796 lines
116 KiB
XML
796 lines
116 KiB
XML
<document ID-DOI="10.1093/zoolinnean/zlac024" ID-GBIF-Dataset="da4f79ec-0c6f-41b0-a822-bbcb7b00aad0" ID-ISSN="0024-4082" ID-Zenodo-Dep="7695193" ID-ZooBank="8C786137-0AAF-40C5-ABDC-51477AFC4023" checkinTime="1677743825978" checkinUser="plazi" docAuthor="Scarpetta, Simon G. & Ledesma, David T." docDate="2023" docId="03FE87CFFFA8933BD4BDF961FAEFE939" docLanguage="en" docName="zlac024.pdf" docOrigin="Zoological Journal of the Linnean Society 197" docStyle="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-.journal_article" docStyleId="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2017-.journal_article" docStyleVersion="7" docTitle="Abronia cuyama Scarpetta & Ledesma 2023, SP. NOV." docType="treatment" docVersion="2" lastPageNumber="761" masterDocId="FFC7FFB7FFAB9332D702FFA3FFBDEB34" masterDocTitle="A strikingly ornamented fossil alligator lizard (Squamata: Abronia) from the Miocene of California" masterLastPageNumber="767" masterPageNumber="752" pageNumber="755" updateTime="1677838144061" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>A strikingly ornamented fossil alligator lizard (Squamata: Abronia) from the Miocene of California</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Scarpetta, Simon G.</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Ledesma, David T.</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>Zoological Journal of the Linnean Society</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2023</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2022-04-30</mods:number>
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</mods:detail>
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<mods:detail type="volume">
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<mods:number>197</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>752</mods:start>
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<mods:end>767</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1093/zoolinnean/zlac024</mods:identifier>
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<mods:identifier type="GBIF-Dataset">da4f79ec-0c6f-41b0-a822-bbcb7b00aad0</mods:identifier>
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<mods:identifier type="ISSN">0024-4082</mods:identifier>
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<mods:identifier type="Zenodo-Dep">7695193</mods:identifier>
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<mods:identifier type="ZooBank">8C786137-0AAF-40C5-ABDC-51477AFC4023</mods:identifier>
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</mods:mods>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.7695195" ID-Zenodo-Dep="7695195" LSID="urn:lsid:plazi:treatment:03FE87CFFFA8933BD4BDF961FAEFE939" httpUri="http://treatment.plazi.org/id/03FE87CFFFA8933BD4BDF961FAEFE939" lastPageId="9" lastPageNumber="761" pageId="3" pageNumber="755">
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<subSubSection box="[959,1275,1730,1754]" pageId="3" pageNumber="755" type="nomenclature">
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<paragraph blockId="3.[959,1275,1730,1754]" box="[959,1275,1730,1754]" pageId="3" pageNumber="755">
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<heading box="[959,1275,1730,1754]" centered="true" fontSize="9" level="2" pageId="3" pageNumber="755" reason="2">
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<taxonomicName authority="Scarpetta & Ledesma, 2023" authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[959,1169,1730,1754]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="763" phylum="Chordata" rank="species" species="cuyama" status="sp. nov.">ABRONIA CUYAMA</taxonomicName>
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<emphasis bold="true" box="[1176,1275,1730,1754]" pageId="3" pageNumber="755">
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<taxonomicNameLabel box="[1176,1275,1730,1754]" pageId="3" pageNumber="755" rank="species">SP. NOV.</taxonomicNameLabel>
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</emphasis>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="3" pageNumber="755" type="description">
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<paragraph blockId="3.[809,1426,1770,1823]" pageId="3" pageNumber="755">
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<emphasis box="[809,1143,1770,1792]" italics="true" pageId="3" pageNumber="755">Z o o b a n k r e g i s t r a t i o n:</emphasis>
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u r n: l s i d: z o o b a n k. org:pub:
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<uuid box="[893,1421,1801,1823]" pageId="3" pageNumber="755">8C786137-0AAF-40C5-ABDC-51477AFC4023</uuid>
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="8" lastPageNumber="760" pageId="3" pageNumber="755" type="materials_examined">
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<paragraph blockId="3.[809,1425,1864,1916]" lastBlockId="4.[163,780,197,434]" lastPageId="4" lastPageNumber="756" pageId="3" pageNumber="755">
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<materialsCitation collectionCode="UCMP" collectorName="Gideon T. James" lastPageId="4" lastPageNumber="756" pageId="3" pageNumber="755" specimenCode="UCMP 54560" specimenCount="2" typeStatus="holotype">
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<emphasis box="[809,922,1864,1885]" italics="true" pageId="3" pageNumber="755">Holotype:</emphasis>
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The
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<typeStatus box="[991,1094,1864,1886]" pageId="3" pageNumber="755">holotype</typeStatus>
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<emphasis box="[1104,1266,1864,1885]" italics="true" pageId="3" pageNumber="755">
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<specimenCode box="[1104,1266,1864,1885]" collectionCode="UCMP" country="USA" httpUri="http://grbio.org/cool/2fxn-eays" name="University of California Museum of Paleontology" pageId="3" pageNumber="755" type="University or college">UCMP 54560</specimenCode>
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</emphasis>
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is housed in the
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<collectionCode box="[854,932,1895,1916]" country="USA" httpUri="http://grbio.org/cool/2fxn-eays" name="University of California Museum of Paleontology" pageId="3" pageNumber="755" type="University or college">UCMP</collectionCode>
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and was collected by
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<collectorName box="[1184,1266,1894,1916]" pageId="3" pageNumber="755">Gideon</collectorName>
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<collectionCode box="[1273,1290,1895,1916]" name="Tavera, Department of Geology and Geophysics" pageId="3" pageNumber="755">T</collectionCode>
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.
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<collectorName box="[1301,1374,1895,1916]" pageId="3" pageNumber="755">James</collectorName>
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and a party of
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<collectionCode box="[290,370,197,218]" country="USA" httpUri="http://grbio.org/cool/2fxn-eays" name="University of California Museum of Paleontology" pageId="4" pageNumber="756" type="University or college">UCMP</collectionCode>
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palaeontologists in 1957 or 1958.
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<collectorName box="[163,207,228,250]" pageId="4" pageNumber="756">The</collectorName>
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specimen consists of a partial cranium including the anterior portion of the skull (
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<figureCitation box="[570,691,258,280]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="4" pageNumber="756">Figs 2A–F</figureCitation>
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,
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<figureCitation captionStart="Figure 3" captionStartId="6.[163,241,1285,1307]" captionTargetBox="[165,1446,197,1244]" captionTargetId="figure-243@6.[163,1446,195,1245]" captionTargetPageId="6" captionText="Figure 3. Selected cranial elements of Abronia cuyama sp. nov. A, B, premaxilla in anterior and posterior view, respectively. C, D, left maxilla in lateral and dorsal view, respectively. E, left septomaxilla in dorsal view. F, right quadrate and separate pieces of right squamosal in lateral view. G, right vomer in lateral view. H, right jugal in lateral view. I, vomerine process of the left palatine in medial view. Scale bars: 1 mm. Abbreviations: a.pr, anterior process; d.o, dorsal ossification; f.pr, facial process; J.s, jugal spur; La, lacrimal; mx.lp, maxillary lappet; mx.pr, maxillary process; n.pr, nasal process; o.pr, orbital process; pa.pr, palatine process; pd.fl, posterodorsal flange; p.pr.Smx, posterior process of the septomaxilla; p.pr.Sq, posterior process of the squamosal; t.r, temporal ramus; v.k, ventral keel; v.l, ventral lamina; vn, vomeronasal region." figureDoi="http://doi.org/10.5281/zenodo.7695201" httpUri="https://zenodo.org/record/7695201/files/figure.png" pageId="4" pageNumber="756">3A–E, G–I</figureCitation>
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,
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<figureCitation box="[225,258,289,310]" captionStart="Figure 4" captionStartId="7.[143,221,1035,1057]" captionTargetBox="[225,1344,197,994]" captionTargetId="figure-481@7.[223,1346,195,995]" captionTargetPageId="7" captionText="Figure 4. Comparison of Abronia cuyama sp. nov. and skulls of extant gerrhonotines, in anterior view. A, Abronia cuyama sp. nov. UCMP 54560. B, Abronia graminea UTA 38831. C, Elgaria multicarinata TNHC 35666. D, Barisia leƲicollis MVZ 68783. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7695203" httpUri="https://zenodo.org/record/7695203/files/figure.png" pageId="4" pageNumber="756">4A</figureCitation>
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), an associated block of mudstone matrix containing a few posterior cranial elements (
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<figureCitation box="[662,773,320,342]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="4" pageNumber="756">Figs 2G–I</figureCitation>
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,
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<figureCitation box="[163,194,351,372]" captionStart="Figure 3" captionStartId="6.[163,241,1285,1307]" captionTargetBox="[165,1446,197,1244]" captionTargetId="figure-243@6.[163,1446,195,1245]" captionTargetPageId="6" captionText="Figure 3. Selected cranial elements of Abronia cuyama sp. nov. A, B, premaxilla in anterior and posterior view, respectively. C, D, left maxilla in lateral and dorsal view, respectively. E, left septomaxilla in dorsal view. F, right quadrate and separate pieces of right squamosal in lateral view. G, right vomer in lateral view. H, right jugal in lateral view. I, vomerine process of the left palatine in medial view. Scale bars: 1 mm. Abbreviations: a.pr, anterior process; d.o, dorsal ossification; f.pr, facial process; J.s, jugal spur; La, lacrimal; mx.lp, maxillary lappet; mx.pr, maxillary process; n.pr, nasal process; o.pr, orbital process; pa.pr, palatine process; pd.fl, posterodorsal flange; p.pr.Smx, posterior process of the septomaxilla; p.pr.Sq, posterior process of the squamosal; t.r, temporal ramus; v.k, ventral keel; v.l, ventral lamina; vn, vomeronasal region." figureDoi="http://doi.org/10.5281/zenodo.7695201" httpUri="https://zenodo.org/record/7695201/files/figure.png" pageId="4" pageNumber="756">3F</figureCitation>
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) and two vials of associated material containing osteoderms in matrix, a probable epipterygoid and unidentified bones.
