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{{Short description|Extinct genus of reptiles}}
{{Short description|Extinct genus of reptiles}}
{{Automatic Taxobox
{{Automatic taxobox
| fossil_range = [[Late Triassic]] - [[Early Jurassic]] {{fossil_range|Carnian|Sinemurian}}
| fossil_range = [[Late Triassic]] - [[Early Jurassic]] {{fossil_range|Carnian|Sinemurian}}
| image = Clevosaurus.svg
| image = Clevosaurus.svg
| image_caption = Skulls of ''C. hudsoni'' (left) and ''C. cambrica'' (right)
| image_caption = Skulls of ''Clevosaurus hudsoni'' (left) and ''C. cambrica'' (right)
| taxon = Clevosaurus
| taxon = Clevosaurus
| authority = [[William Elgin Swinton|Swinton]], 1939
| authority = [[William Elgin Swinton|Swinton]], 1939
| synonyms = * ''Dianosaurus'' <small>Young, 1982</small>
| synonyms = * ''Dianosaurus'' <small>Young, 1982</small>
| subdivision_ranks = Species
| subdivision_ranks = Species
| subdivision = *''C. bairdi'' <small>Sues, Shubin, & Olson, 1994</small>
| subdivision = See text
*''C. brasiliensis'' <small>Bonaparte & Sues, 2006</small>
*''C. cambrica'' <small>Keeble, Whiteside & [[Michael Benton|Benton]], 2018</small><ref>{{cite journal |author1=Emily Keeble |author2=David I. Whiteside |author3=Michael J. Benton |year=2018 |title=The terrestrial fauna of the Late Triassic Pant-y-ffynnon Quarry fissures, South Wales, UK and a new species of ''Clevosaurus'' (Lepidosauria: Rhynchocephalia) |journal=Proceedings of the Geologists' Association |volume=in press |issue=2 |pages= 99–119|doi=10.1016/j.pgeola.2017.11.001 |url=https://research-information.bristol.ac.uk/en/publications/the-terrestrial-fauna-of-the-late-triassic-pantyffynnon-quarry-fissures-south-wales-uk-and-a-new-species-of-clevosaurus-lepidosauria-rhynchocephalia(5afdc677-3ea0-4519-813d-6052ef8370ec).html |hdl=1983/5afdc677-3ea0-4519-813d-6052ef8370ec }}</ref>
*''C. convallis'' <small>Säilä, 2005</small><ref>{{cite journal|last1=Säilä |first1=Laura K. |year=2005 |title=A new species of the sphenodontian reptile ''Clevosaurus'' from the Lower Jurassic of South Wales |journal=Palaeontology |volume=48 |issue=4 |pages=817–831 |doi=10.1111/j.1475-4983.2005.00486.x|doi-access=free }}</ref>
*''C. hadroprodon'' <small>Hsiou ''et al.'' 2019</small>
*''C. hudsoni'' <small>Swinton, 1939</small> (type)
*''C. mcgilli'' <small>Wu, 1994</small>
*''C. minor'' <small>Fraser & Walkden, 1983</small>
*''C. petilus'' <small>(Young, 1982)</small>
*''C. sectumsempra'' <small>[[Jose Bonaparte|Bonaparte]], 2006</small>
*''C. wangi'' <small>Wu, 1994</small>
}}
}}


