199 lines
21 KiB
XML
199 lines
21 KiB
XML
<document id="C1E6505EF65E42404D14B6F14029C6A1" ID-CLB-Dataset="293415" ID-DOI="10.1080/14772019.2021.1976292" ID-GBIF-Dataset="e34069a7-f0b2-4bad-b150-ccf04d6c843d" ID-ISSN="1478-0941" ID-Zenodo-Dep="10959831" ID-ZooBank="A9211C5A-D4F9-472A-B8AB-877D13ABFDD5" IM.bibliography_approvedBy="diego" IM.illustrations_approvedBy="diego" IM.materialsCitations_approvedBy="diego" IM.metadata_approvedBy="diego" IM.taxonomicNames_approvedBy="diego" IM.treatments_approvedBy="diego" checkinTime="1711485146969" checkinUser="felipe" docAuthor="Chambi-Trowell, Sofia A. V., Martinell, Agust ́ ın G., Whitesid, David I., Romo de Viva, Paulo R., Soare, Marina Bento, Schult, Cesar L., Gil, Pamela G., Bento, Michael J. & Rayfiel, Emily J." docDate="2021" docId="038E87CAFFB52732FF44FD9250A2FAE5" docLanguage="en" docName="JSystPalaeontol.19.11.787-820.pdf" docOrigin="Journal of Systematic Palaeontology 19 (11)" docSource="http://dx.doi.org/10.1080/14772019.2021.1976292" docStyle="DocumentStyle:602474738E3D100F0F13788B53F61545.3:JSystPalaeontol.2016-.journal_article.open" docStyleId="602474738E3D100F0F13788B53F61545" docStyleName="JSystPalaeontol.2016-.journal_article.open" docStyleVersion="3" docTitle="Clevosaurus brasiliensis" docType="treatment" docVersion="4" lastPageNumber="813" masterDocId="FFB7FFB2FFAF2729FFDBFB025255FFBD" masterDocTitle="The diversity of Triassic South American sphenodontians: a new basal form, clevosaurs, and a revision of rhynchocephalian phylogeny" masterLastPageNumber="820" masterPageNumber="787" pageNumber="812" updateTime="1712836126280" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title id="7F1358517D71EE9617DF2D5B8C507C5C">The diversity of Triassic South American sphenodontians: a new basal form, clevosaurs, and a revision of rhynchocephalian phylogeny</mods:title>
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<mods:namePart id="67A05D3AB4D011BF6A5EA1E29AE7E28E">Chambi-Trowell, Sofia A. V.</mods:namePart>
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<mods:affiliation id="714F21B0F66C571C747A97745635FEC0">School of Earth Sciences, University of Bristol, Bristol, BS 8 1 TF, UK;</mods:affiliation>
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<mods:namePart id="4F20789BC49B4193F8DDB413CED3966C">Martinell, Agust ́ ın G.</mods:namePart>
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<mods:namePart id="330AEA245C5F6D51D0081235FE808D87">Whitesid, David I.</mods:namePart>
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<mods:namePart id="5700F39C24A5C1EBECC44A28F5348ED3">Romo de Viva, Paulo R.</mods:namePart>
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<mods:namePart id="F64F5D4726AD0201C4DAEA0E0687CEF6">Soare, Marina Bento</mods:namePart>
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<mods:namePart id="26F8A1B3B7FD5FE9260EC04DE8380941">Schult, Cesar L.</mods:namePart>
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<mods:namePart id="680D660661450B4910194701DDBB056D">Gil, Pamela G.</mods:namePart>
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<mods:namePart id="650A8354824A37BF13A2B06D70BD265D">Bento, Michael J.</mods:namePart>
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<mods:namePart id="6E2F20EE32D11C7FB63C147136B781E6">Rayfiel, Emily J.</mods:namePart>
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<mods:date id="85BB507D0D2FFA71253ED54F193A86E9">2021</mods:date>
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<mods:number id="E2D7166D29FACA7632489F68E8890C3E">2021-11-01</mods:number>
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<treatment id="038E87CAFFB52732FF44FD9250A2FAE5" ID-DOI="http://doi.org/10.5281/zenodo.10959841" ID-Zenodo-Dep="10959841" LSID="urn:lsid:plazi:treatment:038E87CAFFB52732FF44FD9250A2FAE5" httpUri="http://treatment.plazi.org/id/038E87CAFFB52732FF44FD9250A2FAE5" lastPageId="27" lastPageNumber="813" pageId="26" pageNumber="812">
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<subSubSection id="C33D6557FFB52733FF44FD92539DF917" box="[159,456,1680,1706]" pageId="26" pageNumber="812" type="nomenclature">
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<paragraph id="8B9836DCFFB52733FF44FD92539DF917" blockId="26.[159,781,1680,1901]" box="[159,456,1680,1706]" pageId="26" pageNumber="812">
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<heading id="D0D081B0FFB52733FF44FD92539DF917" box="[159,456,1680,1706]" fontSize="11" level="2" pageId="26" pageNumber="812" reason="4">
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<taxonomicName id="4C274D5FFFB52733FF44FD92539DF917" baseAuthorityName="Bonaparte & Sues" baseAuthorityYear="2006" box="[159,456,1680,1706]" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Rhynchocephalia" pageId="26" pageNumber="812" phylum="Chordata" rank="species" species="brasiliensis">
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<emphasis id="B953EACEFFB52733FF44FD92539DF917" bold="true" box="[159,456,1680,1706]" italics="true" pageId="26" pageNumber="812">Clevosaurus brasiliensis</emphasis>
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</taxonomicName>
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</heading>
