Discovering the World's Extinct Animals. Text Publishing: Melbourne.

[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.

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

[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.

[PST99] Pereda Suberbiola, X., & P. Taquet. 1999. Restes de Rhabdodon (dinosaure ornithopode) de Transylvanie donnés par Nopcsa au Muséum national d'Histoire naturelle de Paris. Geodiversitas 21 (2): 157–166.

[RB05] Rage, J.-C., & S. Bailon. 2005. Amphibians and squamate reptiles from the late early Miocene (MN 4) of Béon 1 (Montréal-du-Gers, southwestern France). Geodiversitas 27 (3): 413–441.

Last updated: 11 August 2017.

Procoelocryptodira

Cast of skeleton of the Cretaceous Toxochelys latiremis, from Oceans of Kansas.

Belongs within: Centrocryptodira.
Contains: Trionychoidea, Emydidae, Testudinoidae, Cheloniidae, Dermochelyoidae.

The clade Procoelocryptodira includes all living cryptodire turtles and tortoises except for the Chelydridae. Procoelocryptodira have smaller tails than other cryptodires, with small or absent haemal spines and procoelous central articulations (Gaffney & Meylan 1988). They can be divided between the marine Chelonioidea, and the freshwater or terrestrial Chelomacryptodira. Characterising features of the chelonioids include elongation of the forearm digits three and four, with flattened carpals and tarsals and a straightened humerus. The Chelomacryptodira are united by a strong temporal emargination of the skull, with a small postorbital bone, limited parietal postorbital contact, and usually no postorbital-squamosal contact (Gaffney & Meylan 1988).

<==Procoelocryptodira
    |--Chelomacryptodira GM88
    |    |--Trionychoidea GM88
    |    `--Testudinoidea GM88
    |         |--Emydidae GM88
    |         `--Testudinoidae GM88
    `--Chelonioidea GM88
         |  i. s.: Corsochelys GM88
         |         Rhinochelys GM88
         |--Toxochelys [Toxochelyinae] GM88
         |    `--T. latiremis GM88
         `--+--Ctenochelys [Lophochelyinae] GM88
            `--Euchelonioidea GM88
                 |--Cheloniidae GM88
                 `--+--Notochelone costata Owen 1882 GM88, F71
                    |--Allopleuron GM88
                    |--Desmatochelys GM88
                    `--Dermochelyoidae GM88

*Type species of generic name indicated

REFERENCES

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

[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.

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.

Batagurinae

Female Chinese striped-neck turtle Ocadia sinensis, photographed by Tim McCormack.

Belongs within: Testudinoidae.

The Batagurinae are a clade of freshwater turtles found in Asia.

Synapomorphies (from Gaffney & Meylan 1988): Fissura ethmoidalis oval and broad; frontals sometimes excluded from orbital rim by prefrontal-postorbital contact; foramen palatinum posterius small, excluded from pterygoid.

<==Batagurinae
    |--+--Orlitia borneoensis GM88, CP03
    |  `--Siebenrockiella crassicollis GM88, CP03
    `--+--+--Geoclemmys hamiltoni GM88, CP03 [=Damonia hamiltoni USDI77]
       |  `--+--Malayemys subtrijuga GM88, CP03
       |     `--Chinemys GM88
       |          |--C. megalocephala CP03
       |          |--C. nigricans CP03
       |          `--C. reevesi CP03
       `--+--Hieremys annandalii GM88, CP03
          `--+--Ocadia sinensis GM88, CP03
             `--+--+--Batagur baska GM88, CP03
                |  `--Callagur borneoensis GM88, CP03
                |--+--Hardella thurgi GM88, CP03
                |  `--Morenia GM88
                |       |--M. ocellata CP03
                |       `--M. petersi CP03
                `--Kachuga GM88
                     |--*K. kachuga CP03
                     |--K. dhongokha CP03
                     |--K. smithii CP03
                     |--K. tecta CP03
                     `--K. tentoria CP03

Hybrids: Ocadia glyphistoma McCord & Iverson 1994 [=O. sinensis × Mauremys annamensis] D03
         Ocadia philippeni McCord & Iverson 1992 [=O. sinensis × Cuora trifasciata] D03

*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.

[D03] Dalton, R. 2003. Mock turtles. Nature 423: 219-220.

[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.

[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.

Testudinoidae

Indian pond turtles Melanochelys trijuga, photographed by Manoj Sindagi.

Belongs within: Procoelocryptodira.
Contains: Batagurinae, Testudinidae, Geoemyda.

The Testudinoidae is a proposed clade uniting the land tortoises of the Testudinidae with the Asian pond turtles of the paraphyletic 'Bataguridae'. The monophyly of this clade is unsettled, as proposed synapomorphies are also found in other testudinates, and are not universal within testudinoidaens. (Gaffney & Meylan 1988).

Synapomorphies (from Gaffney & Meylan 1988): (Including Echmatemys) Axillary and inguinal musk duct foramina enclosed within peripheral plates or between peripherals and plastral buttresses. (Excluding Echmatemys) Sixth marginal scute often reaching third pleural scute; postorbital-squamosal contact absent.

<==Testudinoidae [Bataguridae, Geoemydidae]
    |  i. s.: Annamemys annamensis CP03
    |--Echmatemys GM88
    `--+--Batagurinae GM88
       `--Geoemydinae GM88
            |  i. s.: Rhinoclemmys GM88
            |           |--R. annulata GM88
            |           |--R. areolata CP03
            |           |--R. diademata CP03
            |           |--R. funerea CP03
            |           |--R. melanosterna CP03
            |           |--R. nasuta CP03
            |           |--R. pulcherrima CP03
            |           |--R. punctularia CP03
            |           `--R. rubida GM88
            |--Mauremys GM88
            |    |--M. annamensis D03
            |    |--M. caspica CP03
            |    |--M. japonica GM88
            |    |--M. leprosa CP03
            |    `--M. mutica GM88
            `--+--+--Notochelys platynota GM88, D03
               |  `--Sacalia GM88
               |       |--S. beali CP03
               |       `--S. quadriocellata D03
               `--+--+--Heosemys GM88
                  |  |    |--H. depressa CP03
                  |  |    |--H. grandis CP03
                  |  |    `--H. spinosa CP03
                  |  `--Cyclemys GM88
                  |       |--C. dentata D03
                  |       |--C. oldhamii D03
                  |       |--C. pulchristriata [incl. C. atripons Iverson & McCord 1997] D03
                  |       `--C. tcheponensis CP03
                  `--+--Melanochelys trijuga GM88, CP03
                     `--+--+--Testudinidae GM88
                        |  `--+--Cistoclemmys GM88
                        |     |    |--C. flavomarginata GM88
                        |     |    `--C. galbinifrons GM88
                        |     `--+--Geoemyda GM88
                        |        `--Pyxidea mouhotii GM88, CP03
                        `--Cuora GM88
                             |--C. amboinensis CP03
                             |--C. aurocapitata CP03
                             |--C. flavomarginata (Gray 1863) I92
                             |--C. galbinifrons D03
                             |--C. mccordi Ernst 1988 D03
                             |--C. pani [incl. C. chriskarannarum Ernst & McCord 1987] D03
                             |--C. trifasciata GM88
                             `--C. zhoui [incl. C. pallidicephala McCord & Iverson 1991] D03

Hybrids: Cuora serrata Iverson & McCord 1992 [Pyxidea mouhotii × Cuora galbinifrons] D03
         Mauremys iversoni Pritchard & McCord 1991 [M. mutica × Cuora trifasciata] D03
         Mauremys pritchardi McCord 1997 [M. mutica × Chinemys reevesii] D03
         Sacalia pseudocellata Iverson & McCord 1992 [Cuora trifasciata × S. quadriocellata] D03

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.

[D03] Dalton, R. 2003. Mock turtles. Nature 423: 219-220.

[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.