'''''Clevosaurus''''' (meaning "Gloucester lizard") is an extinct genus of [[rhynchocephalia]]n reptile from the [[Triassic|Late Triassic]] and the [[Jurassic|Early Jurassic]] [[Period (geology)|periods]].<ref>{{cite web|url=http://novataxa.blogspot.com/2019/08/clevosaurus-hadroprodon.html|title= A New Clevosaurid from the Triassic (Carnian) of Brazil and the Rise of Sphenodontians in Gondwana|date= 19 August 2019}}</ref> Species of ''Clevosaurus'' were widespread across [[Pangaea]], and have been found on all continents except Australia and Antarctica.<ref name=":1">{{cite journal | last1 = Sues | first1 = H. D. | last2 = Reisz | first2 = R. | year = 1995 | title = First record of the early mesozoic sphenodontian clevosaurus (lepidosauria: rhynchocephalia) from the southern hemisphere | journal = Journal of Paleontology | volume = 69 | issue = 1| pages = 123–126 | doi=10.1017/s0022336000026974}}</ref> 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.<ref name=":0">{{Cite web|url=https://phys.org/news/2019-08-sphenodontian-brazil-oldest-group-gondwana.html|title=A new sphenodontian from Brazil is the oldest record of the group in Gondwana|website=phys.org|language=en-us|access-date=2019-08-15}}</ref>[[File:Reconstruction-of-the-skulls-of-lepidosaur-rynchocephalians-A-Clevosaurus-hudsoni.png|left|thumb|Reconstructions of the skulls of ''C. hudsoni'' (A) and ''C. cambrica'' (B), reconstructed areas in orange.]]
'''''Clevosaurus''''' (meaning "Gloucester lizard") is an extinct genus of [[rhynchocephalia]]n reptile from the [[Triassic|Late Triassic]] and the [[Jurassic|Early Jurassic]] [[Period (geology)|periods]]. Species of ''Clevosaurus'' were widespread across [[Pangaea]], and have been found on all continents except Australia and Antarctica.<ref name=":1">{{cite journal | last1 = Sues | first1 = H. D. | last2 = Reisz | first2 = R. | year = 1995 | title = First record of the early mesozoic sphenodontian clevosaurus (lepidosauria: rhynchocephalia) from the southern hemisphere | journal = Journal of Paleontology | volume = 69 | issue = 1| pages = 123–126 | doi=10.1017/s0022336000026974| bibcode = 1995JPal...69..123S | s2cid = 131414921 }}</ref> 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,<ref name=":0">{{Cite web|url=https://phys.org/news/2019-08-sphenodontian-brazil-oldest-group-gondwana.html|title=A new sphenodontian from Brazil is the oldest record of the group in Gondwana|website=phys.org|language=en-us|access-date=2019-08-15}}</ref> though its affinity to ''Clevosaurus'' has been questioned.<ref name="Sofia A. V 2021">{{Cite journal |last1=Chambi-Trowell |first1=Sofia A. V. |last2=Martinelli |first2=Agustín G. |last3=Whiteside |first3=David I. |last4=Vivar |first4=Paulo R. Romo de |last5=Soares |first5=Marina Bento |last6=Schultz |first6=Cesar L. |last7=Gill |first7=Pamela G. |last8=Benton |first8=Michael J. |last9=Rayfield |first9=Emily J. |date=2021-06-03 |title=The diversity of Triassic South American sphenodontians: a new basal form, clevosaurs, and a revision of rhynchocephalian phylogeny |journal=Journal of Systematic Palaeontology |volume=19 |issue=11 |pages=787–820 |doi=10.1080/14772019.2021.1976292 |s2cid=240487298 |issn=1477-2019|doi-access=free |bibcode=2021JSPal..19..787C |hdl=1983/af14affc-a26e-426b-83ca-e1833e355882 |hdl-access=free }}</ref>[[File:Reconstruction-of-the-skulls-of-lepidosaur-rynchocephalians-A-Clevosaurus-hudsoni.png|left|thumb|Reconstructions of the skulls of ''C. hudsoni'' (A) and ''C. cambrica'' (B), reconstructed areas in orange.]]

== History of discovery ==
== 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|Magnesian Conglomerate Formation]]) in the county of [[Gloucestershire]], England, with the name of the county lending its name to the genus.<ref>W. E. Swinton. (1939). A new Triassic rhynchocephalian from Gloucestershire. ''Annals and Magazine of Natural History: Zoology, Botany, and Geology'' '''4''':591-594</ref>