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</paragraph>
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<subSubSection id="C33D6557FFB52732FF44FDB050A2FAE5" lastPageId="27" lastPageNumber="813" pageId="26" pageNumber="812" type="discussion">
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<paragraph id="8B9836DCFFB52732FF44FDB05337FF49" blockId="26.[159,781,1680,1901]" lastBlockId="27.[159,781,188,1368]" lastPageId="27" lastPageNumber="813" pageId="26" pageNumber="812">
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Our anatomical observations and phylogenetic analysis (
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<figureCitation id="131C2A59FFB52733FF73FDD152A8F957" box="[168,253,1747,1770]" captionStart="Figure 16" captionStartId="26.[159,226,1486,1507]" captionTargetBox="[160,1465,188,1468]" captionTargetId="figure-128@26.[159,1466,188,1469]" captionTargetPageId="26" captionText="Figure 16. Phylogeny of Rhynchocephalia, recovered from: A, strict consensus tree from the maximum parsimony analysis with Bremer values above 1 labelled beside the nodes; and B, 50% majority rule consensus tree from the Bayesian-inference analysis with clade credibility values (decimal proportions) labelled beside nodes." figureDoi="http://doi.org/10.5281/zenodo.10959871" httpUri="https://zenodo.org/record/10959871/files/figure.png" pageId="26" pageNumber="812">Fig. 16</figureCitation>
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) confirm that
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<taxonomicName id="4C274D5FFFB52733FE69FDD15097F957" baseAuthorityName="Bonaparte & Sues" baseAuthorityYear="2006" box="[434,706,1747,1770]" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="812" phylum="Chordata" rank="species" species="brasiliensis">
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<emphasis id="B953EACEFFB52733FE69FDD15097F957" box="[434,706,1747,1770]" italics="true" pageId="26" pageNumber="812">Clevosaurus brasiliensis</emphasis>
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</taxonomicName>
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is an early diverging clevosaur, sharing several characteristics with
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<taxonomicName id="4C274D5FFFB52733FF37FC1753AAF891" box="[236,511,1813,1836]" family="Clevosauridae" genus="Brachyrhinodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="812" phylum="Chordata" rank="species" species="taylori">
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<emphasis id="B953EACEFFB52733FF37FC1753AAF891" box="[236,511,1813,1836]" italics="true" pageId="26" pageNumber="812">Brachyrhinodon taylori</emphasis>
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</taxonomicName>
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, and together with
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<taxonomicName id="4C274D5FFFB52733FF44FC37534CF8F1" box="[159,281,1845,1868]" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="812" phylum="Chordata" rank="species" species="bairdii">
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<emphasis id="B953EACEFFB52733FF44FC37534CF8F1" box="[159,281,1845,1868]" italics="true" pageId="26" pageNumber="812">C. bairdii</emphasis>
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</taxonomicName>
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forms a small subclade at the base of
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<taxonomicName id="4C274D5FFFB52733FF0FFC545325F8D0" authorityName="Bonaparte & Sues" authorityYear="2006" box="[212,368,1878,1901]" family="Clevosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="812" phylum="Chordata" rank="family">Clevosauridae</taxonomicName>
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. Unlike later clevosaurs, both
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<taxonomicName id="4C274D5FFFB52733FC96FD5251BCF9DA" baseAuthorityName="Bonaparte & Sues" baseAuthorityYear="2006" box="[845,1001,1616,1639]" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="812" phylum="Chordata" rank="species" species="brasiliensis">
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<emphasis id="B953EACEFFB52733FC96FD5251BCF9DA" box="[845,1001,1616,1639]" italics="true" pageId="26" pageNumber="812">C. brasiliensis</emphasis>
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</taxonomicName>
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and
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<taxonomicName id="4C274D5FFFB52733FBF9FD5256DDF9DA" box="[1058,1160,1616,1639]" family="Clevosauridae" genus="Brachyrhinodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="812" phylum="Chordata" rank="species" species="taylori">
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<emphasis id="B953EACEFFB52733FBF9FD5256DDF9DA" box="[1058,1160,1616,1639]" italics="true" pageId="26" pageNumber="812">B. taylori</emphasis>
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</taxonomicName>
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bear three rows of teeth on their pterygoids (