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</materialsCitation>
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||
</paragraph>
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||
<paragraph blockId="4.[163,779,474,650]" pageId="4" pageNumber="756">
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<emphasis box="[163,297,474,495]" italics="true" pageId="4" pageNumber="756">Etymology:</emphasis>
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The species is named for the Cuyama Valley Badlands where
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<emphasis box="[447,606,505,526]" italics="true" pageId="4" pageNumber="756">
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<collectionCode box="[447,526,505,526]" country="USA" httpUri="http://grbio.org/cool/2fxn-eays" name="University of California Museum of Paleontology" pageId="4" pageNumber="756" type="University or college">UCMP</collectionCode>
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54560
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</emphasis>
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was collected. ‘Cuyama’ is derived from the Chumash word
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<emphasis box="[700,779,536,557]" italics="true" pageId="4" pageNumber="756">kuyam</emphasis>
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(noun) that means ‘a place to come together’ or ‘clam’; the name of the new taxon honours the Chumash people on whose land the fossil was collected.
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</paragraph>
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<paragraph blockId="4.[163,780,690,1540]" pageId="4" pageNumber="756">
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<emphasis box="[163,499,690,712]" italics="true" pageId="4" pageNumber="756">
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Diagnosis:
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<taxonomicName authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[301,499,690,712]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="756" phylum="Chordata" rank="species" species="cuyama">Abronia cuyama</taxonomicName>
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</emphasis>
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||
is a squamate because it has a single premaxilla and pleurodont tooth implantation and an anguimorph because the Meckelian groove is directed ventrally (
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||
<bibRefCitation author="Estes R & De Queiroz K & Gauthier JA" pageId="4" pageNumber="756" pagination="119 - 281" refId="ref9720" refString="Estes R, De Queiroz K, Gauthier JA. 1988. Phylogenetic relationships within Squamata. In: Estes R, Pregill GK, eds. Phylogenetic relationships of the lizard families: essays commemorating Charles L. Camp. Stanford: Stanford University Press, 119 - 281." type="book chapter" year="1988">
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Estes
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<emphasis box="[713,772,782,804]" italics="true" pageId="4" pageNumber="756">et al.</emphasis>
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||
, 1988
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||
</bibRefCitation>
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||
;
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||
<bibRefCitation author="Gauthier JA & Estes R & De Queiroz K" box="[234,480,813,835]" pageId="4" pageNumber="756" pagination="15 - 98" refId="ref10146" refString="Gauthier JA, Estes R, De Queiroz K. 1988. Phylogenetic relationships within Squamata. In: Estes R, Pregill GK, eds. Phylogenetic relationships of the lizard families: essays commemorating Charles L. Camp. Stanford: Stanford University Press, 15 - 98." type="book chapter" year="1988">
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Gauthier
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<emphasis box="[350,409,813,835]" italics="true" pageId="4" pageNumber="756">et al.</emphasis>
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, 1988
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</bibRefCitation>
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). The taxon is an anguid because there are rectangular and laterally imbricating osteoderms, a free margin of the intramandibular septum (
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<bibRefCitation author="Gauthier JA" box="[266,440,905,927]" pageId="4" pageNumber="756" pagination="7 - 54" refId="ref10106" refString="Gauthier JA. 1982. Fossil xenosaurid and anguid lizards from the early Eocene Wasatch Formation, southeast Wyoming, and a revision of the Anguioidea. Contributions to Geology, UniVersity of Wyoming 21: 7 - 54." type="journal article" year="1982">Gauthier, 1982</bibRefCitation>
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) and a squamosal that lacks a posteromedial expansion (
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<bibRefCitation author="Bhullar B-AS" box="[486,641,936,958]" pageId="4" pageNumber="756" pagination="65 - 181" refId="ref9254" refString="Bhullar B-AS. 2011. The power and utility of morphological characters in systematics: a fully resolved phylogeny of Xenosaurus and its fossil relatives (Squamata: Anguimorpha). Bulletin of the Museum of ComparatiVe Zoology 160: 65 - 181." type="journal article" year="2011">Bhullar, 2011</bibRefCitation>
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||
). The taxon is assigned to
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<taxonomicName authorityName="TIHEN" authorityYear="1949" box="[346,526,966,988]" class="Reptilia" family="Gerrhonotinae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="756" phylum="Chordata" rank="family">Gerrhonotinae</taxonomicName>
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because the frontal is fused and lacks suture marks (
|
||
<bibRefCitation author="Klembara J & Bohme M & Rummel M" pageId="4" pageNumber="756" pagination="159 - 196" refId="ref10764" refString="Klembara J, Bohme M, Rummel M. 2010. Revision of the anguine lizard Pseudopus laurillardi (Squamata, Anguidae) from the Miocene of Europe, with comments on paleoecology. Journal of Paleontology 84: 159 - 196." type="journal article" year="2010">
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Klembara
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<emphasis box="[710,772,997,1019]" italics="true" pageId="4" pageNumber="756">et al.</emphasis>
|
||
, 2010
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||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Scarpetta SG & Ledesma DT & Bell CJ" box="[233,484,1028,1049]" pageId="4" pageNumber="756" pagination="1 - 58" refId="ref11478" refString="Scarpetta SG, Ledesma DT, Bell CJ. 2021. A new extinct species of alligator lizard (Squamata: Elgaria) and an expanded perspective on the osteology and phylogeny of Gerrhonotinae. BMC Ecology and EVolution 21: 1 - 58." type="journal article" year="2021">
|
||
Scarpetta
|
||
<emphasis box="[355,413,1028,1049]" italics="true" pageId="4" pageNumber="756">et al.</emphasis>
|
||
, 2021
|
||
</bibRefCitation>
|
||
), there is a raised dorsal ossification on each side of the body of the premaxilla (
|
||
<bibRefCitation author="Scarpetta S" box="[171,356,1089,1110]" pageId="4" pageNumber="756" pagination="1 - 9" refId="ref11399" refString="Scarpetta S. 2018. The earliest known occurrence of Elgaria (Squamata: Anguidae) and a minimum age for crown Gerrhonotinae: fossils from the Split Rock Formation, Wyoming, USA. Palaeontologia Electronica 21.1.1 FC: 1 - 9." type="book chapter" year="2018">Scarpetta, 2018</bibRefCitation>
|
||
), and there are no separate palatal processes of the premaxilla (
|
||
<bibRefCitation author="Evans SE" box="[505,649,1120,1141]" pageId="4" pageNumber="756" pagination="1 - 347" refId="ref9771" refString="Evans SE. 2008. The skull of lizards and tuatara. In: Gans C, Gaunt AS, Adler K, eds. Biology of the Reptilia, Vol. 20, morphology H: the skull of Lepidosauria. Ithaca: Society for the Study of Amphibians and Reptiles, 1 - 347." type="book chapter" year="2008">Evans, 2008</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Scarpetta SG & Ledesma DT & Bell CJ" pageId="4" pageNumber="756" pagination="1 - 58" refId="ref11478" refString="Scarpetta SG, Ledesma DT, Bell CJ. 2021. A new extinct species of alligator lizard (Squamata: Elgaria) and an expanded perspective on the osteology and phylogeny of Gerrhonotinae. BMC Ecology and EVolution 21: 1 - 58." type="journal article" year="2021">
|
||
Scarpetta
|
||
<emphasis box="[163,226,1150,1172]" italics="true" pageId="4" pageNumber="756">et al.</emphasis>
|
||
, 2021
|
||
</bibRefCitation>
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||
). The new taxon shares with
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[680,778,1150,1171]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="756" phylum="Chordata" rank="genus">
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||
<emphasis box="[680,778,1150,1171]" italics="true" pageId="4" pageNumber="756">Abronia</emphasis>
|
||
</taxonomicName>
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||
excluding species previously placed in
|
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<taxonomicName authority="(Scarpetta et al., 2021)" baseAuthorityName="Scarpetta" baseAuthorityYear="2021" class="Reptilia" family="Anguidae" genus="Mesaspis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="756" phylum="Chordata" rank="genus">
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||
<emphasis box="[665,778,1182,1203]" italics="true" pageId="4" pageNumber="756">Mesaspis</emphasis>
|
||
(
|
||
<bibRefCitation author="Scarpetta SG & Ledesma DT & Bell CJ" box="[172,443,1212,1233]" pageId="4" pageNumber="756" pagination="1 - 58" refId="ref11478" refString="Scarpetta SG, Ledesma DT, Bell CJ. 2021. A new extinct species of alligator lizard (Squamata: Elgaria) and an expanded perspective on the osteology and phylogeny of Gerrhonotinae. BMC Ecology and EVolution 21: 1 - 58." type="journal article" year="2021">
|
||
Scarpetta
|
||
<emphasis box="[303,367,1212,1233]" italics="true" pageId="4" pageNumber="756">et al.</emphasis>
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||
, 2021
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</bibRefCitation>
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||
)
|
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</taxonomicName>
|
||
a relatively wide frontal, vermiculate, heavily sculptured osteoderms across the skull, contact of the posterior internasal and prefrontal osteoderms, and the presence of a frontonasal osteoderm (
|
||
<figureCitation box="[303,424,1334,1356]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="4" pageNumber="756">Figs 2A–F</figureCitation>
|
||
,
|
||
<figureCitation box="[439,471,1334,1355]" captionStart="Figure 4" captionStartId="7.[143,221,1035,1057]" captionTargetBox="[225,1344,197,994]" captionTargetId="figure-481@7.[223,1346,195,995]" captionTargetPageId="7" captionText="Figure 4. Comparison of Abronia cuyama sp. nov. and skulls of extant gerrhonotines, in anterior view. A, Abronia cuyama sp. nov. UCMP 54560. B, Abronia graminea UTA 38831. C, Elgaria multicarinata TNHC 35666. D, Barisia leƲicollis MVZ 68783. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7695203" httpUri="https://zenodo.org/record/7695203/files/figure.png" pageId="4" pageNumber="756">4A</figureCitation>
|
||
). The osteoderm keels on and anterior to the frontal and the tall osteoderm keels or ‘sails’ on the posterior cranial osteoderms are autapomorphies of the new taxon (
|
||
<figureCitation box="[563,677,1426,1448]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="4" pageNumber="756">Figs 2H, I</figureCitation>
|
||
,
|
||
<figureCitation box="[691,723,1426,1447]" captionStart="Figure 4" captionStartId="7.[143,221,1035,1057]" captionTargetBox="[225,1344,197,994]" captionTargetId="figure-481@7.[223,1346,195,995]" captionTargetPageId="7" captionText="Figure 4. Comparison of Abronia cuyama sp. nov. and skulls of extant gerrhonotines, in anterior view. A, Abronia cuyama sp. nov. UCMP 54560. B, Abronia graminea UTA 38831. C, Elgaria multicarinata TNHC 35666. D, Barisia leƲicollis MVZ 68783. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7695203" httpUri="https://zenodo.org/record/7695203/files/figure.png" pageId="4" pageNumber="756">4A</figureCitation>
|
||
). We performed phylogenetic analyses (see below) to place the new taxon systematically with respect to extant species of
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[277,369,1518,1539]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="756" phylum="Chordata" rank="genus">
|
||
<emphasis box="[277,369,1518,1539]" italics="true" pageId="4" pageNumber="756">Abronia</emphasis>
|
||
</taxonomicName>
|
||
and other gerrhonotines.
