Sphenodontida (original) (raw)

After Herrera-Flores, Stubbs, Elsler & Benton, 2018 (fig. 3(1)

**<==**o Sphenodontida Estes, 1983 sensu Wu, 1994 [Rhynchocephalia Günther, 1867] (Sphenodontians; alkuliskot) |-- †Gephyrosaurus bridensis Evans, 1980 [paraphyletic Gephyrosauridae] --o Sphenodontia Williston, 1925 |?- †_Diphydontosaurus avonis_ Whiteside, 1986 [paraphyletic Gephyrosauridae] (not treated by Herrera-Flores et al. (2018)] |--o †_Planocephalosaurus_ Fraser, 1982 [paraphyletic Gephyrosauridae] | |-- †_P. robinsonae_ Fraser, 1982 | -- †P. lucasi Heckert, 2004 --+-- †_Rebbanasaurus jaini_ Evans, Prasad & Manhas, 2001 |-- †_Pelecymala robustus_ Fraser, 1986 [=? _Rebbanasaurus_; _Pelecymela_ (lapsus calami)] |--+-- †_Sphenocondor gracilis_ Apestegu�a, G�mez & Rougier, 2012 | -- †Godavarisaurus leteefi Evans, Prasad & Manhas, 2001 --o Eusphenodontia Herrera-Flores, Stubbs, Elsler & Benton, 2018 |-- †_Polysphenodon mulleri_ Jaekel, 1911 [Polysphenodontinae Kuhn, 1969] --+--o †Clevosauridae Bonaparte & Sues, 2006 [“Clevosaurinae” sensu Wu, 1994] | |-- †Brachyrhinodon taylori v. Huene, 1910 | --o †_Clevosaurus_ Swinton, 1939 [_Clevosaurus_ _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._ [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 Neosphenodontia Herrera-Flores, Stubbs, Elsler & Benton, 2018 |--o †Homoeosaurus Meyer, 1850 [Homaeosauridae] | |?- †H. pulchellus | |-- †H. maximiliani | -- †_H. solnhofeni_ Cocude-Michel, 1963 --+--o Sapheosauridae [Sphenodontidae: “Sapheosaurini”] | |--o †Sapheosaurus von Meyer, 1850 [Piocormus] | | |-- †S. thiollierei Meyer, 1850 | | -- †_S. laticeps_ Wagner [_Piocormus laticeps_] | --o †Kallimodon Cocude-Michel, 1963 | |-- †K. pulchellus [= Homoeosaurus pulchellus??] | -- †_K. cerinesis_ --+--o Opisthodontia Apesteguia & Novas, 2003 | |-- †Fraserosphenodon latidens (Fraser, 1993) Herrera-Flores, Stubbs, Elsler & Benton, 2018 [†Clevosaurus sp. Fraser, 1988; Clevosaurus latidens Fraser, 1993] | --+--+-- †_Opisthias rarus_ Gilmore, 1910 [_Theretarius antiquus_ Simpson, 1926 (juv.)]; U.Jur. SWNAm. | | --o †Kawasphenodon Apesteguia, 2005; U.Cret.-L.Paleoc. SAm. | | |-- †K. expectatus Apesteguia, 2005 | | -- †_K. peligrensis_ Apestegu�a, G�mez & Rougier, 2014 | --o †Eilenodontini Rasmussen & Callison, 1981 [Eilenodontinae Rasmussen & Callison, 1981] | |-- †Sphenotitan leyesi Martinez, Apaldetti, Colombi, Praderio, Fernandez, Malnis, Correa, Abelin & Alcober, 2013 | **==**o †_Priosphenodon_ Apesteguia & Novas, 2003 | |-- †_Priosphenodon avelasi_ Apesteguia & Novas, 2003 | |-- †_Priosphenodon minimus_ Apesteguia & Carballido, 2014 | --+-- †Eilenodon robustus Rasmussen & Callison, 1981 | --+-- †_Toxolophosaurus cloudi_ Olson, 1960 [_non_ Throckmorton, Hopson & Parks, 1981] | ?- †Kaikaifilusaurus calvoi Simon & Kellner, 2003 (not treated by Herrera-Flores et al. (2018)] --+--o †Pleurosauridae Lydekker, 1888 | |-- †_Palaeopleurosaurus posidonae_ Carroll, 1985 | --o †Pleurosaurus von Meyer, 1831 | |-- †P. goldfussi von Meyer, 1831 | -- †_P. ginsburgi_ Fabre, 1974 --o Sphenodontidae Cope, 1896 [1870] sensu stricto (Herrera-Flores et al. (2018)) [Sphenodontinae (Cope, 1896 [1870]) Nopcsa, 1928 sensu Wu, 1994] |-- †Pamizinsaurus tlayuaensis Reynoso, 1997 --+--+-- †_Zapatadon ejidoensis_ Reynoso & Clark, 1998 | |-- †_Derasmosaurus pietraroiae_ Barbera & Macuglia, 1988 [_Chometokadmon_ Costa, 1864 (_in partim_)] L.Cret. SEu. | |-- †_Oenosaurus muehlheimensis_ Rauhut, Heyng, L�pez-Arbarello & Hecker, 2012 | -- †Ankylosphenodon pachyostosus Reynoso, 2000 --o Sphenodontini Cope, 1896 [1870] |--o _Sphenodon_ Gray, 1831 (tuataras; tuatarat) | |?- _cf._ †_S. sp._ E.Mioc. NZ. (Jones et al., 2009) | |?- †_S. diversum_ Colenso, 1885 | --o S. punctatus (Gray, 1842) (tuatara) | |-- †S. p. reischeki (South Island tuatara; etel�saaren tuatara) | |-- S. p. guentheri Buller, 1877 (Cook Strait tuatara; Brothers Island tuatara; pikkutuatara; cookinsalmentuatara) | -- _S. p. punctatus_ (Gray, 1842) (North island tuatara; Nortern tuatara; poohjoissaaren tuatara, isotuatara) --+-- †Cynosphenodon huizachalensis Reynoso, 1996 --+-- †_Sphenovipera jimmysjoyi_ Reynoso, 2005; M.Jur. SWNAm. -- †Theretairus antiquus Simpson, 1926; U.Jur. NAm.