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<collectionCode id="ED36AE19FFB52733FBD4FD735621F935" box="[1039,1140,1649,1672]" pageId="26" pageNumber="812">SAVC-T</collectionCode>
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, pers. obs.), the lateral-most of which is reduced to just two to three teeth. In addition, both taxa possess robust jaws, short snouts, broad skulls, a remarkably elongated retroarticular process, two to three large additional teeth on the maxilla, and a high coronoid process. Unfortunately, the nature of the implantation, tooth form, and number – features that mark
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<taxonomicName id="4C274D5FFFB52733FC54FC545678F8D0" baseAuthorityName="Bonaparte & Sues" baseAuthorityYear="2006" box="[911,1069,1878,1901]" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="812" phylum="Chordata" rank="species" species="brasiliensis">
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<emphasis id="B953EACEFFB52733FC54FC545678F8D0" box="[911,1069,1878,1901]" italics="true" pageId="26" pageNumber="812">C. brasiliensis</emphasis>
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</taxonomicName>
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as distinct from other
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<taxonomicName id="4C274D5FFFB52733FAEEFC5457EFF8D0" authorityName="Swinton" authorityYear="1939" box="[1333,1466,1878,1901]" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="812" phylum="Chordata" rank="genus">
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<emphasis id="B953EACEFFB52733FAEEFC5457EFF8D0" box="[1333,1466,1878,1901]" italics="true" pageId="26" pageNumber="812">Clevosaurus</emphasis>
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</taxonomicName>
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– cannot be verified in
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<taxonomicName id="4C274D5FFFB42732FE78FBBF505EFF69" box="[419,523,189,212]" family="Clevosauridae" genus="Brachyrhinodon" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="813" phylum="Chordata" rank="species" species="taylori">
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<emphasis id="B953EACEFFB42732FE78FBBF505EFF69" box="[419,523,189,212]" italics="true" pageId="27" pageNumber="813">B. taylori</emphasis>
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</taxonomicName>
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as these characteristics are not preserved.
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</paragraph>
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<paragraph id="8B9836DCFFB42732FF62FBFF5041FB8A" blockId="27.[159,781,188,1368]" pageId="27" pageNumber="813">
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The teeth of
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<taxonomicName id="4C274D5FFFB42732FE95FBFF53BFFEA9" baseAuthorityName="Bonaparte & Sues" baseAuthorityYear="2006" box="[334,490,253,276]" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="813" phylum="Chordata" rank="species" species="brasiliensis">
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<emphasis id="B953EACEFFB42732FE95FBFF53BFFEA9" box="[334,490,253,276]" italics="true" pageId="27" pageNumber="813">C. brasiliensis</emphasis>
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</taxonomicName>
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differ from those of later clevosaurs in morphology and implantation, appearing to be acrodont but nested deep within the jaw bones so that only a small portion of the tooth is visible. Rather than forming a jagged, narrow, semi-continuous blade, posteriorly the teeth of
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<taxonomicName id="4C274D5FFFB42732FEB3FA9F5051FE09" baseAuthorityName="Bonaparte & Sues" baseAuthorityYear="2006" box="[360,516,413,436]" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="813" phylum="Chordata" rank="species" species="brasiliensis">
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<emphasis id="B953EACEFFB42732FEB3FA9F5051FE09" box="[360,516,413,436]" italics="true" pageId="27" pageNumber="813">C. brasiliensis</emphasis>
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</taxonomicName>
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are closely packed and fused together, and both the cusps and dentary are worn at an angle as the animal ages. This means that the anterior hatchling teeth form a continuous narrow cropping beak-like structure. Most
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<taxonomicName id="4C274D5FFFB42732FE63F91C50C4FD88" box="[440,657,542,565]" pageId="27" pageNumber="813">
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<emphasis id="B953EACEFFB42732FE63F91C506EFD88" box="[440,571,542,565]" italics="true" pageId="27" pageNumber="813">Clevosaurus</emphasis>
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species
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</taxonomicName>
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show additional dentary teeth with a mesiodistally elongated saddle shape and the cusp asymmetrically placed posteriorly on the tooth displaying large anterolateral flanges (