|
||
</paragraph>
|
||
<paragraph blockId="4.[163,779,1577,1906]" pageId="4" pageNumber="756">
|
||
<emphasis box="[163,306,1578,1599]" italics="true" pageId="4" pageNumber="756">Description:</emphasis>
|
||
The specimen is comparable in size to skeletally mature extant gerrhonotines (
|
||
<figureCitation box="[614,675,1608,1630]" captionStart="Figure 4" captionStartId="7.[143,221,1035,1057]" captionTargetBox="[225,1344,197,994]" captionTargetId="figure-481@7.[223,1346,195,995]" captionTargetPageId="7" captionText="Figure 4. Comparison of Abronia cuyama sp. nov. and skulls of extant gerrhonotines, in anterior view. A, Abronia cuyama sp. nov. UCMP 54560. B, Abronia graminea UTA 38831. C, Elgaria multicarinata TNHC 35666. D, Barisia leƲicollis MVZ 68783. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7695203" httpUri="https://zenodo.org/record/7695203/files/figure.png" pageId="4" pageNumber="756">Fig. 4</figureCitation>
|
||
). Several morphological features previously used to determine skeletal maturity in anguimorphs were not preserved on the fossil (e.g. terminal fusions of the braincase, osteodermal crust on the parietal;
|
||
<bibRefCitation author="Bhullar B-AS" box="[562,717,1730,1752]" pageId="4" pageNumber="756" pagination="521 - 530" refId="ref9298" refString="Bhullar B-AS. 2012. A phylogenetic approach to ontogeny and heterochrony in the fossil record: cranial evolution and development in anguimorphan lizards (Reptilia: Squamata). Journal of Experimental Zoology. Part B. Molecular and DeVelopmental EVolution 318: 521 - 530." type="journal article" year="2012">Bhullar, 2012</bibRefCitation>
|
||
), but the robust osteoderms, well-developed osteodermal crust on the frontal and relatively high tooth count (
|
||
<bibRefCitation author="Ledesma DT & Scarpetta SG & Bell CJ" box="[171,404,1822,1844]" pageId="4" pageNumber="756" pagination="11602" refId="ref10854" refString="Ledesma DT, Scarpetta SG, Bell CJ. 2021. Variation in the skulls of Elgaria and Gerrhonotus (Anguidae, Gerrhonotinae) and implications for phylogenetics and fossil identification. PeerJ 9: e 11602." type="journal article" year="2021">
|
||
Ledesma
|
||
<emphasis box="[280,336,1822,1844]" italics="true" pageId="4" pageNumber="756">et al.</emphasis>
|
||
, 2021
|
||
</bibRefCitation>
|
||
) imply a mature individual.
|
||
</paragraph>
|
||
<paragraph blockId="4.[163,779,1577,1906]" lastBlockId="4.[827,1444,197,1905]" pageId="4" pageNumber="756">
|
||
Portions of the alveolar plate, nasal process and incisive process of the premaxilla are preserved (
|
||
<figureCitation box="[836,968,197,219]" captionStart="Figure 3" captionStartId="6.[163,241,1285,1307]" captionTargetBox="[165,1446,197,1244]" captionTargetId="figure-243@6.[163,1446,195,1245]" captionTargetPageId="6" captionText="Figure 3. Selected cranial elements of Abronia cuyama sp. nov. A, B, premaxilla in anterior and posterior view, respectively. C, D, left maxilla in lateral and dorsal view, respectively. E, left septomaxilla in dorsal view. F, right quadrate and separate pieces of right squamosal in lateral view. G, right vomer in lateral view. H, right jugal in lateral view. I, vomerine process of the left palatine in medial view. Scale bars: 1 mm. Abbreviations: a.pr, anterior process; d.o, dorsal ossification; f.pr, facial process; J.s, jugal spur; La, lacrimal; mx.lp, maxillary lappet; mx.pr, maxillary process; n.pr, nasal process; o.pr, orbital process; pa.pr, palatine process; pd.fl, posterodorsal flange; p.pr.Smx, posterior process of the septomaxilla; p.pr.Sq, posterior process of the squamosal; t.r, temporal ramus; v.k, ventral keel; v.l, ventral lamina; vn, vomeronasal region." figureDoi="http://doi.org/10.5281/zenodo.7695201" httpUri="https://zenodo.org/record/7695201/files/figure.png" pageId="4" pageNumber="756">Fig. 3A, B</figureCitation>
|
||
). There are three teeth and several replacement teeth. The nasal process is narrow relative to the width of the alveolar plate and tapers distally almost to a point, and the anterior surface of the process is relatively flat. The alveolar plate lacks discrete palatal processes. The ventral portion of the nasal process is broken, hence the presence of an anterior foramen or an ossified bridge between the nasal process and the alveolar plate cannot be determined. The nasals are present but were difficult to segment from the matrix, and osteoderms are fused to the dorsal surface of the nasals. Some morphological features of the nasals could be observed from the CT slices. Specifically, it was possible to ascertain that the nasals have an anteromedial process that articulates with the nasal process of the premaxilla (xy slice 239; Supporting Information,
|
||
<figureCitation box="[1111,1207,687,709]" captionStart="Figure 1" captionStartId="1.[145,224,1678,1700]" captionTargetBox="[284,1287,197,1636]" captionTargetId="figure-26@1.[283,1287,195,1638]" captionTargetPageId="1" captionText="Figure 1. A, geographical context. B, local Oligocene and Miocene sedimentary units. In A, distribution data for Abronia are from GBIF.org (2021). Distribution data were filtered manually to remove selected outliers using the study by Gutiérrez- Rodríguez et al. (2021) as a reference. Grey shading in A indicates elevation (i.e. mountainous areas). Geological data in B are based on the studies by Hoyt et al. (2018) and Jennings (2010). The location of the UCMP V-5847 fossiliferous site was obtained from the study by James (1963)." figureDoi="http://doi.org/10.5281/zenodo.7695197" httpUri="https://zenodo.org/record/7695197/files/figure.png" pageId="4" pageNumber="756">Fig. S1A</figureCitation>
|
||
) and that the nasals are broadly separated for most of their length, especially anteriorly (xy slice 272; Supporting Information,
|
||
<figureCitation captionStart="Figure 1" captionStartId="1.[145,224,1678,1700]" captionTargetBox="[284,1287,197,1636]" captionTargetId="figure-26@1.[283,1287,195,1638]" captionTargetPageId="1" captionText="Figure 1. A, geographical context. B, local Oligocene and Miocene sedimentary units. In A, distribution data for Abronia are from GBIF.org (2021). Distribution data were filtered manually to remove selected outliers using the study by Gutiérrez- Rodríguez et al. (2021) as a reference. Grey shading in A indicates elevation (i.e. mountainous areas). Geological data in B are based on the studies by Hoyt et al. (2018) and Jennings (2010). The location of the UCMP V-5847 fossiliferous site was obtained from the study by James (1963)." figureDoi="http://doi.org/10.5281/zenodo.7695197" httpUri="https://zenodo.org/record/7695197/files/figure.png" pageId="4" pageNumber="756">Fig. S1B</figureCitation>
|
||
) and near their anterior–posterior midpoint (xy slice 448 XY; Supporting Information,
|
||
<figureCitation box="[1262,1358,810,832]" captionStart="Figure 1" captionStartId="1.[145,224,1678,1700]" captionTargetBox="[284,1287,197,1636]" captionTargetId="figure-26@1.[283,1287,195,1638]" captionTargetPageId="1" captionText="Figure 1. A, geographical context. B, local Oligocene and Miocene sedimentary units. In A, distribution data for Abronia are from GBIF.org (2021). Distribution data were filtered manually to remove selected outliers using the study by Gutiérrez- Rodríguez et al. (2021) as a reference. Grey shading in A indicates elevation (i.e. mountainous areas). Geological data in B are based on the studies by Hoyt et al. (2018) and Jennings (2010). The location of the UCMP V-5847 fossiliferous site was obtained from the study by James (1963)." figureDoi="http://doi.org/10.5281/zenodo.7695197" httpUri="https://zenodo.org/record/7695197/files/figure.png" pageId="4" pageNumber="756">Fig. S1C</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph blockId="4.[827,1444,197,1905]" pageId="4" pageNumber="756">
|
||
Both septomaxillae are present, but only the left septomaxilla was segmented from the surrounding matrix (
|
||
<figureCitation box="[925,1011,902,924]" captionStart="Figure 3" captionStartId="6.[163,241,1285,1307]" captionTargetBox="[165,1446,197,1244]" captionTargetId="figure-243@6.[163,1446,195,1245]" captionTargetPageId="6" captionText="Figure 3. Selected cranial elements of Abronia cuyama sp. nov. A, B, premaxilla in anterior and posterior view, respectively. C, D, left maxilla in lateral and dorsal view, respectively. E, left septomaxilla in dorsal view. F, right quadrate and separate pieces of right squamosal in lateral view. G, right vomer in lateral view. H, right jugal in lateral view. I, vomerine process of the left palatine in medial view. Scale bars: 1 mm. Abbreviations: a.pr, anterior process; d.o, dorsal ossification; f.pr, facial process; J.s, jugal spur; La, lacrimal; mx.lp, maxillary lappet; mx.pr, maxillary process; n.pr, nasal process; o.pr, orbital process; pa.pr, palatine process; pd.fl, posterodorsal flange; p.pr.Smx, posterior process of the septomaxilla; p.pr.Sq, posterior process of the squamosal; t.r, temporal ramus; v.k, ventral keel; v.l, ventral lamina; vn, vomeronasal region." figureDoi="http://doi.org/10.5281/zenodo.7695201" httpUri="https://zenodo.org/record/7695201/files/figure.png" pageId="4" pageNumber="756">Fig. 3E</figureCitation>
|
||
). The posterior process is long. The presence of an anterolateral process could not be determined. The maxillae preserve the facial, palatine, premaxillary and orbital processes (
|
||
<figureCitation box="[1256,1348,994,1016]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="4" pageNumber="756">Figs 2D</figureCitation>
|
||
,
|
||
<figureCitation box="[1362,1427,994,1016]" captionStart="Figure 3" captionStartId="6.[163,241,1285,1307]" captionTargetBox="[165,1446,197,1244]" captionTargetId="figure-243@6.[163,1446,195,1245]" captionTargetPageId="6" captionText="Figure 3. Selected cranial elements of Abronia cuyama sp. nov. A, B, premaxilla in anterior and posterior view, respectively. C, D, left maxilla in lateral and dorsal view, respectively. E, left septomaxilla in dorsal view. F, right quadrate and separate pieces of right squamosal in lateral view. G, right vomer in lateral view. H, right jugal in lateral view. I, vomerine process of the left palatine in medial view. Scale bars: 1 mm. Abbreviations: a.pr, anterior process; d.o, dorsal ossification; f.pr, facial process; J.s, jugal spur; La, lacrimal; mx.lp, maxillary lappet; mx.pr, maxillary process; n.pr, nasal process; o.pr, orbital process; pa.pr, palatine process; pd.fl, posterodorsal flange; p.pr.Smx, posterior process of the septomaxilla; p.pr.Sq, posterior process of the squamosal; t.r, temporal ramus; v.k, ventral keel; v.l, ventral lamina; vn, vomeronasal region." figureDoi="http://doi.org/10.5281/zenodo.7695201" httpUri="https://zenodo.org/record/7695201/files/figure.png" pageId="4" pageNumber="756">3C, D</figureCitation>
|
||
). There are six or seven nutrient foramina on the labial surface of both maxillae. There are 18 teeth on the left maxilla, filling all available positions, and on the right maxilla there are 16 teeth and 20 tooth positions. The maxillary lappet projects anterodorsally from the medial surface of the premaxillary process. There is a single foramen (the anterior inferior alveolar foramen) near the anterior edge of the facial process where it meets the premaxillary process. The facial process is inflected medially but does not contact the frontal. The anteroposterior dimension of the facial process is narrow. The palatine process has a slightly rounded subtriangular medial projection. The orbital process is long relative to the facial and premaxillary processes. The posteriormost portion of the left orbital process is broken. Rugose texturing is present on the lateral surface of the maxilla, particularly on the facial process, and some osteoderms are fused to the facial process. The lacrimal is long, extending for nearly half the length of the orbital process of the maxilla, and has rugose sculpturing on its lateral surface. The lacrimal extends both medially and dorsally to enclose, in part, the lacrimal foramen.
|
||
</paragraph>
|
||
<paragraph blockId="4.[827,1444,197,1905]" lastBlockId="6.[163,779,1521,1911]" lastPageId="6" lastPageNumber="758" pageId="4" pageNumber="756">
|
||
The right jugal is present but is missing the posterior portion of the orbital process and dorsal portion of the temporal ramus (
|
||
<figureCitation box="[1185,1269,1791,1813]" captionStart="Figure 3" captionStartId="6.[163,241,1285,1307]" captionTargetBox="[165,1446,197,1244]" captionTargetId="figure-243@6.[163,1446,195,1245]" captionTargetPageId="6" captionText="Figure 3. Selected cranial elements of Abronia cuyama sp. nov. A, B, premaxilla in anterior and posterior view, respectively. C, D, left maxilla in lateral and dorsal view, respectively. E, left septomaxilla in dorsal view. F, right quadrate and separate pieces of right squamosal in lateral view. G, right vomer in lateral view. H, right jugal in lateral view. I, vomerine process of the left palatine in medial view. Scale bars: 1 mm. Abbreviations: a.pr, anterior process; d.o, dorsal ossification; f.pr, facial process; J.s, jugal spur; La, lacrimal; mx.lp, maxillary lappet; mx.pr, maxillary process; n.pr, nasal process; o.pr, orbital process; pa.pr, palatine process; pd.fl, posterodorsal flange; p.pr.Smx, posterior process of the septomaxilla; p.pr.Sq, posterior process of the squamosal; t.r, temporal ramus; v.k, ventral keel; v.l, ventral lamina; vn, vomeronasal region." figureDoi="http://doi.org/10.5281/zenodo.7695201" httpUri="https://zenodo.org/record/7695201/files/figure.png" pageId="4" pageNumber="756">Fig. 3H</figureCitation>
|
||
). The temporal ramus has a broad anteroposterior dimension, but the orbital process is more gracile. The orbital process has a ventral lamina that overhangs the dorsal margin of the orbital process of the maxilla. The jugal spur (quadratojugal process) is large and has a pointed posterior projection. The palpebral is triangular and externally visible.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.7695199" ID-Zenodo-Dep="7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="5" pageNumber="757" startId="5.[145,225,1678,1700]" targetBox="[243,1328,197,1630]" targetPageId="5">
|
||
<paragraph blockId="5.[145,1425,1678,1906]" pageId="5" pageNumber="757">
|
||
<emphasis bold="true" box="[145,251,1678,1700]" pageId="5" pageNumber="757">Figure 2.</emphasis>
|
||
Holotype of
|
||
<taxonomicName authority="Scarpetta & Ledesma, 2023" authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[391,565,1679,1701]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="757" phylum="Chordata" rank="species" species="cuyama" status="sp. nov.">
|
||
<emphasis box="[391,565,1679,1701]" italics="true" pageId="5" pageNumber="757">Abronia cuyama</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[573,650,1679,1701]" pageId="5" pageNumber="757" rank="species">sp. nov.</taxonomicNameLabel>
|
||
<emphasis box="[656,799,1679,1701]" italics="true" pageId="5" pageNumber="757">UCMP 54560</emphasis>
|
||
. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest.
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.7695201" ID-Zenodo-Dep="7695201" httpUri="https://zenodo.org/record/7695201/files/figure.png" pageId="6" pageNumber="758" startId="6.[163,241,1285,1307]" targetBox="[165,1446,197,1244]" targetPageId="6">
|
||
<paragraph blockId="6.[163,1444,1285,1483]" pageId="6" pageNumber="758">
|
||
<emphasis bold="true" box="[163,266,1285,1307]" pageId="6" pageNumber="758">Figure 3.</emphasis>
|
||
Selected cranial elements of
|
||
<taxonomicName authority="Scarpetta & Ledesma, 2023" authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[572,743,1285,1307]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="758" phylum="Chordata" rank="species" species="cuyama" status="sp. nov.">
|
||
<emphasis box="[572,743,1285,1307]" italics="true" pageId="6" pageNumber="758">Abronia cuyama</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[748,823,1285,1307]" pageId="6" pageNumber="758" rank="species">sp. nov.</taxonomicNameLabel>
|
||
A, B, premaxilla in anterior and posterior view, respectively. C, D, left maxilla in lateral and dorsal view, respectively. E, left septomaxilla in dorsal view. F, right quadrate and separate pieces of right squamosal in lateral view. G, right vomer in lateral view. H, right jugal in lateral view. I, vomerine process of the left palatine in medial view. Scale bars: 1 mm. Abbreviations: a.pr, anterior process; d.o, dorsal ossification; f.pr, facial process; J.s, jugal spur; La, lacrimal; mx.lp, maxillary lappet; mx.pr, maxillary process; n.pr, nasal process; o.pr, orbital process; pa.pr, palatine process; pd.fl, posterodorsal flange; p.pr.Smx, posterior process of the septomaxilla; p.pr.Sq, posterior process of the squamosal; t.r, temporal ramus; v.k, ventral keel; v.l, ventral lamina; vn, vomeronasal region.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="6.[163,779,1521,1911]" lastBlockId="6.[827,1443,1521,1911]" pageId="6" pageNumber="758">
|
||
Both prefrontals are present but were difficult to separate from the matrix. The frontal process is long and projects posterodorsally. The ventral process is short and lacks an anteroventral projection (xy slices 740–750). The prefrontal has a large articulation facet for the facial process of the maxilla. The frontal is complete but broken into an anterior and a posterior piece that are slightly separated from each other (
|
||
<figureCitation box="[171,285,1889,1911]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="6" pageNumber="758">Fig. 2E, F</figureCitation>
|
||
). The frontal is azygous and wide relative to its length. An osteodermal crust covers the dorsal exposure of the frontal, and some osteoderms are fused to the frontal. The anterior process of the frontal is triradiate. The interorbital region is constricted. The facet for the prefrontal on the anterolateral surface of the frontal has small, anterolaterally facing processes at its posterior margin.