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 [[Slickstones Quarry, Cromhall|Cromhall Quarry]] ([[Magnesian Conglomerate|Magnesian Conglomerate Formation]]) in the county of [[Gloucestershire]], England, with the name of the county lending its name to the genus.<ref>W. E. Swinton. (1939). A new Triassic rhynchocephalian from Gloucestershire. ''Annals and Magazine of Natural History: Zoology, Botany, and Geology'' '''4''':591-594</ref>

Another notable specimen was discovered in 1953 in Cromhall Quarry alongside the holotype of ''[[Cryptovaranoides|Cryptovaranoides microlanius]]''.<ref>{{Cite journal |last1=Whiteside |first1=D. I. |last2=Chambi-Trowell |first2=S. A. V. |last3=Benton |first3=M. J. |date=2022 |title=A Triassic crown squamate |url=https://www.science.org/doi/10.1126/sciadv.abq8274 |journal=Science Advances |language=en |volume=8 |issue=48 |pages=eabq8274 |doi=10.1126/sciadv.abq8274 |pmid=36459546 |bibcode=2022SciA....8.8274W |issn=2375-2548|hdl=1983/a3c7a019-cfe6-4eb3-9ac0-d50c61c5319e |hdl-access=free }}</ref>


== Description ==
== Description ==
[[File:Clevosaurus brasiliensis.jpg|thumb|Skull of ''Clevosaurus brasiliensis''|left]]
[[File:Clevosaurus brasiliensis.jpg|thumb|Skull of ''Clevosaurus brasiliensis''|left]]
The teeth of ''Clevosaurus'' species are elongated and occlude precisely with the opposite pair of teeth, leaving conspicuous wear facets and acting as a self-sharpening cutting surface.<ref name=":2">{{Cite journal|last1=Chambi‐Trowell|first1=Sofia A. V.|last2=Whiteside|first2=David I.|last3=Benton|first3=Michael J.|last4=Rayfield|first4=Emily J.|date=16 June 2020|editor-last=Lautenschlager|editor-first=Stephan|title=Biomechanical properties of the jaws of two species of Clevosaurus and a reanalysis of rhynchocephalian dentary morphospace|url=https://onlinelibrary.wiley.com/doi/10.1111/pala.12493|journal=Palaeontology|language=en|volume=63|issue=6|pages=919–939|doi=10.1111/pala.12493|s2cid=220902843|issn=0031-0239|via=}}</ref>
Species of ''Clevosaurus'' varied in body size, with ''Clevosaurus sectumsemper'' having an estimated total length of {{Convert|12|cm}}, while ''C. hudsoni'' had a total length of around {{Convert|25|cm}}''.'' The skull length could range from as little as {{Convert|1.4|cm}} in ''C. sectumsemper'' and up to {{Convert|4|cm}} in ''C. hudsoni''.<ref name=":3">{{Cite journal |last1=Keeble |first1=Emily |last2=Whiteside |first2=David I. |last3=Benton |first3=Michael J. |date=April 2018 |title=The terrestrial fauna of the Late Triassic Pant-y-ffynnon Quarry fissures, South Wales, UK and a new species of Clevosaurus (Lepidosauria: Rhynchocephalia) |url=https://linkinghub.elsevier.com/retrieve/pii/S0016787817301529 |journal=Proceedings of the Geologists' Association |language=en |volume=129 |issue=2 |pages=99–119 |doi=10.1016/j.pgeola.2017.11.001 |bibcode=2018PrGA..129...99K |hdl-access=free |hdl=1983/5afdc677-3ea0-4519-813d-6052ef8370ec}}</ref> 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.<ref name=":2">{{Cite journal|last1=Chambi-Trowell|first1=Sofia A. V.|last2=Whiteside|first2=David I.|last3=Benton|first3=Michael J.|last4=Rayfield|first4=Emily J.|date=16 June 2020|editor-last=Lautenschlager|editor-first=Stephan|title=Biomechanical properties of the jaws of two species of Clevosaurus and a reanalysis of rhynchocephalian dentary morphospace|url=https://onlinelibrary.wiley.com/doi/10.1111/pala.12493|journal=Palaeontology|language=en|volume=63|issue=6|pages=919–939|doi=10.1111/pala.12493|bibcode=2020Palgy..63..919C |s2cid=220902843|issn=0031-0239|via=}}</ref> 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.<ref name="Sofia A. V 2021"/>