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<bibRefCitation id="EFB64B2DFFB42732FD74F97C5324FD08" author="Chambi-Trowell, S. A. V. & Whiteside, D. I. & Benton, M. J. & Rayfield, E. J." pageId="27" pageNumber="813" pagination="919 - 939" refId="ref21413" refString="Chambi-Trowell, S. A. V., Whiteside, D. I., Benton, M. J. & Rayfield, E. J. 2020. Biomechanical properties of the jaws of two species of Clevosaurus and a reanalysis of rhynchocephalian dentary morphospace. Palaeontology, 63, 919 - 939. doi: 10.1111 / pala. 12493" type="journal article" year="2020">
|
||
Chambi-Trowell
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<emphasis id="B953EACEFFB42732FF27F99C5378FD08" box="[252,301,670,693]" italics="true" pageId="27" pageNumber="813">et al</emphasis>
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. 2020
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</bibRefCitation>
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). However, the additional teeth of
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<taxonomicName id="4C274D5FFFB42732FD2EF99C5342FD68" baseAuthorityName="Bonaparte & Sues" baseAuthorityYear="2006" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="813" phylum="Chordata" rank="species" species="brasiliensis">
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<emphasis id="B953EACEFFB42732FD2EF99C5342FD68" italics="true" pageId="27" pageNumber="813">C. brasiliensis</emphasis>
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</taxonomicName>
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are mostly conical, with only the ultimate tooth – which can be up to two to three times bigger than any of the other teeth
|
||
<emphasis id="B953EACEFFB42732FE1CF9FD5381FCAB" box="[455,468,767,790]" italics="true" pageId="27" pageNumber="813">–</emphasis>
|
||
elongated mesiodistally, and with its cusp placed anteriorly rather than posteriorly (two features unique to
|
||
<taxonomicName id="4C274D5FFFB42732FE6CF83D5003FCEB" baseAuthorityName="Bonaparte & Sues" baseAuthorityYear="2006" box="[439,598,831,854]" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="813" phylum="Chordata" rank="species" species="brasiliensis">
|
||
<emphasis id="B953EACEFFB42732FE6CF83D5003FCEB" box="[439,598,831,854]" italics="true" pageId="27" pageNumber="813">C. brasiliensis</emphasis>
|
||
</taxonomicName>
|
||
amongst clevosaurs, but also shared with
|
||
<taxonomicName id="4C274D5FFFB42732FE15F85D50F0FCCB" baseAuthorityName="Fraser" baseAuthorityYear="1982" box="[462,677,863,886]" class="Sphenodontia" family="Sphenodontidae" genus="Planocephalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="813" phylum="Chordata" rank="genus">
|
||
<emphasis id="B953EACEFFB42732FE15F85D50F0FCCB" box="[462,677,863,886]" italics="true" pageId="27" pageNumber="813">Planocephalosaurus</emphasis>
|
||
</taxonomicName>
|
||
). In contrast to other clevosaurs, the teeth on the dentary of
|
||
<taxonomicName id="4C274D5FFFB42732FD2EF87D5342FC0B" baseAuthorityName="Bonaparte & Sues" baseAuthorityYear="2006" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="813" phylum="Chordata" rank="species" species="brasiliensis">
|
||
<emphasis id="B953EACEFFB42732FD2EF87D5342FC0B" italics="true" pageId="27" pageNumber="813">C. brasiliensis</emphasis>
|
||
</taxonomicName>
|
||
do not appear to bear flanges at all. But despite these differences,
|
||
<taxonomicName id="4C274D5FFFB42732FE44F8BD5068FC6B" baseAuthorityName="Bonaparte & Sues" baseAuthorityYear="2006" box="[415,573,959,982]" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="813" phylum="Chordata" rank="species" species="brasiliensis">
|
||
<emphasis id="B953EACEFFB42732FE44F8BD5068FC6B" box="[415,573,959,982]" italics="true" pageId="27" pageNumber="813">C. brasiliensis</emphasis>
|
||
</taxonomicName>
|
||
is otherwise very similar to later clevosaurs, with a short robust snout, broad skull, supratemporal bone, premaxillary beak and an elongated retroarticular process.
|
||
</paragraph>
|
||
<paragraph id="8B9836DCFFB42732FF62FF4250A2FAE5" blockId="27.[159,781,188,1368]" pageId="27" pageNumber="813">
|
||
Some of the skull traits of
|
||
<taxonomicName id="4C274D5FFFB42732FE3EFF4250D7FBEA" baseAuthorityName="Bonaparte & Sues" baseAuthorityYear="2006" box="[485,642,1088,1111]" family="Clevosauridae" genus="Clevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="813" phylum="Chordata" rank="species" species="brasiliensis">
|
||
<emphasis id="B953EACEFFB42732FE3EFF4250D7FBEA" box="[485,642,1088,1111]" italics="true" pageId="27" pageNumber="813">C. brasiliensis</emphasis>
|
||
</taxonomicName>
|
||
suggest that it must have been capable of a strong bite, such as its robust and short snout that seems to be further reinforced by its brace-like premaxillae and the slot-like facet on the prefrontal so that the maxilla is supported both medially and laterally. A similar slot-like facet is observed between the postorbital and jugal, with the bone interdigitating along their meeting edge rather than the simpler overlapping facets seen in later clevosaurs.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |