Clevosaurus

Abstract: The sphenodontid genus Clevosaurus from the late Triassic of south west Britain is represented by at least two different species. Exceptionally abundant remains of Clevosaurus occur as part of a diverse assemblage of microvertebrates lodged within fissure deposits at localities in Glamorgan and Avon. Although the majority of specimens occur as beautifully preserved disassociated elements, certain associated and articulated remains have been recovered. Contemporaneous vertebrates include other sphenodontians, archosaurs and procolophonids. C. hudsoni is fully restored as a lizard-like reptile approximately 25 cm long. It was essentially insectivorous, although it may have also been facultatively herbivorous, breaking down food with a well-developed precision shear bite. A smaller species, C. minor, is less widespread and its remains are generally not as well preserved. It seems probable that a third species might be distinguished on the basis of dental characteristics, and the occurrence of a fourth species cannot be discounted. A distinct suite of apomorphic characters separates the Sphenodontida from other lepidosauromorphs and these apomorphies are readily identifiable in Clevosaurus. In certain late Triassic sphenodontians the lower temporal arcade was apparently incomplete, but, contrary to Robinson's (J. geol. Soc. Lond. 129, 457 (1973)) description, the lower temporal arcade was unbroken in Clevosaurus. However, the articulation between the jugal, squamosal and quadratojugal does not conform to the configuration exhibited by all other sphenodontians in which both temporal arcades are complete. In addition, the possession of a supratemporal in Clevosaurus is unique among known sphenodontians. The parietal table in Clevosaurus is intermediate between the broad, flat plesiomorphic condition, seen in forms such as Polysphenodon and Homoeosaurus, and the derived narrow median ridge displayed by Kallimodon and Sphenodon. Consequently, within the Sphenodontida the position of Clevosaurus is unclear; however, on the basis of the prominently flanged additional teeth, some reduction in tooth numbers and at least some partial reduction in the width of the parietal table, it would appear to share certain characteristics with Sapheosaurus and Kallimodon. Palaeopleurosaurus, described by Carroll (1985) as a primitive pleurosaur, may also have affinities with this grouping. Some of the earliest sphenodontians, such as Planocephalosaurus and Diphydontosaurus, exhibit fusion of the median skull roofing elements, a short temporal region and retention of a variable degree of pleurodonty in the marginal dentition. Together with Gephyrosaurus these genera may constitute a monophyletic assemblage forming the sister group of all other sphenodontians.

Abstract: The skull and lower jaw of a new sphenodontid reptile Diphydontosaurus avonis is described from disarticulated bones. The fossils were recovered from a detrital limestone of Rhaetian age deposited within solutional and tectonically formed fissures within the Carboniferous Limestone of Tytherington quarry near Bristol. The bone remains, numbering over 1000, are exquisitely preserved with intact facets. Diphydontosaurus was the smallest member of an insular fauna, was primarily insectivorous, and probably formed locally high-density populations. Uniquely for a sphenodontid, Diphydontosaurus had pleurodont teeth on the premaxilla and on the anterior regions of the dentary and maxilla. However, Diphydontosaurus also has the series of acrodont teeth alternating in size on the maxilla and dentary which is characteristic of the Sphenodontidae and particularly Sphenodon. An analysis of 49 synapomorphs in the Diapsida emphasises the sphenodontid nature of Diphydontosaurus. A study of these synapomorphs among the other Triassic sphenodontids Clevosaurus and Planocephalosaurus and the eosuchian Gephyrosaurus suggests that the lack of a quadrate-quadratojugal conch and the complete lower temporal bar are secondarily derived in the `living fossil' Sphenodon punctatus. The tuatara is therefore much less archaic than hitherto proposed. Functional reasons are advocated for the loss of the conch and the regrowth of a complete lower temporal bar. Transformation series are described which could have led to the shape of the maxilla, dentary, premaxilla, palatine and to the loss of the lacrimal in Sphenodon.

Abstract: The Rhynchocephalia are a group of small diapsid reptiles that were globally distributed during the early Mesozoic. By contrast, the only extant representatives, Sphenodon punctatus and S. guntheri (Tuatara), are restricted to New Zealand off-shore islands. The Rhynchocephalia are widely considered to be morphologically uniform but research over the past 30 years has revealed unexpected phenotypic and taxonomic diversity. Phylogenetically "basal taxa" generally possess relatively simple conical or columnar teeth whereas more derived taxa possessed stouter flanged teeth and sophisticated shearing mechanisms: orthal in some (e.g., Clevosaurus hudsoni) and propalinal in others (e.g., S. punctatus). This variation in feeding apparatus suggests a wide range of feeding niches were exploited by rhynchocephalians. The relationship of skull shape to skull length, phylogenetic grouping, habit, and characters relating to the feeding apparatus are explored here with geometric morphometric analysis on two-dimensional landmarks. Principle components analysis demonstrates that there are significant differences between phylogenetic groups. In particular, Sphenodon differs significantly from all well known fossil taxa including the most phylogenetically basal forms. Therefore, it is not justifiable to use Sphenodon as a solitary outgroup when studying skull shape and feeding strategy in squamates; rhynchocephalian fossil taxa also need to be considered. There are also significant differences between the skull shapes of aquatic taxa and those of terrestrial taxa. Of the observed variation in skull shape, most variation is subsumed by variation in dentary tooth base shape, the type of jaw movement employed (e.g., orthal vs. propalinal) and the number of palatal tooth rows. By comparison, the presence or absence of flanges, dentary tooth number and palatal tooth row orientation subsume much less. Skull length was also found to be a poor descriptor of overall skull shape. Compared to basal rhynchocephalians members of more derived terrestrial radiations possess an enlarged postorbital area, a high parietal, and a jaw joint positioned ventral to the tooth row. Modification of these features is closely associated with increased biting performance and thus access to novel food items. Some of these same trends are apparent during Sphenodon ontogeny where skull growth is allometric and there is evidence for ontogenetic variation in diet.