Sphenodontia incertae sedis:

Sphenodontians yet unassignated to phylogeny.

†**Sphenodontia** _genera_ et _species_ indet. [Richmond basin, Virginia, USA]
†**Sphenodontidae** gen. et sp. indet. [Evans & Sigogneau-Russell, 1997] [Early Cretaceous, Western North Africa]
†**Sphenodontidae** gen. et sp. indet. [CMSN 5639] [Cau, Baiano & Raia, 2014] [Early Cretaceous, Pietraroia, Italy]
†**Sphenodontidae** gen. et sp. indet. [MPN A01/82] [Cau, Baiano & Raia, 2014] [Early Cretaceous, Pietraroia, Italy]
†_Elachistosuchus hueni_ Janensch, 1949; U.Trias. CEu. [Elachistosuchoidea v. Huene, 1956; Elachistosuchidae Romer, 1966]
†_Lamarquesaurus cabazai_ Apesteguia & Rougier, 2007; U.Cret. SAm.
†_Leptosaurus_ Fitzinger, 1837
    †_Leptosaurus neptunius_ Fitzinger, 1837
    †_Leptosaurus pulchellus_ Zittel 1887
†_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 Wu, 1994, Evans et al., 2001, Apesteguia & Novas, 2003, Bonaparte & Sues, 2006, and Apesteguia & Carballido, 2014 (old Archive combine tree) **<==**o Sphenodontida |-- †Gephyrosaurus `--o Sphenodontia | -- †Diphydontosaurus_ `--+-- †_Planocephalosaurus | --o †Pleurosauridae | | -- †Palaeopleurosaurus | `-- †Pleurosaurus_ `--o Sphenodontidae | -- †_Rebbanasaurus `--+--+-- †Sphenocondor_ | `-- †_Godavarisaurus `--+--o †Clevosauridae | | -- †Polysphenodon_ | `--+-- †_Brachyrhinodon | `-- †Clevosaurus_ `--o Sphenodontidae | -- †_Homoeosaurus `--+-- †Kallimodon_ `--+--o “Sapheosaurini” | | -- †_Ankylosphenodon | `-- †Sapheosaurus_ `--+-- †_Pamizinsaurus `--+--o Sphenodontinae | | --+-- †Zapatadon_ | | | -- †Derasmosaurus | | `-- †_Oenosaurus | `--o Sphenodontini | | -- †Cynosphenodon_ | | -- Sphenodon | `--+-- †_Sphenovipera | `--+-- †Theretairus_ | `-- †_Kawasphenodon `--o Opisthodontia | -- †Opisthias_ `--o †Eilenodontini | --+-- †_Kaikaifilusaurus | `-- †Toxolophosaurus_ `--+-- †_Eilenodon `-- †Priosphenodon_ | After Apesteguia & Carballido, 2014 **<==**o Rhynchocephalia |-- †Gephyrosaurus `--o Sphenodontia | -- †_Diphydontosaurus `--+-- †Planocephalosaurus_ `--+-- †_Rebbanasaurus `--+--+-- †Sphenocondor_ | `-- †_Godavarisaurus `--+--+-- †Brachyrhinodon_ | `-- †_Clevosaurus `--+-- †Palaeopleurosaurus_ `--+--+-- †_Kallimodon | `-- †Sapheosaurus_ `--+-- †_Theretairus | -- †Sphenovipera `--+-- †Homoeosaurus_ | -- †Zapatodon | -- †Cynosphenodon | -- Sphenodon | -- †Ankylosphenodon `--o †Opisthodontia | -- †_Kawasphenodon | -- †Opisthias `--o †Eilenodontinae | -- †Toxolophosaurus_ | -- †Eilenodon `-- †_Priosphenodon Reference(s): Apesteguia, S. & Carballido, J. L., 2014: A New Eilenodontine (Lepidosauria, Sphenodontidae) from the Lower Cretaceous of Central Patagonia.–Journal of Vertebrate Paleontology: Vol. 34, #2, pp. 303-317 [**doi: 10.1080/02724634.2013.803974**] | After Rauhut, Heyng, L�pez-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): Rauhut, O. W. M., Heyng, A. M., L�pez-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**] | | ------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------ | ---------------------------------------------------------- | ------------------- | | ------------------------ | -------------------------------------------------- | ---------------------------------------------------------- | ------------------------------------------------------------- | | ------------------- | ------------------------- | ---------------------------------------------------------- | -------------------------------------------------------------------------------------------------------------- | | --------------------- | ------------------------------------------------------------------------------------------------------------------------------ | | ------------------ | | | ------------------- | | ------------------ | ------------------- | | ------------------- | | -------------- | ----------------------- | ---------------------- | -------------------------------------------------------------------------- | ------------------------------------------------------------------------ | ------------------------- | --------------------------------------------------------------------------------------------------------------------------------------------------- | ----------------------------------------------------------------------------------------------------------- | ----------------------------------------------------------------------------------------------------------------------------------- | ------------------------------------------------------------------ | ----------------------------------------------------------------------------------------------------------------- | ------------------------------------------------------------------- | --------------------------------------------------------------------- | --------------- | ------------------- | -------------- | ------------------------------------------------------------------------ | ------------------- | ---------------------------------------------------------------------- | --------------------- | --------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------- | --------------------------------------------------------------------------------------------------------------------------- | ---------------------------------------------------------------------------------------- | --------------------- | ------------------ | | -------------------- | | ------------------------- | -------------------------------------------------------------- | | ------------------------ | ------------------------------------------------------------ | -------------------------------------------------------------- | | -------------- | --------------------- | ------------------------ | ----------------------------------------------------------- | ------------------------------------------------------------- | --------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------- |