== Paleobiology ==
== Paleobiology ==
[[File:Clevosaurus restoration.jpg|left|thumb|Life restoration of ''Clevosaurus hadroprodon'']]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.<ref name=":2" />
[[File:Clevosaurus restoration.jpg|left|thumb|Life restoration of ''Clevosaurus hadroprodon'']]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.<ref name=":2" />


==Taxonomy==
== List of species ==
At least 9 species of ''Clevosaurus'' are considered valid:

* †''Clevosaurus bairdi'' Sues et al. 1994 [[McCoy Brook Formation]], Canada, [[Hettangian]]
*†''Clevosaurus bairdi'' Sues et al. 1994 [[McCoy Brook Formation]], Canada, [[Hettangian]]
* †''Clevosaurus brasiliensis'' Bonaparte and Sues 2006<ref>{{cite journal | last1 = Bonaparte | first1 = J. F. | last2 = Sues | first2 = H. D. | year = 2006 | title = A new species of clevosaurus (lepidosauria: rhynchocephalia) from the upper Triassic of Rio Grande do Sul, Brazil | journal = Palaeontology | volume = 49 | issue = 1| pages = 917–923 | doi = 10.1111/j.1475-4983.2006.00568.x }}</ref> [[Caturrita Formation]], Brazil, [[Norian]]
* †''Clevosaurus brasiliensis'' Bonaparte and Sues 2006<ref>{{cite journal |last1=Bonaparte |first1=J. F. |last2=Sues |first2=H. D. |year=2006 |title=A new species of clevosaurus (lepidosauria: rhynchocephalia) from the upper Triassic of Rio Grande do Sul, Brazil |journal=Palaeontology |volume=49 |issue=1 |pages=917–923 |doi=10.1111/j.1475-4983.2006.00568.x |s2cid=26483165 |doi-access=free|bibcode=2006Palgy..49..917B }}</ref> [[Caturrita Formation]], Brazil, [[Norian]]
* †''Clevosaurus cambrica'' Keeble et al. 2018 [[Pant-y-Ffynnon Quarry|Pant-y-ffynnon Quarry]] fissure fill, Wales, [[Rhaetian]]
* †''Clevosaurus cambrica'' Keeble et al. 2018<ref name=":3" /> [[Pant-y-Ffynnon Quarry|Pant-y-ffynnon Quarry]] fissure fill, Wales, [[Rhaetian]]
* †''Clevosaurus convallis'' Saila 2005 St. Bride's Island fissure fill, Wales, Hettangian
* †''Clevosaurus convallis'' Saila 2005<ref>{{cite journal |last1=Säilä |first1=Laura K. |year=2005 |title=A new species of the sphenodontian reptile ''Clevosaurus'' from the Lower Jurassic of South Wales |journal=Palaeontology |volume=48 |issue=4 |pages=817–831 |doi=10.1111/j.1475-4983.2005.00486.x |doi-access=free|bibcode=2005Palgy..48..817S }}</ref> St. Bride's Island fissure fill, Wales, Hettangian
* †''Clevosaurus hadroprodon'' Hsiou et al. 2019<ref>{{Cite journal |last1=Hsiou |first1=Annie S. |last2=Nydam |first2=Randall L. |last3=Simões |first3=Tiago R. |last4=Pretto |first4=Flávio A. |last5=Onary |first5=Silvio |last6=Martinelli |first6=Agustín G. |last7=Liparini |first7=Alexandre |last8=Martínez |first8=Paulo R. Romo de Vivar |last9=Soares |first9=Marina B. |last10=Schultz |first10=Cesar L. |last11=Caldwell |first11=Michael W. |date=2019-08-14 |title=A New Clevosaurid from the Triassic (Carnian) of Brazil and the Rise of Sphenodontians in Gondwana |journal=Scientific Reports |language=en |volume=9 |issue=1 |page=11821 |doi=10.1038/s41598-019-48297-9 |issn=2045-2322 |pmc=6694142 |pmid=31413294|bibcode=2019NatSR...911821H }}</ref> [[Santa Maria Formation]], Brazil, [[Carnian]]
* †''Clevosaurus hadroprodon'' Hsiou et al. 2019 [[Santa Maria Formation]], Brazil, [[Carnian]]
* †''Clevosaurus hudsoni'' Swinton 1939 Cromhall Quarry fissure fill, England, Rhaetian
* †''Clevosaurus hudsoni'' Swinton 1939 [[Slickstones Quarry, Cromhall|Cromhall Quarry fissure fill]], England, Rhaetian
* †''Clevosaurus minor'' Fraser 1988 Cromhall Quarry fissure fill, England, Rhaetian
* †''Clevosaurus minor'' Fraser 1988 Cromhall Quarry fissure fill, England, Rhaetian
* †''Clevosaurus sectumsemper'' Klein et al. 2015<ref>{{cite journal |last1=Klein |first1=Catherine G. |last2=Whiteside |first2=David I. |last3=de Lucas |first3=Victor Selles |last4=Viegas |first4=Pedro A. |last5=Benton |first5=Michael J. |title=A distinctive Late Triassic microvertebrate fissure fauna and a new species of ''Clevosaurus'' (Lepidosauria: Rhynchocephalia) from Woodleaze Quarry, Gloucestershire, UK |journal=Proceedings of the Geologists' Association |date=2015 |volume=126 |issue=3 |pages=402–416 |doi=10.1016/j.pgeola.2015.05.003}}</ref> Woodleaze Quarry fissure fill, England, Rhaetian
* †''Clevosaurus sectumsemper'' Klein et al. 2015<ref>{{cite journal |last1=Klein |first1=Catherine G. |last2=Whiteside |first2=David I. |last3=de Lucas |first3=Victor Selles |last4=Viegas |first4=Pedro A. |last5=Benton |first5=Michael J. |date=2015 |title=A distinctive Late Triassic microvertebrate fissure fauna and a new species of ''Clevosaurus'' (Lepidosauria: Rhynchocephalia) from Woodleaze Quarry, Gloucestershire, UK |journal=Proceedings of the Geologists' Association |volume=126 |issue=3 |pages=402–416 |doi=10.1016/j.pgeola.2015.05.003|bibcode=2015PrGA..126..402K }}</ref> Woodleaze Quarry fissure fill, England, Rhaetian
* †''Clevosaurus nicholasi'' Bhat et al. 2023<ref>{{Cite journal |last1=Bhat |first1=Mohd Shafi |last2=Datta |first2=Debajit |last3=Ray |first3=Sanghamitra |last4=Datta |first4=P. M. |date=2023-01-02 |title=A new clevosaurid (Lepidosauria: Rhynchocephalia) from the Upper Triassic of IndiaCitation for this article: Bhat, M. S., Datta, D., Ray, S., & Datta, P. M. (2023) A new clevosaurid (Lepidosauria: Rhynchocephalia) from the Upper Triassic of India. Journal of Vertebrate Paleontology |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2023.2232833 |journal=Journal of Vertebrate Paleontology |language=en |volume=43 |issue=1 |doi=10.1080/02724634.2023.2232833 |bibcode=2023JVPal..43E2833B |s2cid=260525273 |issn=0272-4634}}</ref> [[Tiki Formation]], India, late Carnian-middle Norian
The three species known from the [[Sinemurian]] aged [[Lufeng Formation]] of China (''C. mcgilli, C.wangi'' and ''C. petilus'') are now considered indeterminate within the genus.<ref>Jones MEH. 2006 [https://www.researchgate.net/publication/230865919_The_Early_Jurassic_clevosaurs_from_China_Diapsida_Lepidosauria The Early Jurassic clevosaurs from China (Diapsida: Lepidosauria).] ''New Mex. Museum Nat. Hist. Sci. Bull''. 37, 548–562.</ref> Indeterminate remains are also known from the [[Stormberg Group]] (either [[Elliot Formation|Elliot]] or [[Clarens Formation]]) of South Africa, dating to the Hettangian.<ref name=":1" />
The three species known from the [[Sinemurian]] aged [[Lufeng Formation]] of China (''C. mcgilli, C.wangi'' and ''C. petilus'') are now considered indeterminate within the genus.<ref>Jones MEH. 2006 [https://www.researchgate.net/publication/230865919_The_Early_Jurassic_clevosaurs_from_China_Diapsida_Lepidosauria The Early Jurassic clevosaurs from China (Diapsida: Lepidosauria).] ''New Mex. Museum Nat. Hist. Sci. Bull''. 37, 548–562.</ref> Indeterminate remains are also known from the [[Stormberg Group]] (either [[Elliot Formation|Elliot]] or [[Clarens Formation]]) of South Africa, dating to the Hettangian.<ref name=":1" />

==Species==
Below is a [[cladogram]] of the relationships within Clevosauridae based on the [[phylogenetic analysis]] of Hsiou et al. (2015):<ref name=Hsiouetal2005>{{cite journal |author=Annie Schmaltz Hsiou |author2=Marco Aurélio Gallo De França |author3=Jorge Ferigolo |year=2015 |title=New Data on the ''Clevosaurus'' (Sphenodontia: Clevosauridae) from the Upper Triassic of Southern Brazil |journal=PLOS ONE |volume=10 |issue=9 |pages=e0137523 |doi=10.1371/journal.pone.0137523 |pmid=26355294 |pmc=4565693|bibcode=2015PLoSO..1037523H |doi-access=free }}</ref>
Below is a [[cladogram]] of the relationships within Clevosauridae based on the [[phylogenetic analysis]] of Hsiou et al. (2015):<ref name=Hsiouetal2005>{{cite journal |author=Annie Schmaltz Hsiou |author2=Marco Aurélio Gallo De França |author3=Jorge Ferigolo |year=2015 |title=New Data on the ''Clevosaurus'' (Sphenodontia: Clevosauridae) from the Upper Triassic of Southern Brazil |journal=PLOS ONE |volume=10 |issue=9 |pages=e0137523 |doi=10.1371/journal.pone.0137523 |pmid=26355294 |pmc=4565693|bibcode=2015PLoSO..1037523H |doi-access=free }}</ref>


{{clade| style=font-size:90%;line-height:90%
{{clade| style=font-size:90%;line-height:90%
|label1=&nbsp;[[Clevosauridae]]&nbsp;
|label1=[[Clevosauridae]]
|1={{clade
|1={{clade
|1=''[[Polysphenodon mulleri]]''
|1=''[[Polysphenodon mulleri]]''
Line 63: Line 58:
|3=''Clevosaurus cambrica''}} }} }} }} }}
|3=''Clevosaurus cambrica''}} }} }} }} }}


''"Clevosaurus" latidens'' was recovered outside of Clevosauridae, as the [[sister taxon]] of [[Opisthodontia (rhynchocephalians)|Opisthodontia]].<ref name=Hsiouetal2005/> It was subsequently assigned to a new genus, ''[[Fraserosphenodon]]'', in 2018.<ref>{{cite journal |author1=Jorge A. Herrera-Flores |author2=Thomas L. Stubbs |author3=Armin Elsler |author4=Michael J. Benton |year=2018 |title=Taxonomic reassessment of ''Clevosaurus latidens'' Fraser, 1993 (Lepidosauria, Rhynchocephalia) and rhynchocephalian phylogeny based on parsimony and Bayesian inference |journal=Journal of Paleontology |volume=Online edition |issue= 4|pages= 734–742|doi=10.1017/jpa.2017.136 |doi-access=free }}</ref>
''"Clevosaurus" latidens'' was recovered outside of Clevosauridae, as the [[sister taxon]] of [[Opisthodontia (rhynchocephalians)|Opisthodontia]].<ref name=Hsiouetal2005/> It was subsequently assigned to a new genus, ''[[Fraserosphenodon]]'', in 2018.<ref name=":4">{{cite journal |author1=Jorge A. Herrera-Flores |author2=Thomas L. Stubbs |author3=Armin Elsler |author4=Michael J. Benton |year=2018 |title=Taxonomic reassessment of ''Clevosaurus latidens'' Fraser, 1993 (Lepidosauria, Rhynchocephalia) and rhynchocephalian phylogeny based on parsimony and Bayesian inference |journal=Journal of Paleontology |volume=92 |issue= 4|pages= 734–742|doi=10.1017/jpa.2017.136 |doi-access=free |bibcode=2018JPal...92..734H |hdl=1983/59126b60-16d8-46d2-b657-954693a39d4e |hdl-access=free }}</ref>


''Clevosaurus'' is considered to be a member of the group Eusphenodontia by the groups definition, due to it possessing characters not shared with more primitive sphenodontians. By definition, it is excluded from Neosphenodontia.<ref name=":4" />
==Notes==

{{Reflist}}
Position of ''Clevosaurus'' within Rhynchocephalia, after DeMar et al. 2022.<ref name=":14">{{Cite journal |last1=DeMar |first1=David G. |last2=Jones |first2=Marc E. H. |last3=Carrano |first3=Matthew T. |date=2022-12-31 |title=A nearly complete skeleton of a new eusphenodontian from the Upper Jurassic Morrison Formation, Wyoming, USA, provides insight into the evolution and diversity of Rhynchocephalia (Reptilia: Lepidosauria) |url=https://www.tandfonline.com/doi/full/10.1080/14772019.2022.2093139 |journal=Journal of Systematic Palaeontology |language=en |volume=20 |issue=1 |pages=1–64 |doi=10.1080/14772019.2022.2093139 |issn=1477-2019 |s2cid=252325953 |hdl-access=free |hdl=2440/136608}}</ref>

