Rhynchocephalia

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Rhynchocephalians
Temporal range: Late Triassic - Recent, 228–0Ma
Tuatara, Sphenodon punctatus
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Superorder: Lepidosauria
Order: Rhynchocephalia
Williston, 1925
Families

Rhynchocephalia is an order of lizard-like reptiles that includes only one living genus and species, the tuatara (Sphenodon punctatus). Despite its current lack of diversity, the Rhynchocephalia at one time included a wide array of genera in several families, and represents a lineage stretching back to the Mesozoic Era. Many of the niches occupied by lizards today were then held by sphenodontians. There was even a successful group of aquatic sphenodontians known as pleurosaurs. They are thought to only inhabit parts of New Zealand.[1]

Systematics

Fossil specimen of Kallimodon pulchellus

Sphenodonts, and their sister group Squamata (which includes lizards, snakes and amphisbaenians), belong to the superorder Lepidosauria, the only surviving taxon within Lepidosauromorpha. Squamates and sphenodonts both show caudal autotomy (loss of the tail-tip when threatened), and have transverse cloacal slits.[2] The origin of the sphenodonts probably lies close to the split between the Lepidosauromorpha and the Archosauromorpha. Though they resemble lizards, the similarity is superficial, because the group has several characteristics unique among reptiles. The typical lizard shape is very common for the early amniotes; the oldest known fossil of a reptile, the Hylonomus, resembles a modern lizard.[3] R.L. Ditmars, Litt.D, says; "The Tuatara resembles in form stout-bodies modern lizards, which we might call iguanas; this resemblance is further intensified by a row of spines upon the back. It is dark olive, the sides sprinkled with pale dots. The eye has a cat-like pupil. Large specimens are two and a half feet long. While superficial resemblance might tend to group this reptile with lizards, its skeleton and anatomy show it to belong to a different part of a technical classification."[4]

Tuatara were originally classified as lizards in 1831 when the British Museum received a skull.[5] The genus remained misclassified until 1867, when Albert Günther of the British Museum noted features similar to birds, turtles, and crocodiles. He proposed the order Rhynchocephalia (meaning "beak head") for the tuatara and its fossil relatives.[6] Many disparately related species were subsequently added to the Rhynchocephalia, resulting in what taxonomists call a "wastebasket taxon".[7] Williston proposed the Sphenodontia to include only tuatara and their closest fossil relatives in 1925.[7] Sphenodon is derived from the Greek for "wedge" (σφηνος/sphenos) and "tooth" (δόντι/odon(t)).[8] However, today Rhynchocephalia is used to include Gephyrosaurus and Sphenodontia, while Sphenodontia excludes the former.[9][10]

Taxonomy

Classification follows Wu (1994),[11] Evans et al. (2001),[9] Apesteguia & Novas (2003)[12] and Evans & Borsuk−Białynicka (2009).[10]

Phylogeny

The following is a cladogram of Rhynchocephalia after Wu (1994),[11] Evans et al. (2001),[9] and Apesteguia & Novas (2003).[12][13]

Rhynchocephalia

 Gephyrosaurus 


Sphenodontia

 Diphydontosaurus 




 Planocephalosaurus 


Pleurosauridae

 Palaeopleurosaurus



 Pleurosaurus



Sphenodontidae

 Rebbanasaurus



"Clevosaurinae"

 Polysphenodon



 Brachyrhinodon



 Clevosaurus



Sphenodontinae

 Homoeosaurus




 Kallimodon



"Sapheosaurini"

 Sapheosaurus



 Ankylosphenodon





 Pamazinsaurus





 Zapatadon




 Theretairus



 Sphenovipera



Sphenodontini

 Cynosphenodon



 Sphenodon (Tuatara)




Opisthodontia

 Opisthias


Eilenodontini

 Toxolophosaurus




 Priosphenodon



 Eilenodon















References

  1. Ditmars, Raymond L., "Reptiles of the World" The MacMillan Co., New York, 1936, p. xii
  2. Cree, Alison. 2002. Tuatara. In: Halliday, Tim and Adler, Kraig (eds.), The new encyclopedia of reptiles and amphibians, Oxford University Press, Oxford, pp. 210–211. ISBN 0-19-852507-9
  3. "Hylonomus lyelli". Symbols. Province of Nova Scotia. May 2003. Retrieved 24 May 2007. 
  4. Ditmars, 1936, p. xiv
  5. Lutz 2005, p. 42.
  6. Lutz 2005, p. 43.
  7. 7.0 7.1 Fraser, Nicholas; Sues, Hans-Dieter; (eds) (1994). "Phylogeny" In the Shadow of the Dinosaurs: Early Mesozoic Tetrapods. Cambridge University Press. ISBN 0-521-45242-2. 
  8. "Sphenodon". Dictionary.com Unabridged (v 1.1). Random House, Inc. Retrieved 8 January 2007. 
  9. 9.0 9.1 9.2 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
  10. 10.0 10.1 Susan E. Evans and Magdalena Borsuk−Białynicka (2009). "A small lepidosauromorph reptile from the Early Triassic of Poland". Paleontologica Polonica 65: 179–202. 
  11. 11.0 11.1 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
  12. 12.0 12.1 Apesteguía S, Novas FE (2003) Large Cretaceous sphenodontian from Patagonia provides insight into lepidosaur evolution in Gondwana. Nature, 425:609–612
  13. "Sphenodontida". Retrieved May 17, 2012. 
  • Daugherty CH, Cree A, Hay JM, Thompson MB (1990). Neglected taxonomy and continuing extinctions of tuatara (Sphenodon). Nature 347, 177–179.
  • Evans SE. 2003. At the feet of the dinosaurs: the early history and radiation of lizards. Biological Reviews, 78:513-551. doi:10.1017/S1464793103006134
  • Jones MEH. 2008. Skull shape and feeding strategy in Sphenodon and other Rhynchocephalia (Diapsida: Lepidosauria). Journal of Morphology. 269: 945–966. doi:10.1002/jmor.10634
  • Jones MEH. 2009. Dentary tooth shape in Sphenodon and its fossil relatives (Diapsida: Lepidosauria: Rhynchocephalia). In Koppe T, Meyer G, Alt KW, eds. Interdisciplinary Dental Morphology, Frontiers of Oral Biology (vol 13). Greifswald, Germany; Karger. 9–15.
  • Evans SE, Jones MEH (2010) The Origin, early history and diversification of lepidosauromorph reptiles. In Bandyopadhyay S. (ed.), New Aspects of Mesozoic Biodiversity, 27 Lecture Notes in Earth Sciences 132, 27-44. doi:10.1007/978-3-642-10311-7_2,

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