|
||
</paragraph>
|
||
<paragraph blockId="6.[827,1443,1521,1911]" lastBlockId="7.[145,762,1154,1912]" lastPageId="7" lastPageNumber="759" pageId="6" pageNumber="758">
|
||
Both vomers are present, and the right vomer was segmented (
|
||
<figureCitation box="[973,1061,1766,1788]" captionStart="Figure 3" captionStartId="6.[163,241,1285,1307]" captionTargetBox="[165,1446,197,1244]" captionTargetId="figure-243@6.[163,1446,195,1245]" captionTargetPageId="6" captionText="Figure 3. Selected cranial elements of Abronia cuyama sp. nov. A, B, premaxilla in anterior and posterior view, respectively. C, D, left maxilla in lateral and dorsal view, respectively. E, left septomaxilla in dorsal view. F, right quadrate and separate pieces of right squamosal in lateral view. G, right vomer in lateral view. H, right jugal in lateral view. I, vomerine process of the left palatine in medial view. Scale bars: 1 mm. Abbreviations: a.pr, anterior process; d.o, dorsal ossification; f.pr, facial process; J.s, jugal spur; La, lacrimal; mx.lp, maxillary lappet; mx.pr, maxillary process; n.pr, nasal process; o.pr, orbital process; pa.pr, palatine process; pd.fl, posterodorsal flange; p.pr.Smx, posterior process of the septomaxilla; p.pr.Sq, posterior process of the squamosal; t.r, temporal ramus; v.k, ventral keel; v.l, ventral lamina; vn, vomeronasal region." figureDoi="http://doi.org/10.5281/zenodo.7695201" httpUri="https://zenodo.org/record/7695201/files/figure.png" pageId="6" pageNumber="758">Fig. 3G</figureCitation>
|
||
). There is a dorsally extending posterolateral flange. The anterior (vomeronasal) portion of the bone extends to a lower ventral level than the rest of the element. The foramen for the medial palatine nerve penetrates the right vomer posteriorly and exits anteriorly (xy slices 528–573). The palatines were difficult to segment from the matrix, especially posteriorly. Only the vomerine process was segmented (
|
||
<figureCitation box="[154,232,1246,1268]" captionStart="Figure 3" captionStartId="6.[163,241,1285,1307]" captionTargetBox="[165,1446,197,1244]" captionTargetId="figure-243@6.[163,1446,195,1245]" captionTargetPageId="6" captionText="Figure 3. Selected cranial elements of Abronia cuyama sp. nov. A, B, premaxilla in anterior and posterior view, respectively. C, D, left maxilla in lateral and dorsal view, respectively. E, left septomaxilla in dorsal view. F, right quadrate and separate pieces of right squamosal in lateral view. G, right vomer in lateral view. H, right jugal in lateral view. I, vomerine process of the left palatine in medial view. Scale bars: 1 mm. Abbreviations: a.pr, anterior process; d.o, dorsal ossification; f.pr, facial process; J.s, jugal spur; La, lacrimal; mx.lp, maxillary lappet; mx.pr, maxillary process; n.pr, nasal process; o.pr, orbital process; pa.pr, palatine process; pd.fl, posterodorsal flange; p.pr.Smx, posterior process of the septomaxilla; p.pr.Sq, posterior process of the squamosal; t.r, temporal ramus; v.k, ventral keel; v.l, ventral lamina; vn, vomeronasal region." figureDoi="http://doi.org/10.5281/zenodo.7695201" httpUri="https://zenodo.org/record/7695201/files/figure.png" pageId="7" pageNumber="759">Fig. 3I</figureCitation>
|
||
), but from the CT slices it was evident that there are no palatine teeth. The vomerine process lacks a posteroventral ridge demarcating the facet for the vomer. There is no anterior flange dorsal to the choana.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.7695203" ID-Zenodo-Dep="7695203" httpUri="https://zenodo.org/record/7695203/files/figure.png" pageId="7" pageNumber="759" startId="7.[143,221,1035,1057]" targetBox="[225,1344,197,994]" targetPageId="7">
|
||
<paragraph blockId="7.[143,1425,1035,1116]" pageId="7" pageNumber="759">
|
||
<emphasis bold="true" box="[143,247,1035,1057]" pageId="7" pageNumber="759">Figure 4.</emphasis>
|
||
Comparison of
|
||
<taxonomicName authority="Scarpetta & Ledesma, 2023" authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[418,590,1035,1057]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="759" phylum="Chordata" rank="species" species="cuyama" status="sp. nov.">
|
||
<emphasis box="[418,590,1035,1057]" italics="true" pageId="7" pageNumber="759">Abronia cuyama</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[597,673,1035,1057]" pageId="7" pageNumber="759" rank="species">sp. nov.</taxonomicNameLabel>
|
||
and skulls of extant gerrhonotines, in anterior view. A,
|
||
<taxonomicName authority="Scarpetta & Ledesma, 2023" authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[1253,1424,1035,1057]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="759" phylum="Chordata" rank="species" species="cuyama" status="sp. nov.">
|
||
<emphasis box="[1253,1424,1035,1057]" italics="true" pageId="7" pageNumber="759">Abronia cuyama</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel box="[145,222,1064,1086]" pageId="7" pageNumber="759" rank="species">sp. nov.</taxonomicNameLabel>
|
||
<emphasis box="[227,369,1064,1086]" italics="true" pageId="7" pageNumber="759">UCMP 54560</emphasis>
|
||
. B,
|
||
<emphasis box="[409,728,1064,1086]" italics="true" pageId="7" pageNumber="759">
|
||
<taxonomicName authority="UTA" authorityName="UTA" box="[409,658,1064,1086]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="759" phylum="Chordata" rank="species" species="graminea">Abronia graminea UTA</taxonomicName>
|
||
38831
|
||
</emphasis>
|
||
. C,
|
||
<taxonomicName authority="TNHC" authorityName="TNHC" box="[769,1078,1064,1086]" class="Reptilia" family="Anguidae" genus="Elgaria" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="759" phylum="Chordata" rank="species" species="multicarinata">
|
||
<emphasis box="[769,1003,1064,1086]" italics="true" pageId="7" pageNumber="759">Elgaria multicarinata</emphasis>
|
||
TNHC
|
||
</taxonomicName>
|
||
35666. D,
|
||
<taxonomicName authority="MVZ" authorityName="MVZ" box="[1189,1425,1064,1086]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="759" phylum="Chordata" rank="species" species="levicollis">
|
||
<emphasis box="[1189,1366,1064,1086]" italics="true" pageId="7" pageNumber="759">Barisia leƲicollis</emphasis>
|
||
MVZ
|
||
</taxonomicName>
|
||
68783. Scale bars: 1 mm.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="7.[145,762,1154,1912]" lastBlockId="7.[809,1426,1154,1912]" pageId="7" pageNumber="759">
|
||
The dentaries preserve all but the posterior portion of the bone (
|
||
<figureCitation box="[304,429,1399,1421]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="7" pageNumber="759">Fig. 2B, D</figureCitation>
|
||
). The Meckelian groove is open and is directed ventromedially and ventrally in the anterior and posterior portions of the dentary, respectively. Neither dentary is preserved well enough to ascertain the presence of a surangular process or a posterior groove ventral to the parapet. The free margin of the intramandibular septum is incomplete but present on both sides (for the right dentary, see xy slice 1044; Supporting Information,
|
||
<figureCitation box="[593,673,1644,1666]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="7" pageNumber="759">Fig. S2</figureCitation>
|
||
). There are 22 or 23 tooth positions and 17 teeth on the right dentary and at least ten teeth on the left dentary. Dentition is pleurodont. Crowns on the mesial teeth are unicuspid and sharp, and the crowns on distal teeth are near bicuspid. Teeth are recurved throughout the tooth row of both the dentary and maxilla except for the distalmost teeth. Some of the crowns are missing from the mesial teeth of the right dentary and left maxilla. The mesial teeth and some of the mid-tooth row teeth of the right maxilla are particularly long, recurved and sharp.
|
||
</paragraph>
|
||
<paragraph blockId="7.[809,1426,1154,1912]" pageId="7" pageNumber="759">
|
||
The distal portion of the right squamosal is preserved in the separate mudstone block (
|
||
<figureCitation box="[1305,1419,1276,1298]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="7" pageNumber="759">Figs 2G–I</figureCitation>
|
||
,
|
||
<figureCitation box="[809,840,1307,1328]" captionStart="Figure 3" captionStartId="6.[163,241,1285,1307]" captionTargetBox="[165,1446,197,1244]" captionTargetId="figure-243@6.[163,1446,195,1245]" captionTargetPageId="6" captionText="Figure 3. Selected cranial elements of Abronia cuyama sp. nov. A, B, premaxilla in anterior and posterior view, respectively. C, D, left maxilla in lateral and dorsal view, respectively. E, left septomaxilla in dorsal view. F, right quadrate and separate pieces of right squamosal in lateral view. G, right vomer in lateral view. H, right jugal in lateral view. I, vomerine process of the left palatine in medial view. Scale bars: 1 mm. Abbreviations: a.pr, anterior process; d.o, dorsal ossification; f.pr, facial process; J.s, jugal spur; La, lacrimal; mx.lp, maxillary lappet; mx.pr, maxillary process; n.pr, nasal process; o.pr, orbital process; pa.pr, palatine process; pd.fl, posterodorsal flange; p.pr.Smx, posterior process of the septomaxilla; p.pr.Sq, posterior process of the squamosal; t.r, temporal ramus; v.k, ventral keel; v.l, ventral lamina; vn, vomeronasal region." figureDoi="http://doi.org/10.5281/zenodo.7695201" httpUri="https://zenodo.org/record/7695201/files/figure.png" pageId="7" pageNumber="759">3F</figureCitation>
|
||
). The ventral process curves anterolaterally to articulate with the right quadrate. The element is uniform in width. The right quadrate also is preserved in the block. The quadrate has a narrow mediolateral dimension, with roughly parallel lateral and medial margins. The cephalic condyle is well developed. The mandibular condyle is shaped like a hyperbolic paraboloid and is slightly narrower than the rest of the quadrate. The anteromedial surface of the quadrate is concave, especially dorsally. The pterygoid lamina is shallow, and there is a slightly protruding flange of bone dorsal to it. The column and the tympanic crest are both well developed. The dorsal and ventral portions of the conch are roughly the same width.
|
||
</paragraph>
|
||
<paragraph blockId="7.[809,1426,1154,1912]" lastBlockId="8.[163,780,197,1047]" lastPageId="8" lastPageNumber="760" pageId="7" pageNumber="759">
|
||
The osteoderms are robust and have heavy vermiculate sculpturing (
|
||
<figureCitation box="[1103,1220,1767,1789]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="7" pageNumber="759">Figs 2A–F</figureCitation>
|
||
,
|
||
<figureCitation box="[1233,1265,1767,1788]" captionStart="Figure 4" captionStartId="7.[143,221,1035,1057]" captionTargetBox="[225,1344,197,994]" captionTargetId="figure-481@7.[223,1346,195,995]" captionTargetPageId="7" captionText="Figure 4. Comparison of Abronia cuyama sp. nov. and skulls of extant gerrhonotines, in anterior view. A, Abronia cuyama sp. nov. UCMP 54560. B, Abronia graminea UTA 38831. C, Elgaria multicarinata TNHC 35666. D, Barisia leƲicollis MVZ 68783. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7695203" httpUri="https://zenodo.org/record/7695203/files/figure.png" pageId="7" pageNumber="759">4A</figureCitation>
|
||
). Osteoderms become more rectangular posteriorly. Many osteoderms have midline keels, including osteoderms anterior to the frontal and the osteodermal crust of the frontal itself. An articulated series of osteoderms (potential temporal osteoderms) surround the quadrate dorsally, laterally and ventrolaterally (
|
||
<figureCitation box="[402,510,228,250]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="8" pageNumber="760">Fig. 2G–I</figureCitation>
|
||
). The osteoderms must have slid ventrally during fossilization, because osteoderms could not occur directly ventral to the quadrate in a live lizard owing to articulation between the quadrate and the mandible. The posterior osteoderms above the quadrate have keels or ‘sails’ that are more than twice as tall as the osteoderm itself (
|
||
<figureCitation box="[171,280,442,464]" captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="8" pageNumber="760">Fig. 2H, I</figureCitation>
|
||
). The prefrontal, frontonasal and posterior internasal osteoderms are present, and the prefrontals and posterior internasals are in broad contact (
|
||
<figureCitation captionStart="Figure 2" captionStartId="5.[145,225,1678,1700]" captionTargetBox="[243,1328,197,1630]" captionTargetId="figure-26@5.[241,1328,195,1639]" captionTargetPageId="5" captionText="Figure 2. Holotype of Abronia cuyama sp. nov. UCMP 54560. Images on the left are of the physical specimen and those on the right are digital renderings; B, D and F are volume renderings, whereas H and I are surface renderings. A, B, skull in right lateral view. C, D, skull in left lateral view. E, F, skull in dorsal view. G, quadrate, squamosal and osteoderms from mudstone block in posterior view. H, I, segmented quadrate, squamosal and osteoderms dorsal to the quadrate in posterior and anterior view, respectively. Scale bars: 1 mm. Abbreviations: am.s, anteromedial surface; co, conch; De, dentary; fn, frontonasal; f.pr, facial process; fp, frontoparietal; Fr, frontal; Ju, jugal; k, keel; La, lacrimal; m.so, median supraocular; Me.gr, Meckelian groove; Mx, maxilla; or.pr, orbital process; p.in, posterior internasal; Pp, palpebral; prf, prefrontal; pt.lm, pterygoid lamina; Px, premaxilla; px.pr, premaxillary process; Qu, quadrate; Smx, septomaxilla; Sq, squamosal; ty.cr, tympanic crest." figureDoi="http://doi.org/10.5281/zenodo.7695199" httpUri="https://zenodo.org/record/7695199/files/figure.png" pageId="8" pageNumber="760">Fig. 2E, F</figureCitation>
|
||
). The left prefrontal osteoderm appears to have slipped ventrally from its natural position. Supranasal and anterior internasal osteoderms are not preserved. There is a frontoparietal osteoderm on the left side of the frontal; the osteoderm might have fallen off on the right side. No anteroventral or sublabial osteoderms were preserved.