{{clade|{{clade
|1=''[[Younginia|Younginia capensis]]''
|2=''[[Prolacerta|Prolacerta broomi]]''
|label3=[[Lepidosauria]]
|3={{clade
|label1=[[Squamata|Pan-Squamata]]
|1={{clade
|1=''[[Sophineta|Sophineta cracoviensis]]''
|2=''[[Pristidactylus]]''
|3=''[[Eichstaettisaurus|Eichstaettisaurus schroederi]]''
|4={{clade
|1=''[[Megachirella|Megachirella wachtleri]]''
|2=''[[Marmoretta|Marmoretta oxoniensis]]''}}
}}
|label2=[[Rhynchocephalia]]
|2={{clade
|1=''[[Gephyrosaurus|Gephyrosaurus bridensis]]''
|label2=Sphenodontia
|2={{clade
|1=''[[Diphydontosaurus|Diphydontosaurus avonis]]''
|label2=Acrosphenodontia
|2={{clade
|1=''[[Planocephalosaurus|Planocephalosaurus robinsonae]]''
|2={{clade
|1=''[[Rebbanasaurus|Rebbanasaurus jaini]]''
|2={{clade
|1={{clade
|1=''[[Godavarisaurus|Godavarisaurus lateefi]]''
|2=''[[Theretairus|Theretairus antiquus]]''}}
|label2=Eusphenodontia
|2={{clade
|1=''[[Polysphenodon|Polysphenodon mulleri]]''
|2={{clade
|1=''[[Opisthiamimus|Opisthiamimus gregori]]''
|2={{clade
|label1='''''Clevosaurus'''''
|1={{clade
|1={{clade
|1=''Clevosaurus convallis''
|2={{clade
|1=''Clevosaurus brasiliensis''
|2=''Clevosaurus hadroprodon''}}}}
|2={{clade
|1=''Clevosaurus bairdi''
|2={{clade
|1=''Clevosaurus hudsoni''
|2=''Clevosaurus cambrica''}}}}}}
|label2=Neosphenodontia
|2={{clade
|1=''[[Brachyrhinodon|Brachyrhinodon taylori]]''
|2={{clade
|1=''[[Colobops|Colobops noviportensis]]''
|2={{clade
|label1=[[Sphenodontidae]]
|1={{clade
|1=''[[Sphenodon|Sphenodon punctatus]]''
|2=''[[Cynosphenodon|Cynosphenodon huizachalensis]]''
|3=''[[Sphenovipera|Sphenovipera jimmysjoyi]]''
|4={{clade
|1=''[[Kawasphenodon|Kawasphenodon expectatus]]''
|2=''[[Kawasphenodon|Kawasphenodon peligrensis]]''}}
|5=''[[Pelecymala|Pelecymala robustus]]''
|label6=[[Eilenodontinae]]
|6={{clade
|1={{clade
|1=''[[Fraserosphenodon|Fraserosphenodon latidens]]''
|2=''[[Opisthias|Opisthias rarus]]''}}
|2={{clade
|1=''[[Eilenodon|Eilenodon robustus]]''
|2={{clade
|1=''[[Sphenotitan|Sphenotitan leyesi]]''
|2={{clade
|1=''[[Toxolophosaurus|Toxolophosaurus cloudi]]''
|2=''[[Priosphenodon|Priosphenodon avelasi]]''}}}}}}}}}}
|label2=Leptorhynchia
|2={{clade
|1={{clade
|1=''[[Homoeosaurus|Homoeosaurus maximiliani]]''
|2=''[[Kallimodon|Kallimodon pulchellus]]''}}
|2={{clade
|1={{clade
|1=''[[Sigmala|Sigmala sigmala]]''
|2=''[[Vadasaurus|Vadasaurus herzogi]]''}}
|label2=[[Pleurosauridae]]
|2={{clade
|1=''[[Palaeopleurosaurus|Palaeopleurosaurus posidonae]]''
|2={{clade
|1=''[[Pleurosaurus|Pleurosaurus goldfussi]]''
|2=''[[Pleurosaurus|Pleurosaurus ginsburgi]]''}}}}}}
|3=''[[Kallimodon|Kallimodon cerinensis]]''
|label4=[[Sapheosauridae]]
|4={{clade
|1=''[[Sapheosaurus|Sapheosaurus thiollierei]]''
|2={{clade
|1=''[[Ankylosphenodon|Ankylosphenodon pachyostosus]]''
|2=''[[Oenosaurus|Oenosaurus muehlheimensis]]''}}}}}}}}}}}}}}}}}}}}}}}}}}}}}}}}|style=font-size:85%; line-height:85%}}


==References==
==References==
{{Reflist}}

==Additional reading==
* [https://web.archive.org/web/20080602115308/http://www.paleofile.com/Demo/Mainpage/Taxalist/Sphenodonts.htm Paleofile]
* [https://web.archive.org/web/20080602115308/http://www.paleofile.com/Demo/Mainpage/Taxalist/Sphenodonts.htm 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.
* 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.
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{{Taxonbar|from=Q720817}}


{{Portal|Paleontology}}

[[Category:Mesozoic reptiles of Asia]]
[[Category:Fossils of China]]
[[Category:Fossils of China]]
[[Category:Fossils of Canada]]
[[Category:Fossils of Canada]]
[[Category:Triassic lepidosaurs]]
[[Category:Triassic lepidosaurs]]
[[Category:Jurassic lepidosaurs]]
[[Category:Jurassic lepidosaurs]]
[[Category:Sphenodontia]]
[[Category:Rhynchocephalia]]
[[Category:Late Triassic first appearances]]
[[Category:Late Triassic first appearances]]
[[Category:Early Jurassic extinctions]]
[[Category:Early Jurassic extinctions]]
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[[Category:Prehistoric reptile genera]]
[[Category:Prehistoric reptile genera]]
[[Category:Paleontology in Nova Scotia]]
[[Category:Paleontology in Nova Scotia]]

{{Portal|Paleontology}}

{{triassic-reptile-stub}}
{{Jurassic-reptile-stub}}

Latest revision as of 03:38, 5 December 2024

Clevosaurus
Temporal range: Late Triassic - Early Jurassic Carnian–Sinemurian
Skulls of Clevosaurus hudsoni (left) and C. cambrica (right)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Reptilia
Order: Rhynchocephalia
Family: †Clevosauridae
Genus: †Clevosaurus
Swinton, 1939
Species

See text

Synonyms
  • Dianosaurus Young, 1982

Clevosaurus (meaning "Gloucester lizard") is an extinct genus of rhynchocephalian reptile from the Late Triassic and the Early Jurassic periods. Species of Clevosaurus were widespread across Pangaea, and have been found on all continents except Australia and Antarctica.[1] 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,[2] though its affinity to Clevosaurus has been questioned.[3]

Reconstructions of the skulls of C. hudsoni (A) and C. cambrica (B), reconstructed areas in orange.

History of discovery

[edit]

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, England, with the name of the county lending its name to the genus.[4]

Another notable specimen was discovered in 1953 in Cromhall Quarry alongside the holotype of Cryptovaranoides microlanius.[5]

Description

[edit]
Skull of Clevosaurus brasiliensis

Species of Clevosaurus varied in body size, with Clevosaurus sectumsemper having an estimated total length of 12 centimetres (4.7 in), while C. hudsoni had a total length of around 25 centimetres (9.8 in). The skull length could range from as little as 1.4 centimetres (0.55 in) in C. sectumsemper and up to 4 centimetres (1.6 in) in C. hudsoni.[6] 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.[7] 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.[3]

Paleobiology

[edit]
Life restoration of Clevosaurus hadroprodon

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.[7]

Taxonomy

[edit]

At least 9 species of Clevosaurus are considered valid:

The three species known from the Sinemurian aged Lufeng Formation of China (C. mcgilli, C.wangi and C. petilus) are now considered indeterminate within the genus.[13] Indeterminate remains are also known from the Stormberg Group (either Elliot or Clarens Formation) of South Africa, dating to the Hettangian.[1]

Below is a cladogram of the relationships within Clevosauridae based on the phylogenetic analysis of Hsiou et al. (2015):[14]

Clevosauridae

Polysphenodon mulleri

Brachyrhinodon taylori

Clevosaurus sp. (South Africa)

Clevosaurus convallis

Clevosaurus hudsoni

Clevosaurus petilus

Clevosaurus bairdi

Clevosaurus mcgilli

Clevosaurus wangi

Clevosaurus brasiliensis

Clevosaurus cambrica

"Clevosaurus" latidens was recovered outside of Clevosauridae, as the sister taxon of Opisthodontia.[14] It was subsequently assigned to a new genus, Fraserosphenodon, in 2018.[15]

Clevosaurus is considered to be a member of the group Eusphenodontia by the groups definition, due to it possessing characters not shared with more primitive sphenodontians. By definition, it is excluded from Neosphenodontia.[15]

Position of Clevosaurus within Rhynchocephalia, after DeMar et al. 2022.[16]