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. & Carballido, J. L., 2014: A New Eilenodontine (Lepidosauria, Sphenodontidae) from the Lower Cretaceous of Central Patagonia.
  –Journal of Vertebrate Paleontology: Vol. 34, #2, pp. 303-317 [**doi: 10.1080/02724634.2013.803974**]
• Apestegu�a, S., G�mez, R. O. & Rougier, G. W., 2014: The youngest South American rhynchocephalian, a survivor of the K/Pg extinction.
  –Proceedings of the Royal Society B: Biological Sciences: Vol. 281, #1792, pp. [**doi: 10.1098/rspb.2014.0811**]
• Apestegu�a, S., G�mez, R. O. & Rougier, G. W., 2012: A basal sphenodontian (Lepidosauria) from the Jurassic of Patagonia: new insights on the phylogeny and biogeography of Gondwanan rhynchocephalians.
  –Zoological Journal of the Linnean Society: Vol. 166, #2, pp. 342-360 [**doi: 10.1111/j.1096-3642.2012.00837.x**]
• 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]
• Cau, A., Baiano, M. A. & Raia, P., 2014: A new sphenodontian (Reptilia, Lepidosauria) from the Lower Cretaceous of Southern Italy and the phylogenetic affinities of the Pietraroia Plattenkalk rhynchocephalians.
  –Cretaceous Research: Vol. 49, pp. 172-180 [**doi: http://dx.doi.org/10.1016/j.cretres.2014.02.001**]
• Daugherty, C. H., Cree, A., Hay, J. M. & Thompson, M. B., 1990: Neglected taxonomy and continuing extinctions of tuatara (Sphenodon).
  –Nature: Vol. 347, #6289, pp. 177-179 [**doi: 10.1038/347177a0**]
• 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
• Hay, J. M., Sarre, S. D., Lambert, D. M., Allendorf, F. W. & Daugherty, C. H., 2010: Genetic diversity and taxonomy: a reassessment of species designation in tuatara (Sphenodon: Reptilia).
  –Conservation Genetics: Vol. 11, #3, pp. 1063-1081 [**doi: 10.1007/s10592-009-9952-7**]
• 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]
• Herrera-Flores, J. A., Stubbs, T. L., Elsler, A. & Benton, M. J., 2018: Taxonomic reassessment of Clevosaurus latidens Fraser, 1993 (Lepidosauria, Rhynchocephalia) and rhynchocephalian phylogeny based on parsimony and Bayesian inference.
  –Journal of Paleontology: in press, pp. 1-9 [**doi: 10.1017/jpa.2017.136**]
• 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_]
• Mart�nez, R. N., Apaldetti, C., Colombi, C. E., Praderio, A., Fernandez, E., Malnis, P. S., Correa, G. A., Abelin, D. & Alcober, O., 2013: A new sphenodontian (Lepidosauria: Rhynchocephalia) from the Late Triassic of Argentina and the early origin of the herbivore opisthodontians.
  –Proceedings of the Royal Society B: Biological Sciences: Vol. 280, #1772, Art.nr. 20132057, pp. (1-7) [**doi: 10.1098/rspb.2013.2057**]
• 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., L�pez-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
• Simon, M. E. & Kellner, A. W. A., 2003: New sphenodontid (Lepidosauria, Rhynchocephalia, Eilenodontinae) from the Candeleros Formation, Cenomanian of Patagonia, Argentina.
  –Boletim do Museu Nacional, Geologia, nova s�rie: #68, pp. 1-12
• 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