|
||
</paragraph>
|
||
<paragraph blockId="8.[163,780,197,1047]" pageId="8" pageNumber="760">One of the vials of associated material contains two unidentified bones, which might be weathered osteoderms. The other vial contains two small pieces of mudstone matrix, each with several visible and apparently articulated osteoderms. One of the matrix pieces also contains a tubular bone that is probably an epipteryogid. The osteoderms have the same heavy, vermiculate sculpturing as do those on the skull and in the large mudstone block, and several osteoderms preserve distinct keels.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="9" lastPageNumber="761" pageId="8" pageNumber="760" type="discussion">
|
||
<paragraph blockId="8.[163,780,1088,1907]" pageId="8" pageNumber="760">
|
||
<emphasis box="[163,330,1088,1109]" italics="true" pageId="8" pageNumber="760">Comparisons:</emphasis>
|
||
Comparisons are largely based on the specimens examined by
|
||
<bibRefCitation author="Scarpetta SG & Ledesma DT & Bell CJ" box="[503,772,1118,1140]" pageId="8" pageNumber="760" pagination="1 - 58" refId="ref11478" refString="Scarpetta SG, Ledesma DT, Bell CJ. 2021. A new extinct species of alligator lizard (Squamata: Elgaria) and an expanded perspective on the osteology and phylogeny of Gerrhonotinae. BMC Ecology and EVolution 21: 1 - 58." type="journal article" year="2021">
|
||
Scarpetta
|
||
<emphasis box="[628,688,1118,1140]" italics="true" pageId="8" pageNumber="760">et al.</emphasis>
|
||
(2021)
|
||
</bibRefCitation>
|
||
. If no specimen number is listed, the comparison accommodates all specimens of a given taxon that we examined for that dataset.
|
||
</paragraph>
|
||
<paragraph blockId="8.[163,780,1088,1907]" lastBlockId="8.[827,1444,197,1905]" pageId="8" pageNumber="760">
|
||
<taxonomicName authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[187,382,1241,1263]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="cuyama">
|
||
<emphasis box="[187,382,1241,1263]" italics="true" pageId="8" pageNumber="760">Abronia cuyama</emphasis>
|
||
</taxonomicName>
|
||
has the wide frontal and heavily sculptured vermiculate-textured osteoderms that are characteristic of
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[436,537,1302,1323]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[436,537,1302,1323]" italics="true" pageId="8" pageNumber="760">Abronia</emphasis>
|
||
</taxonomicName>
|
||
, excluding, among examined specimens,
|
||
<taxonomicName authority="Cope, 1878" authorityName="Cope" authorityYear="1878" box="[419,775,1333,1354]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="monticola">
|
||
<emphasis box="[419,636,1333,1354]" italics="true" pageId="8" pageNumber="760">Abronia monticola</emphasis>
|
||
Cope, 1878
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authority="Bocourt, 1872" authorityName="Bocourt" authorityYear="1872" box="[163,534,1364,1385]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="moreletti">
|
||
<emphasis box="[163,367,1364,1385]" italics="true" pageId="8" pageNumber="760">Abronia moreletti</emphasis>
|
||
Bocourt, 1872
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authority="Boulenger, 1913" authorityName="Boulenger" authorityYear="1913" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="gadovii">
|
||
<emphasis box="[592,778,1364,1385]" italics="true" pageId="8" pageNumber="760">Abronia gadoƲii</emphasis>
|
||
Boulenger, 1913
|
||
</taxonomicName>
|
||
(i.e. species previously placed in
|
||
<taxonomicName box="[163,272,1426,1447]" class="Reptilia" family="Anguidae" genus="Mesaspis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[163,272,1426,1447]" italics="true" pageId="8" pageNumber="760">Mesaspis</emphasis>
|
||
</taxonomicName>
|
||
). The anterodorsal osteoderms of
|
||
<taxonomicName authority="Wiegmann, 1828" authorityName="Wiegmann" authorityYear="1828" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="imbricata">
|
||
<emphasis italics="true" pageId="8" pageNumber="760">Barisia imbricata</emphasis>
|
||
Wiegmann, 1828
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authority="Smith, 1942" authorityName="Smith" authorityYear="1942" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ciliaris">
|
||
<emphasis box="[525,696,1456,1477]" italics="true" pageId="8" pageNumber="760">Barisia ciliaris</emphasis>
|
||
Smith, 1942
|
||
</taxonomicName>
|
||
are also heavily sculptured. The facial process of the maxilla is narrow in
|
||
<taxonomicName authority="Brodie & Savage, 1993" authorityName="Brodie & Savage" authorityYear="1993" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="campbelli">
|
||
<emphasis box="[481,694,1517,1538]" italics="true" pageId="8" pageNumber="760">Abronia campbelli</emphasis>
|
||
Brodie & Savage, 1993
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authority="Smith & Alvarez" authorityName="Smith & Alvarez" box="[354,778,1548,1570]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="lythrochila">
|
||
<emphasis box="[354,579,1548,1569]" italics="true" pageId="8" pageNumber="760">Abronia lythrochila</emphasis>
|
||
Smith & Alvarez
|
||
</taxonomicName>
|
||
del Toro, 1963,
|
||
<taxonomicName authorityName="Bocourt" authorityYear="1872" box="[341,478,1578,1600]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="moreletti">
|
||
<emphasis box="[341,478,1578,1600]" italics="true" pageId="8" pageNumber="760">A. moreletti</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Cope" authorityYear="1878" box="[492,639,1578,1600]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="monticola">
|
||
<emphasis box="[492,639,1578,1600]" italics="true" pageId="8" pageNumber="760">A. monticola</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Boulenger" authorityYear="1913" box="[654,772,1578,1600]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="gadovii">
|
||
<emphasis box="[654,772,1578,1600]" italics="true" pageId="8" pageNumber="760">A. gadoƲii</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authority="SRSU" authorityName="SRSU" box="[163,355,1609,1630]" class="Reptilia" family="Anguidae" genus="Gerrhonotus" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="parvus">
|
||
<emphasis box="[163,276,1609,1630]" italics="true" pageId="8" pageNumber="760">G. parƲus</emphasis>
|
||
SRSU
|
||
</taxonomicName>
|
||
5538 and some observed specimens of
|
||
<taxonomicName authority="Stebbins, 1958" authorityName="Stebbins" authorityYear="1958" box="[193,627,1639,1662]" class="Reptilia" family="Anguidae" genus="Elgaria" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="panamintina">
|
||
<emphasis box="[193,445,1640,1661]" italics="true" pageId="8" pageNumber="760">Elgaria panamintina</emphasis>
|
||
Stebbins, 1958
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authority="Grismer & Hollingsworth, 2001" authorityName="Grismer & Hollingsworth" authorityYear="2001" class="Reptilia" family="Anguidae" genus="Elgaria" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="velazquezi">
|
||
<emphasis italics="true" pageId="8" pageNumber="760">Elgaria Ʋelazquezi</emphasis>
|
||
Grismer & Hollingsworth, 2001
|
||
</taxonomicName>
|
||
(
|
||
<collectionCode box="[718,778,1670,1691]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34777" lsid="urn:lsid:biocol.org:col:34777" name="Museum of Vertebrate Zoology, University of California Berkeley" pageId="8" pageNumber="760" type="Museum">MVZ</collectionCode>
|
||
191076 and
|
||
<collectionCode box="[314,416,1701,1722]" country="USA" httpUri="http://grbio.org/cool/4ksk-xsw5" name="San Diego Natural History Museum" pageId="8" pageNumber="760">SDNHM</collectionCode>
|
||
68678, respectively). Several species of
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[279,371,1732,1753]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[279,371,1732,1753]" italics="true" pageId="8" pageNumber="760">Abronia</emphasis>
|
||
</taxonomicName>
|
||
(e.g.
|
||
<taxonomicName authority="Cope, 1864" authorityName="Cope" authorityYear="1864" box="[432,775,1731,1753]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="graminea">
|
||
<emphasis box="[432,640,1732,1753]" italics="true" pageId="8" pageNumber="760">Abronia graminea</emphasis>
|
||
Cope, 1864
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Campbell" authorityYear="1984" box="[163,287,1762,1784]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ornelasi">
|
||
<emphasis box="[163,287,1762,1784]" italics="true" pageId="8" pageNumber="760">A. ornelasi</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Boulenger" authorityYear="1913" box="[345,462,1762,1784]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="gadovii">
|
||
<emphasis box="[345,462,1762,1784]" italics="true" pageId="8" pageNumber="760">A. gadoƲii</emphasis>
|
||
</taxonomicName>
|
||
),
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[484,569,1763,1784]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[484,569,1763,1784]" italics="true" pageId="8" pageNumber="760">Barisia</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<taxonomicName authority="Stejneger, 1890" authorityName="Stejneger" authorityYear="1890" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="levicollis">
|
||
<emphasis box="[585,779,1762,1784]" italics="true" pageId="8" pageNumber="760">Barisia leƲicollis</emphasis>
|
||
Stejneger, 1890
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Smith" authorityYear="1942" box="[371,493,1793,1815]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ciliaris">
|
||
<emphasis box="[371,493,1793,1815]" italics="true" pageId="8" pageNumber="760">B. ciliaris</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Wiegmann" authorityYear="1828" box="[559,713,1793,1815]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="imbricata">
|
||
<emphasis box="[559,713,1793,1815]" italics="true" pageId="8" pageNumber="760">B. imbricata</emphasis>
|
||
</taxonomicName>
|
||
) and
|
||
<emphasis box="[163,258,1824,1845]" italics="true" pageId="8" pageNumber="760">
|
||
D.
