After Wu, 1994, Evans et al., 2001, Apesteguia & Novas, 2003, and Bonaparte & Sues, 2006
<==o Sphenodontida Estes, 1983 sensu Wu, 1994 [Rhynchocephalia Günther, 1867] (Sphenodontians; alkuliskot) |-- †Gephyrosaurus bridensis Evans, 1980 [paraphyletic Gephyrosauridae] `--o Sphenodontia Williston, 1925 |?- Sphenodontia genera et species indet. [Richmond basin, Virginia, USA] |-- †Diphydontosaurus avonis Whiteside, 1986 [paraphyletic Gephyrosauridae] `--+--o †Planocephalosaurus Fraser, 1982 [paraphyletic Gephyrosauridae] | |-- †P. robinsonae Fraser, 1982 | `-- †P. lucasi Heckert, 2004 |--o †Pleurosauridae Lydekker, 1888 | |-- †Palaeopleurosaurus posidonae Carroll, 1985 | `--o †Pleurosaurus Meyer | |-- †P. goldfussi | `-- †P. ginsburgi `--o Sphenodontidae Cope, 1896 [1870] sensu lato (sphenodontids, tuataras; tuatarat) |-- †Rebbanasaurus jaini Evans, Prasad & Manhas, 2001 `--+--o †Clevosauridae Bonaparte & Sues, 2006 [“Clevosaurinae” sensu Wu, 1994] | |-- †Polysphenodon mulleri Jaekel, 1911 [Polysphenodontinae Kuhn, 1969] | `--+-- †Brachyrhinodon taylori v. Huene, 1910 | `--o †Clevosaurus Swinton, 1939 [Glevosaurus sensu Murry, 1987] [incl. Asiacephalosaurus; Rarojugalosaurus; Dianosaurus Young, 1982] L.Trias.-E.Jur. WEu. ENAm. EAs. | |?- †C. convallis Säilä, 2005 | |?- †C. minor Fraser, 1988 | |?- †C. sp. [England] [Fraser, 1988] | |?- †C.? sp. [South Africa] [Gow & Raath, 1977] | |?- †C. sp. [South Africa] [Sues & Reisz, unpubl. data (Sues, Shubin & Olson, 1994)] | |?- †c. brasiliensis Bonaparte & Sues, 2006 | `--+--+-- †C. petilus (Young, 1982) [Dianosaurus petilus Young, 1982] | | `-- †C. hudsoni Swinton, 1939 | `--+?- †C. wangi Wu, 1994 [Asiacephalosaurus wangi (Wu)] | |-- †C. bairdi Sues, Shubin & Olsen, 1994 [was C. sp. (Sues, Shubin & Olsen)] | `-- †C. mcgilli Wu, 1994 [Rarojugalosaurus mcgilli] `--o Sphenodontidae Cope, 1896 [1870] sensu stricto? [Sphenodontinae (Cope, 1896 [1870]) Nopcsa, 1928 sensu Wu, 1994] |--o †Homoeosaurus Meyer, 1850 | |?- †H. pulchellus | |-- †H. maximiliani | `-- †H. solnhofeni Cocude-Michel, 1963 `--+--o †Kallimodon Cocude-Michel, 1963 | |-- †K. pulchellus [= Homoeosaurus pulchellus??] | `-- †K. cerinesis `--+--o “Sapheosaurini” | |-- †Ankylosphenodon pachyostosus Reynoso, 2000 | `--o †Sapheosaurus von Meyer, 1850 [Piocormus] | |-- †S. thiollierei Meyer, 1850 | `-- †S. laticeps Wagner [Piocormus laticeps] `--+-- †Pamizinsaurus tlayuaensis Reynoso, 1997 `--+--o Sphenodontinae (Cope, 1896) sensu Rauhut, Heyng, López-Arbarello & Hecker, 2012 | |-- †Zapatadon ejidoensis Reynoso & Clark, 1998 | |-- †Oenosaurus muehlheimensis Rauhut, Heyng, López-Arbarello & Hecker, 2012 | |?-+-- †Theretairus antiquus (Simpson, 1926) | | `-- †Sphenovipera jimmysjoyi Reynoso, 2005; M.Jur. SWNAm. | `--o Sphenodontini Cope, 1896 [1870] | |-- †Cynosphenodon huizachalensis Reynoso, 1996 | `--o Sphenodon Gray, 1831 (tuataras; tuatarat) | |?- cf. †S. sp. E.Mioc. NZ. (Jones et al., 2009) | |?- †S. diversum Colenso, 1885 | |-- S. guentheri Buller, 1877 (Brothers Island tuatara; pikkutuatara) | `--o S. punctatus (Gray, 1842) (Northern tuatara; isotuatara, tuatara) | |-- †S. p. reischeki | |-- S. p. ssp. (Cook Strait tuatara; cookinsalmentuatara) | `-- S. p. punctatus (northern tuatara) `--o Opisthodontia Apesteguia & Novas, 2003 |-- †Opisthias rarus Gilmore, 1910 [Theretarius antiquus Simpson, 1926 (juv.)]; U.Jur. SWNAm. `--o †Eilenodontini Rasmussen & Callison, 1981 [Eilenodontinae Rasmussen & Callison, 1981] |-- †Toxolophosaurus cloudi Olson, 1960 [non Throckmorton, Hopson & Parks, 1981] `--+-- †Priosphenodon avelasi Apesteguia & Novas, 2003 `-- †Eilenodon robustus Rasmussen & Callison, 1981
Sphenodontia incertae sedis:
Sphenodontian yet unassignated to phylogeny.
†Sphenodontidae gen. et sp. indet. [Evans & Sigogneau-Russell, 1997] [Early Cretaceous, Western North Africa]
†Derasmosaurus Barbera & Macuglia, 1988 [Chometokadmon Costa, 1864 (in partim)] L.Cret. SEu.
†Elachistosuchus hueni Janensch, 1949; U.Trias. CEu. [Elachistosuchoidea v. Huene, 1956; Elachistosuchidae Romer, 1966]
†Godavarisaurus leteefi Evans, Prasad & Manhas, 2001
†Kawasphenodon expectatus Apesteguia, 2005; U.Cret. SAm.
†Lamarquesaurus cabazai Apesteguia & Rougier, 2007; U.Cret. SAm.
†Leptosaurus Fitzinger, 1837
†Leptosaurus neptunius Fitzinger, 1837
†Leptosaurus pulchellus Zittel 1887
†Pelecymela Fraser, 1986
†Paleollanosaurus fraseri Heckert, 2004
†Sigmala sigmala Fraser, 1986 [Sigmala sigmala Fraser & Walkden, 1983 (nomen nudum)]; U.Trias. WEu.
†Tingitana anoualae Evans & Sigogneau-Russell, 1997; L. Cret. WNAf.
–Whitakersaurus bermani Heckert, Lucas, Rinehart & Hunt, 2008: L.Trias. SWNAm.
Alternative phylogenies of Sphenodontia
After Rauhut, Heyng, López-Arbarello & Hecker, 2012<==o Rhynchocephalia |-- †Gephyrosaurus `--o Sphenodontia |-- †Diphydontosaurus `--+-- †Planocephalosaurus `--o Sphenodontidae |--+-- †Homoeosaurus | `--o “Clevosaurs” | |-- †Brachyrhinodon | |-- †Clevosaurus hudsoni | `-- †Clevosaurus wangi `--+--o †Pleurosauridae | |-- †Palaeopleurosaurus | `-- †Pleurosaurus `--+--+-- †Kallimodon | `-- †Sapheosaurus `--+--o Sphenodontinae | |-- Sphenodon | `--+-- †Oenosaurus | `--+-- †Cynosphenodon | `-- †Zapatodon `--o †Opisthodontia |-- †Opisthias `--o †Eilenodontinae |-- †Toxolophosaurus `--+-- †Eilenodon `-- †Priosphenodon Reference(s):
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Reference(s):
- Apesteguia, S., 2005: A late Campanian sphenodontid (Reptilia, Diapsida) from northern Patagonia.
–Comptes Rendus de l'Académie des Sciencies, Paris: Palevol: Preliminary Online Publication, pp. 1-7 - Apesteguia, S. & Novas, F. E., 2003: Large Cretaceous sphenodontian from Patagonia provides insight into lepidosaur evolution in Gondwana.
–Nature: Vol. 425, pp. 609-612 - Apesteguia, S. & Rougier, G. W., 2007: A Late Campanian sphenodontid maxilla from northern Patagonia.
–American Museum Novitates: #3581, pp. 1-11 - Bonaparte, J. F. & Sues, H.-D., 2006: A new species of Clevosaurus (Lepidosauria: Rhynchocephalia) from the upper Triassic of Rio Grande do Sul, Brazil.
–Palaeontology: Vol. 49, #4, pp. 917-923 [doi: 10.1111/j.1475-4983.2006.00568.x] - Daugherty, C., H., Cree, A., Hay, J., M. & Thompson, M., B., 1990: Neglected taxonomy and continuing extinctions of tuatara (Sphenodon).
–Nature: Vol. 347, 13 September, pp. 177-179 - Evans, S. E., Prasad, G. V. R. & Manhas, B. K., 2001: Rhynchocephalians (Diapsida: Lepidosauria) from the Jurassic Kota Formation of India.