Younginia capensis

Prolacerta broomi

Lepidosauria
Pan-Squamata
Rhynchocephalia

Gephyrosaurus bridensis

Sphenodontia

Diphydontosaurus avonis

Acrosphenodontia

Planocephalosaurus robinsonae

Rebbanasaurus jaini

Eusphenodontia

Polysphenodon mulleri

Opisthiamimus gregori

Clevosaurus

Clevosaurus convallis

Clevosaurus brasiliensis

Clevosaurus hadroprodon

Clevosaurus bairdi

Clevosaurus hudsoni

Clevosaurus cambrica

Neosphenodontia

References

[edit]
  1. ^ a b Sues, H. D.; Reisz, R. (1995). "First record of the early mesozoic sphenodontian clevosaurus (lepidosauria: rhynchocephalia) from the southern hemisphere". Journal of Paleontology. 69 (1): 123–126. Bibcode:1995JPal...69..123S. doi:10.1017/s0022336000026974. S2CID 131414921.
  2. ^ "A new sphenodontian from Brazil is the oldest record of the group in Gondwana". phys.org. Retrieved 2019-08-15.
  3. ^ a b Chambi-Trowell, Sofia A. V.; Martinelli, Agustín G.; Whiteside, David I.; Vivar, Paulo R. Romo de; Soares, Marina Bento; Schultz, Cesar L.; Gill, Pamela G.; Benton, Michael J.; Rayfield, Emily J. (2021-06-03). "The diversity of Triassic South American sphenodontians: a new basal form, clevosaurs, and a revision of rhynchocephalian phylogeny". Journal of Systematic Palaeontology. 19 (11): 787–820. Bibcode:2021JSPal..19..787C. doi:10.1080/14772019.2021.1976292. hdl:1983/af14affc-a26e-426b-83ca-e1833e355882. ISSN 1477-2019. S2CID 240487298.
  4. ^ W. E. Swinton. (1939). A new Triassic rhynchocephalian from Gloucestershire. Annals and Magazine of Natural History: Zoology, Botany, and Geology 4:591-594
  5. ^ Whiteside, D. I.; Chambi-Trowell, S. A. V.; Benton, M. J. (2022). "A Triassic crown squamate". Science Advances. 8 (48): eabq8274. Bibcode:2022SciA....8.8274W. doi:10.1126/sciadv.abq8274. hdl:1983/a3c7a019-cfe6-4eb3-9ac0-d50c61c5319e. ISSN 2375-2548. PMID 36459546.
  6. ^ a b Keeble, Emily; Whiteside, David I.; Benton, Michael J. (April 2018). "The terrestrial fauna of the Late Triassic Pant-y-ffynnon Quarry fissures, South Wales, UK and a new species of Clevosaurus (Lepidosauria: Rhynchocephalia)". Proceedings of the Geologists' Association. 129 (2): 99–119. Bibcode:2018PrGA..129...99K. doi:10.1016/j.pgeola.2017.11.001. hdl:1983/5afdc677-3ea0-4519-813d-6052ef8370ec.
  7. ^ a b Chambi-Trowell, Sofia A. V.; Whiteside, David I.; Benton, Michael J.; Rayfield, Emily J. (16 June 2020). Lautenschlager, Stephan (ed.). "Biomechanical properties of the jaws of two species of Clevosaurus and a reanalysis of rhynchocephalian dentary morphospace". Palaeontology. 63 (6): 919–939. Bibcode:2020Palgy..63..919C. doi:10.1111/pala.12493. ISSN 0031-0239. S2CID 220902843.
  8. ^ 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. 49 (1): 917–923. Bibcode:2006Palgy..49..917B. doi:10.1111/j.1475-4983.2006.00568.x. S2CID 26483165.
  9. ^ Säilä, Laura K. (2005). "A new species of the sphenodontian reptile Clevosaurus from the Lower Jurassic of South Wales". Palaeontology. 48 (4): 817–831. Bibcode:2005Palgy..48..817S. doi:10.1111/j.1475-4983.2005.00486.x.
  10. ^ Hsiou, Annie S.; Nydam, Randall L.; Simões, Tiago R.; Pretto, Flávio A.; Onary, Silvio; Martinelli, Agustín G.; Liparini, Alexandre; Martínez, Paulo R. Romo de Vivar; Soares, Marina B.; Schultz, Cesar L.; Caldwell, Michael W. (2019-08-14). "A New Clevosaurid from the Triassic (Carnian) of Brazil and the Rise of Sphenodontians in Gondwana". Scientific Reports. 9 (1): 11821. Bibcode:2019NatSR...911821H. doi:10.1038/s41598-019-48297-9. ISSN 2045-2322. PMC 6694142. PMID 31413294.
  11. ^ Klein, Catherine G.; Whiteside, David I.; de Lucas, Victor Selles; Viegas, Pedro A.; Benton, Michael J. (2015). "A distinctive Late Triassic microvertebrate fissure fauna and a new species of Clevosaurus (Lepidosauria: Rhynchocephalia) from Woodleaze Quarry, Gloucestershire, UK". Proceedings of the Geologists' Association. 126 (3): 402–416. Bibcode:2015PrGA..126..402K. doi:10.1016/j.pgeola.2015.05.003.
  12. ^ Bhat, Mohd Shafi; Datta, Debajit; Ray, Sanghamitra; Datta, P. M. (2023-01-02). "A new clevosaurid (Lepidosauria: Rhynchocephalia) from the Upper Triassic of IndiaCitation for this article: Bhat, M. S., Datta, D., Ray, S., & Datta, P. M. (2023) A new clevosaurid (Lepidosauria: Rhynchocephalia) from the Upper Triassic of India. Journal of Vertebrate Paleontology". Journal of Vertebrate Paleontology. 43 (1). Bibcode:2023JVPal..43E2833B. doi:10.1080/02724634.2023.2232833. ISSN 0272-4634. S2CID 260525273.
  13. ^ Jones MEH. 2006 The Early Jurassic clevosaurs from China (Diapsida: Lepidosauria). New Mex. Museum Nat. Hist. Sci. Bull. 37, 548–562.
  14. ^ a b Annie Schmaltz Hsiou; Marco Aurélio Gallo De França; Jorge Ferigolo (2015). "New Data on the Clevosaurus (Sphenodontia: Clevosauridae) from the Upper Triassic of Southern Brazil". PLOS ONE. 10 (9): e0137523. Bibcode:2015PLoSO..1037523H. doi:10.1371/journal.pone.0137523. PMC 4565693. PMID 26355294.
  15. ^ a b Jorge A. Herrera-Flores; Thomas L. Stubbs; Armin Elsler; Michael J. Benton (2018). "Taxonomic reassessment of Clevosaurus latidens Fraser, 1993 (Lepidosauria, Rhynchocephalia) and rhynchocephalian phylogeny based on parsimony and Bayesian inference". Journal of Paleontology. 92 (4): 734–742. Bibcode:2018JPal...92..734H. doi:10.1017/jpa.2017.136. hdl:1983/59126b60-16d8-46d2-b657-954693a39d4e.
  16. ^ DeMar, David G.; Jones, Marc E. H.; Carrano, Matthew T. (2022-12-31). "A nearly complete skeleton of a new eusphenodontian from the Upper Jurassic Morrison Formation, Wyoming, USA, provides insight into the evolution and diversity of Rhynchocephalia (Reptilia: Lepidosauria)". Journal of Systematic Palaeontology. 20 (1): 1–64. doi:10.1080/14772019.2022.2093139. hdl:2440/136608. ISSN 1477-2019. S2CID 252325953.

Additional reading

[edit]
  • 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.
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