|
||
<taxonomicName authorityName="McCoy" authorityYear="1970" box="[198,258,1824,1845]" class="Magnoliopsida" family="Cannabaceae" genus="Desertum" kingdom="Plantae" order="Rosales" pageId="8" pageNumber="760" phylum="Tracheophyta" rank="species" species="lugoi">lugoi</taxonomicName>
|
||
</emphasis>
|
||
have nasals that are separated near their longitudinal midpoint. The quadrate conch is largely uniform in width in
|
||
<taxonomicName authorityName="Brodie & Savage" authorityYear="1993" box="[425,581,1885,1906]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="campbelli">
|
||
<emphasis box="[425,581,1885,1906]" italics="true" pageId="8" pageNumber="760">A. campbelli</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<emphasis box="[600,954,197,1906]" italics="true" pageId="8" pageNumber="760">
|
||
<taxonomicName box="[600,773,1885,1906]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="lythrochila">A. lythrochila</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Boulenger" authorityYear="1913" box="[827,954,197,218]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="gadovii">A. gadoƲii</taxonomicName>
|
||
</emphasis>
|
||
and several specimens of
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1291,1382,198,219]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1291,1382,198,219]" italics="true" pageId="8" pageNumber="760">Barisia</emphasis>
|
||
</taxonomicName>
|
||
(e.g.
|
||
<taxonomicName authority="FMNH" authorityName="FMNH" box="[827,1040,228,249]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ciliaris">
|
||
<emphasis box="[827,947,228,249]" italics="true" pageId="8" pageNumber="760">B. ciliaris</emphasis>
|
||
FMNH
|
||
</taxonomicName>
|
||
30707). The vomeronasal region of the vomer is ventrally displaced in
|
||
<taxonomicName authorityName="Brodie & Savage" authorityYear="1993" box="[1287,1435,258,279]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="campbelli">
|
||
<emphasis box="[1287,1435,258,279]" italics="true" pageId="8" pageNumber="760">A. campbelli</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName box="[827,981,289,310]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="lythrochila">
|
||
<emphasis box="[827,981,289,310]" italics="true" pageId="8" pageNumber="760">A. lythrochila</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Cope" authorityYear="1864" box="[993,1133,289,310]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="graminea">
|
||
<emphasis box="[993,1133,289,310]" italics="true" pageId="8" pageNumber="760">A. graminea</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Campbell" authorityYear="1984" box="[1145,1266,289,310]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ornelasi">
|
||
<emphasis box="[1145,1266,289,310]" italics="true" pageId="8" pageNumber="760">A. ornelasi</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Boulenger" authorityYear="1913" box="[1322,1437,289,310]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="gadovii">
|
||
<emphasis box="[1322,1437,289,310]" italics="true" pageId="8" pageNumber="760">A. gadoƲii</emphasis>
|
||
</taxonomicName>
|
||
. The vomerine process of the palatine lacks a ventral ridge in most
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1012,1113,350,371]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1012,1113,350,371]" italics="true" pageId="8" pageNumber="760">Abronia</emphasis>
|
||
</taxonomicName>
|
||
, but the ridge is present in
|
||
<taxonomicName authority="Bogert & Porter, 1967" authorityName="Bogert & Porter" authorityYear="1967" box="[865,1383,381,403]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="mixteca">
|
||
<emphasis box="[865,1078,381,402]" italics="true" pageId="8" pageNumber="760">Abronia mixteca</emphasis>
|
||
Bogert & Porter, 1967
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Boulenger" authorityYear="1913" box="[827,949,411,433]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="gadovii">
|
||
<emphasis box="[827,949,411,433]" italics="true" pageId="8" pageNumber="760">A. gadoƲii</emphasis>
|
||
</taxonomicName>
|
||
. Species previously referred to
|
||
<taxonomicName box="[1334,1442,412,433]" class="Reptilia" family="Anguidae" genus="Mesaspis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1334,1442,412,433]" italics="true" pageId="8" pageNumber="760">Mesaspis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authority="Wiegmann, 1828" authorityName="Wiegmann" authorityYear="1828" box="[877,1267,442,464]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="taeniata">
|
||
<emphasis box="[877,1070,442,464]" italics="true" pageId="8" pageNumber="760">Abronia taeniata</emphasis>
|
||
Wiegmann, 1828
|
||
</taxonomicName>
|
||
(
|
||
<collectionCode box="[1281,1359,442,463]" country="USA" httpUri="http://grbio.org/cool/dj6p-xxc3" name="Texas Cooperative Wildlife Collection" pageId="8" pageNumber="760">TCWC</collectionCode>
|
||
30660) have an osteoderm overlying the frontoparietal scute. Examined specimens of
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1192,1280,504,525]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1192,1280,504,525]" italics="true" pageId="8" pageNumber="760">Barisia</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1341,1436,503,524]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1341,1436,503,524]" italics="true" pageId="8" pageNumber="760">Abronia</emphasis>
|
||
</taxonomicName>
|
||
, excluding
|
||
<taxonomicName authorityName="Campbell" authorityYear="1984" box="[944,1067,534,555]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ornelasi">
|
||
<emphasis box="[944,1067,534,555]" italics="true" pageId="8" pageNumber="760">A. ornelasi</emphasis>
|
||
</taxonomicName>
|
||
and species previously placed in
|
||
<taxonomicName box="[827,933,565,586]" class="Reptilia" family="Anguidae" genus="Mesaspis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[827,933,565,586]" italics="true" pageId="8" pageNumber="760">Mesaspis</emphasis>
|
||
</taxonomicName>
|
||
, display contact of the posterior internasal and prefrontal osteoderms.
|
||
<taxonomicName authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[1167,1367,595,617]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="cuyama">
|
||
<emphasis box="[1167,1367,595,617]" italics="true" pageId="8" pageNumber="760">Abronia cuyama</emphasis>
|
||
</taxonomicName>
|
||
has a frontonasal osteoderm, which is absent in
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1302,1386,627,648]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1302,1386,627,648]" italics="true" pageId="8" pageNumber="760">Barisia</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation captionStart="Figure 4" captionStartId="7.[143,221,1035,1057]" captionTargetBox="[225,1344,197,994]" captionTargetId="figure-481@7.[223,1346,195,995]" captionTargetPageId="7" captionText="Figure 4. Comparison of Abronia cuyama sp. nov. and skulls of extant gerrhonotines, in anterior view. A, Abronia cuyama sp. nov. UCMP 54560. B, Abronia graminea UTA 38831. C, Elgaria multicarinata TNHC 35666. D, Barisia leƲicollis MVZ 68783. Scale bars: 1 mm." figureDoi="http://doi.org/10.5281/zenodo.7695203" httpUri="https://zenodo.org/record/7695203/files/figure.png" pageId="8" pageNumber="760">Fig. 4D</figureCitation>
|
||
), and has a flat anterior surface of the nasal process of the premaxilla, unlike
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1114,1198,688,709]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1114,1198,688,709]" italics="true" pageId="8" pageNumber="760">Barisia</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph blockId="8.[827,1444,197,1905]" pageId="8" pageNumber="760">
|
||
The extinct gerrhonotines
|
||
<taxonomicName authorityName="Estes" authorityYear="1963" box="[1205,1383,718,740]" class="Squamata" family="Anguidae" genus="Paragerrhonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ricardensis">
|
||
<emphasis box="[1205,1383,718,740]" italics="true" pageId="8" pageNumber="760">P. ricardensis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Wilson" authorityYear="1968" box="[827,1023,749,770]" class="Reptilia" family="Anguidae" genus="Gerrhonotus" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="mungerorum">
|
||
<emphasis box="[827,1023,749,770]" italics="true" pageId="8" pageNumber="760">G. mungerorum</emphasis>
|
||
</taxonomicName>
|
||
were previously suggested to be part of or closely related to
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1154,1248,779,800]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1154,1248,779,800]" italics="true" pageId="8" pageNumber="760">Abronia</emphasis>
|
||
</taxonomicName>
|
||
based primarily on the presence of heavily sculptured osteoderms and relatively long, fang-like teeth (
|
||
<bibRefCitation author="Good DA" box="[1277,1427,841,863]" pageId="8" pageNumber="760" pagination="188 - 195" refId="ref10291" refString="Good DA. 1988 a. The phylogenetic position of fossils assigned to the Gerrhonotinae (Squamata: Anguidae). Journal of Vertebrate Paleontology 8: 188 - 195." type="journal article" year="1988">Good, 1988a</bibRefCitation>
|
||
), but neither was ever formally placed in
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1337,1435,871,892]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1337,1435,871,892]" italics="true" pageId="8" pageNumber="760">Abronia</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
<taxonomicName authorityName="Estes" authorityYear="1963" box="[827,1173,902,923]" class="Squamata" family="Anguidae" genus="Paragerrhonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ricardensis">
|
||
<emphasis box="[827,1173,902,923]" italics="true" pageId="8" pageNumber="760">Paragerrhonotus ricardensis</emphasis>
|
||
</taxonomicName>
|
||
was described from a partial skull (
|
||
<bibRefCitation author="Estes R" box="[989,1122,933,955]" pageId="8" pageNumber="760" pagination="676 - 680" refId="ref9679" refString="Estes R. 1963. A new gerrhonotine lizard from the Pliocene of California. Copeia 1963: 676 - 680." type="journal article" year="1963">Estes, 1963</bibRefCitation>
|
||
). Phylogenetic studies that included
|
||
<taxonomicName authorityName="Estes" authorityYear="1963" box="[931,1084,963,985]" class="Squamata" family="Anguidae" genus="Paragerrhonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ricardensis">
|
||
<emphasis box="[931,1084,963,985]" italics="true" pageId="8" pageNumber="760">P. ricardensis</emphasis>
|
||
</taxonomicName>
|
||
placed the taxon in a polytomy with
|
||
<taxonomicName authorityName="Wiegmann" authorityYear="1828" box="[888,1034,994,1015]" class="Reptilia" family="Anguidae" genus="Gerrhonotus" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[888,1034,994,1015]" italics="true" pageId="8" pageNumber="760">Gerrhonotus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1048,1135,995,1016]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1048,1135,995,1016]" italics="true" pageId="8" pageNumber="760">Barisia</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1196,1290,994,1015]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1196,1290,994,1015]" italics="true" pageId="8" pageNumber="760">Abronia</emphasis>
