Clevosaurus Restoration
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''Clevosaurus'' (meaning "Gloucester lizard") is an extinct genus of rhynchocephalian reptile from the
Late Triassic The Late Triassic is the third and final epoch of the Triassic Period in the geologic time scale, spanning the time between Ma and Ma (million years ago). It is preceded by the Middle Triassic Epoch and followed by the Early Jurassic Epoch. ...
and the
Early Jurassic The Early Jurassic Epoch (geology), Epoch (in chronostratigraphy corresponding to the Lower Jurassic series (stratigraphy), Series) is the earliest of three epochs of the Jurassic Period. The Early Jurassic starts immediately after the Triassic-J ...
periods. Species of ''Clevosaurus'' were widespread across
Pangaea Pangaea or Pangea () was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous approximately 335 million y ...
, and have been found on all continents except Australia and Antarctica. Five species of ''Clevosaurus'' have been found in ancient fissure fill deposits in south-west England and Wales, alongside other sphenodontians, early mammals and dinosaurs. In regards to its Pangaean distribution, ''C. hadroprodon'' is the oldest record of a sphenodontian from Gondwana, though its affinity to ''Clevosaurus'' has been questioned.


History of discovery

The first species of ''Clevosaurus'' to be described was ''C. hudsoni'', which was described by William Elgin Swinton in 1939 from a fissure fill deposit in Cromhall Quarry ( Magnesian Conglomerate Formation) in the county of
Gloucestershire Gloucestershire ( abbreviated Glos) is a county in South West England. The county comprises part of the Cotswold Hills, part of the flat fertile valley of the River Severn and the entire Forest of Dean. The county town is the city of Gl ...
, England, with the name of the county lending its name to the genus.


Description

''Clevosaurus'' was a medium-sized sphenodontian that was estimated to reach full length of ~25 cm (''Clevosaurus hudsoni''). The skull length could range from as little as 14 mm in ''C. sectumsemper'' and up to 40 mm in ''C. hudsoni''. Notable for their greatly reduced number of teeth (3-6 per jaw quadrant), broad skulls and shortened snouts. The teeth of european clevosaurs tended to be mesio-distally elongated, blade-like, and occluded precisely with the opposite pair of teeth, leaving conspicuous diagonal wear facets and acting as a self-sharpening cutting surface. However, the teeth of ''C. brasiliensis'' have a very different morphology with no diagonal wear facets, the teeth of the dentary are all conical excluding the posterior-most tooth which can be up to three-times bigger than any of the other teeth, they also have a unique form of implantation, where the base of the teeth sit deeply within the jaw bones, which is not known of in any other rhynchocephalian.


Paleobiology

Species of ''Clevosaurus'' were likely insectivorous. Biomechanical modelling suggests that they had high enough tooth pressures and strong enough bite force to crush chitin, indicating that they had the ability to feed on thick-shelled beetles as well as possibly small vertebrates.


List of species

* †''Clevosaurus bairdi'' Sues et al. 1994 McCoy Brook Formation, Canada, Hettangian * †''Clevosaurus brasiliensis'' Bonaparte and Sues 2006
Caturrita Formation The Caturrita Formation is a rock formation found in Rio Grande do Sul, Brazil. Its sediments were deposited in the Paraná Basin. The formation is from the Upper Triassic and forms part of the Santa Maria Supersequence in the upper section of th ...
, Brazil, Norian * †''Clevosaurus cambrica'' Keeble et al. 2018 Pant-y-ffynnon Quarry fissure fill, Wales, Rhaetian * †''Clevosaurus convallis'' Saila 2005 St. Bride's Island fissure fill, Wales, Hettangian * †''Clevosaurus hadroprodon'' Hsiou et al. 2019
Santa Maria Formation The Santa Maria Formation is a sedimentary rock formation found in Rio Grande do Sul, Brazil. It is primarily Carnian in age (Late Triassic), and is notable for its fossils of cynodonts, "rauisuchian" pseudosuchians, and early dinosaurs and other ...
, Brazil,
Carnian The Carnian (less commonly, Karnian) is the lowermost stage (stratigraphy), stage of the Upper Triassic series (stratigraphy), Series (or earliest age (geology), age of the Late Triassic Epoch (reference date), Epoch). It lasted from 237 to 227 m ...
* †''Clevosaurus hudsoni'' Swinton 1939 Cromhall Quarry fissure fill, England, Rhaetian * †''Clevosaurus minor'' Fraser 1988 Cromhall Quarry fissure fill, England, Rhaetian * †''Clevosaurus sectumsemper'' Klein et al. 2015 Woodleaze Quarry fissure fill, England, Rhaetian The three species known from the
Sinemurian In the geologic timescale, the Sinemurian is an age and stage in the Early or Lower Jurassic Epoch or Series. It spans the time between 199.3 ± 2 Ma and 190.8 ± 1.5 Ma (million years ago). The Sinemurian is preceded by the Hettangian and ...
aged
Lufeng Formation The Lufeng Formation (formerly Lower Lufeng Series) is a Lower Jurassic sedimentary rock formation found in Yunnan, China. It has two units: the lower Dull Purplish Beds/Shawan Member are of Hettangian age, and Dark Red Beds/Zhangjia'ao Member ar ...
of China (''C. mcgilli, C.wangi'' and ''C. petilus'') are now considered indeterminate within the genus. Indeterminate remains are also known from the Stormberg Group (either
Elliot Elliot (also spelled Eliot, Elliotte, Elliott, Eliott and Elyot) is a personal name which can serve as either a surname or a given name. Although the given name has historically been given to males, females have increasingly been given the name ...
or
Clarens Formation The Clarens Formation is a geological formation found in several localities in Lesotho and in the Free State, KwaZulu-Natal, and Eastern Cape provinces in South Africa. It is the uppermost of the three formations found in the Stormberg Group of ...
) of South Africa, dating to the Hettangian.


Phylogeny

Below is a
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
of the relationships within Clevosauridae based on the
phylogenetic analysis In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
of Hsiou et al. (2015): ''"Clevosaurus" latidens'' was recovered outside of Clevosauridae, as the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of Opisthodontia. It was subsequently assigned to a new genus, '' Fraserosphenodon'', in 2018.


References


Additional reading


Paleofile
* Gill PG, Säilä LK, Corfe IJ, Challands TJ, Williams M, Clemens WA (2006). The fauna and palaeoenvironment of St. Brides Island: Evidence from the lower Jurassic fissure fills of South Wales. In Barrett PM, Evans SE (eds.). Ninth international symposium on Mesozoic terrestrial ecosystems and biota. pp 48−51. London: Natural History Museum. * Jones MEH (2006) The Early Jurassic clevosaurs from China (Diapsida: Lepidosauria). Natl Mus Nat Hist Sci Bull, 37:548–562. * 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. {{Portal, Paleontology Fossils of China Fossils of Canada Triassic lepidosaurs Jurassic lepidosaurs Sphenodontia Late Triassic first appearances Early Jurassic extinctions Taxa named by William Elgin Swinton Prehistoric reptile genera Paleontology in Nova Scotia