–Zoological Journal of the Linnean Society: Vol. 133, #3, pp. 309-334 - Evans, S. E. & Sigogneau-Russell, D., 1997: New sphenodontians (Diapsida: Lepidosauria: Rhynchocephalia) from the Early Cretaceous of North Africa.
–Journal of Vertebrate Paleontology: Vol. 17, #1, pp. 45-51 - Heckert, A. B., Lucas, S. G., Rinehart, L. F. & Hunt, A. P., 2008: A new genus and species of sphenodontian from the Ghost Ranch Coelophysis Quarry (upper Triassic: Apachean), Rock Point Formation, New Mexico, USA.
–Palaeontology: Vol. 51, #4, pp. 827-845 [doi: 10.1111/j.1475-4983.2008.00786.x] - Jones, M. E. H., Tennyson, A. J. D., Worthy, J. P., Evans, S. E. & Worthy, T. H., 2009: A sphenodontine(Rhynchocephalia) from the Miocene of New Zealand and palaeobiogeography of the tuatara (Sphenodon).
–Proceedings of the Royal Society of London: Biological series: Vol. 276, pp. 1385-1390 [doi: 10.1098/rspb.2008.1785] - Murry, P. A., 1987: New reptiles from the Upper Triassic Chinle Formation of Arizona.
–Journal of Paleontology: Vol. 61, #4, pp. 773-786 [permanent link] - Rauhut, O. W. M., Heyng, A. M., López-Arbarello, A. & Hecker, A., 2012: A New Rhynchocephalian from the Late Jurassic of Germany with a Dentition That Is Unique amongst Tetrapods.
–Plos One: Vol. 7, #10, pp. e46839 [doi: 10.1371/journal.pone.0046839] - Reynoso, V.-H., 1996: A Middle Jurassic Sphenodon-like sphenodontian (Diapsida: Lepidosauria) from Huizachal Canyon, Tamaulipas, Mexico.
–Journal of Vertebrate Paleontology: Vol. 16, #2, pp. 210-221 - Reynoso, V.-H., 1997: A "beaded" sphenodontian (Diapsida: Lepidosauria) from the early Cretaceous of Central Mexico.
–Journal of Vertebrate Paleontology: Vol. 17, #1, pp. 52-59 - Reynoso, V.-H., 2000: An unusual aquatic sphenodontian (Reptilia: Diapsida) from the Tlayua Formation (Albian), Central Mexico.
–Journal of Paleontology: Vol. 74, #1, pp. 133-148 - Reynoso, V.-H., 2005: Possibly evidence of a venom apparatus in a Middle Jurassic sphenodontian from the Huizachal Red Beds of Tamaulipas, Mexico.
–Journal of Vertebrate Paleontology: Vol. 25, #3, pp. 646-654 - Reynoso, V.-H. & Clark, J. M., 1998: A dwarf sphenodontian from the Jurassic La Boca Formation of Tamaulipas, Mexico.
–Journal of Vertebrate Paleontology: Vol. 18, #2, pp. 333-339 - Reynoso, V.-H. & Gómez-Bonilla, A., 2000: The Albian sphenodontian Ankylosphenodon pachyostosus of Tlayua, Mexico, is not a eilenodontine but a sapheosaur.
–Journal of Vertebrate Paleontology: Vol. 20, #3, suppl. to #3, 64A - Rieppel, O. 1994: The Lepidosauromorpha: an overview with special emphasis on the Squamata. 23-37.
in Fraser, N. C. & Sues, H-D. 1994: In the Shadow of the Dinosaurs.
–Cambridge University Press, New York. 1994, pp. x-435 - Sues, H.-D., Olsen, P. E. & Kroehler, P. A., 1994: Small tetrapods from the Upper Triassic of the Richmond basin (Newark Supergroup), Virginia. 161-170
in Fraser, N. C. & Sues, H-D. (eds.) 1994: In the Shadow of the Dinosaurs.
–Cambridge University Press, New York. 1994, pp. x-435 - Sues, H.-D., Shubin, N. H. & Olsen, P. E., 1994: A new sphenodontian (Lepidosauria: Rhynchocephalia) from the McCoy Brook Formation (Lower Jurassic) of Nova Scotia, Canada.
–Journal of Vertebrate Paleontology: Vol. 14, #3, pp. 327-340 - Säilä, L. K., 2005: A new species of the sphenodontian reptile Clevosaurus from the lower Jurassic of south Wales.
–Palaeontology: Vol. 48, #4, pp. 817-831 - Wu, X-C. 1994: Late Triassic-Early Jurassic sphenodontians from China and the phylogeny of the Sphenodontia.
in Fraser, N. C. & Sues, H-D. 1994: In the Shadow of the Dinosaurs.
–Cambridge University Press, New York. 1994, pp. x-435