|
||
</taxonomicName>
|
||
or as a stem gerrhonotine (
|
||
<bibRefCitation author="Conrad JL & Norell MA" box="[1000,1275,1025,1047]" pageId="8" pageNumber="760" pagination="1 - 24" refId="ref9595" refString="Conrad JL, Norell MA. 2008. The braincases of two glyptosaurines (Anguidae, Squamata) and anguid phylogeny. American Museum NoVitates 3613: 1 - 24." type="journal article" year="2008">Conrad & Norell, 2008</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Norell MA & Gao K & Conrad J" pageId="8" pageNumber="760" pagination="1 - 22" refId="ref10979" refString="Norell MA, Gao K, Conrad J. 2008. A new platynotan lizard (Diapsida: Squamata) from the Late Cretaceous Gobi Desert. American Museum NoVitates 3605: 1 - 22." type="journal article" year="2008">
|
||
Norell
|
||
<emphasis box="[1376,1436,1025,1046]" italics="true" pageId="8" pageNumber="760">et al.</emphasis>
|
||
, 2008
|
||
</bibRefCitation>
|
||
). Based on examination of the
|
||
<typeStatus box="[1237,1334,1055,1077]" pageId="8" pageNumber="760">holotype</typeStatus>
|
||
and with reference to the diagnostic characters listed by
|
||
<bibRefCitation author="Estes R" pageId="8" pageNumber="760" pagination="676 - 680" refId="ref9679" refString="Estes R. 1963. A new gerrhonotine lizard from the Pliocene of California. Copeia 1963: 676 - 680." type="journal article" year="1963">Estes (1963)</bibRefCitation>
|
||
,
|
||
<taxonomicName authorityName="Estes" authorityYear="1963" box="[917,1081,1117,1138]" class="Squamata" family="Anguidae" genus="Paragerrhonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ricardensis">
|
||
<emphasis box="[917,1081,1117,1138]" italics="true" pageId="8" pageNumber="760">P. ricardensis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[1144,1271,1117,1138]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="cuyama">
|
||
<emphasis box="[1144,1271,1117,1138]" italics="true" pageId="8" pageNumber="760">A. cuyama</emphasis>
|
||
</taxonomicName>
|
||
share heavily sculptured osteoderms (although the texture is less vermiculate in
|
||
<taxonomicName authorityName="Estes" authorityYear="1963" box="[1000,1191,1178,1199]" class="Squamata" family="Anguidae" genus="Paragerrhonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1000,1191,1178,1199]" italics="true" pageId="8" pageNumber="760">Paragerrhonotus</emphasis>
|
||
</taxonomicName>
|
||
, similar to
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1323,1408,1179,1200]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1323,1408,1179,1200]" italics="true" pageId="8" pageNumber="760">Barisia</emphasis>
|
||
</taxonomicName>
|
||
), a large jugal spur, and sharp, recurved mesial teeth on the maxilla. A large jugal spur and sharp and recurved maxillary teeth are present in many gerrhonotines (
|
||
<bibRefCitation author="Gauthier JA" box="[835,1004,1301,1323]" pageId="8" pageNumber="760" pagination="7 - 54" refId="ref10106" refString="Gauthier JA. 1982. Fossil xenosaurid and anguid lizards from the early Eocene Wasatch Formation, southeast Wyoming, and a revision of the Anguioidea. Contributions to Geology, UniVersity of Wyoming 21: 7 - 54." type="journal article" year="1982">Gauthier, 1982</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Ledesma DT & Scarpetta SG & Bell CJ" box="[1017,1250,1301,1323]" pageId="8" pageNumber="760" pagination="11602" refId="ref10854" refString="Ledesma DT, Scarpetta SG, Bell CJ. 2021. Variation in the skulls of Elgaria and Gerrhonotus (Anguidae, Gerrhonotinae) and implications for phylogenetics and fossil identification. PeerJ 9: e 11602." type="journal article" year="2021">
|
||
Ledesma
|
||
<emphasis box="[1126,1183,1301,1322]" italics="true" pageId="8" pageNumber="760">et al.</emphasis>
|
||
, 2021
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Scarpetta SG & Ledesma DT & Bell CJ" pageId="8" pageNumber="760" pagination="1 - 58" refId="ref11478" refString="Scarpetta SG, Ledesma DT, Bell CJ. 2021. A new extinct species of alligator lizard (Squamata: Elgaria) and an expanded perspective on the osteology and phylogeny of Gerrhonotinae. BMC Ecology and EVolution 21: 1 - 58." type="journal article" year="2021">
|
||
Scarpetta
|
||
<emphasis box="[1381,1437,1301,1322]" italics="true" pageId="8" pageNumber="760">et al.</emphasis>
|
||
, 2021
|
||
</bibRefCitation>
|
||
), and neither was used here as a phylogenetic character.
|
||
<taxonomicName authorityName="Estes" authorityYear="1963" box="[953,1292,1362,1383]" class="Squamata" family="Anguidae" genus="Paragerrhonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ricardensis">
|
||
<emphasis box="[953,1292,1362,1383]" italics="true" pageId="8" pageNumber="760">Paragerrhonotus ricardensis</emphasis>
|
||
</taxonomicName>
|
||
differs from
|
||
<taxonomicName authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[827,946,1393,1414]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="cuyama">
|
||
<emphasis box="[827,946,1393,1414]" italics="true" pageId="8" pageNumber="760">A. cuyama</emphasis>
|
||
</taxonomicName>
|
||
in possessing an elongate frontal, in lacking osteoderm keels, in lacking a ventral lamina of the orbital process of the jugal, in possessing a distinctive arrangement of small osteoderms on the posterior portion of the frontal (not adapted as a phylogenetic character) and in possessing a premaxillary nasal process with a concave dorsal surface.
|
||
<taxonomicName authorityName="Scarpetta & Ledesma" authorityYear="2023" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="cuyama">
|
||
<emphasis italics="true" pageId="8" pageNumber="760">Abronia cuyama</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Estes" authorityYear="1963" box="[980,1142,1607,1629]" class="Squamata" family="Anguidae" genus="Paragerrhonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ricardensis">
|
||
<emphasis box="[980,1142,1607,1629]" italics="true" pageId="8" pageNumber="760">P. ricardensis</emphasis>
|
||
</taxonomicName>
|
||
do not share any unique synapomorphies or apomorphic character state combinations.
|
||
</paragraph>
|
||
<paragraph blockId="8.[827,1444,197,1905]" lastBlockId="9.[145,761,197,1200]" lastPageId="9" lastPageNumber="761" pageId="8" pageNumber="760">
|
||
We also note that
|
||
<taxonomicName authorityName="Estes" authorityYear="1963" box="[1073,1236,1699,1721]" class="Squamata" family="Anguidae" genus="Paragerrhonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ricardensis">
|
||
<emphasis box="[1073,1236,1699,1721]" italics="true" pageId="8" pageNumber="760">P. ricardensis</emphasis>
|
||
</taxonomicName>
|
||
has a single row of four pterygoid teeth, as originally stated by
|
||
<bibRefCitation author="Estes R" pageId="8" pageNumber="760" pagination="676 - 680" refId="ref9679" refString="Estes R. 1963. A new gerrhonotine lizard from the Pliocene of California. Copeia 1963: 676 - 680." type="journal article" year="1963">Estes (1963)</bibRefCitation>
|
||
and
|
||
<emphasis box="[957,1030,1761,1782]" italics="true" pageId="8" pageNumber="760">contra</emphasis>
|
||
<bibRefCitation author="Good DA" box="[1038,1190,1761,1783]" pageId="8" pageNumber="760" pagination="188 - 195" refId="ref10291" refString="Good DA. 1988 a. The phylogenetic position of fossils assigned to the Gerrhonotinae (Squamata: Anguidae). Journal of Vertebrate Paleontology 8: 188 - 195." type="journal article" year="1988">Good (1988a)</bibRefCitation>
|
||
. The loss of pterygoid teeth is another feature mentioned by
|
||
<bibRefCitation author="Good DA" box="[1264,1414,1791,1813]" pageId="8" pageNumber="760" pagination="188 - 195" refId="ref10291" refString="Good DA. 1988 a. The phylogenetic position of fossils assigned to the Gerrhonotinae (Squamata: Anguidae). Journal of Vertebrate Paleontology 8: 188 - 195." type="journal article" year="1988">Good (1988a)</bibRefCitation>
|
||
to suggest a close relationship between
|
||
<taxonomicName authorityName="Estes" authorityYear="1963" box="[1242,1394,1822,1844]" class="Squamata" family="Anguidae" genus="Paragerrhonotus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="species" species="ricardensis">
|
||
<emphasis box="[1242,1394,1822,1844]" italics="true" pageId="8" pageNumber="760">P. ricardensis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[827,921,1853,1874]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[827,921,1853,1874]" italics="true" pageId="8" pageNumber="760">Abronia</emphasis>
|
||
</taxonomicName>
|
||
. The loss of pterygoid teeth was interpreted by several authors as a derived feature of
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[1347,1443,1883,1904]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="760" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1347,1443,1883,1904]" italics="true" pageId="8" pageNumber="760">Abronia</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[195,279,198,219]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="genus">
|
||
<emphasis box="[195,279,198,219]" italics="true" pageId="9" pageNumber="761">Barisia</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Good DA" box="[295,421,197,219]" pageId="9" pageNumber="761" pagination="285 - 297" refId="ref10266" refString="Good DA. 1987. A phylogenetic analysis of cranial osteology in the gerrhonotine lizards. Journal of Herpetology 21: 285 - 297." type="journal article" year="1987">Good, 1987</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Scarpetta SG & Ledesma DT & Bell CJ" box="[434,676,197,219]" pageId="9" pageNumber="761" pagination="1 - 58" refId="ref11478" refString="Scarpetta SG, Ledesma DT, Bell CJ. 2021. A new extinct species of alligator lizard (Squamata: Elgaria) and an expanded perspective on the osteology and phylogeny of Gerrhonotinae. BMC Ecology and EVolution 21: 1 - 58." type="journal article" year="2021">
|
||
Scarpetta
|
||
<emphasis box="[552,608,197,219]" italics="true" pageId="9" pageNumber="761">et al.</emphasis>
|
||
, 2021
|
||
</bibRefCitation>
|
||
). A few examined specimens of
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[416,508,228,249]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="genus">
|
||
<emphasis box="[416,508,228,249]" italics="true" pageId="9" pageNumber="761">Abronia</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Gray" authorityYear="1838" box="[567,652,228,249]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="genus">
|
||
<emphasis box="[567,652,228,249]" italics="true" pageId="9" pageNumber="761">Barisia</emphasis>
|
||
</taxonomicName>
|
||
have one to three irregularly arranged pterygoid teeth (e.g.
|
||
<taxonomicName authority="UTA" authorityName="UTA" box="[145,347,289,310]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="campbelli">
|
||
<emphasis box="[145,287,289,310]" italics="true" pageId="9" pageNumber="761">A. campbelli</emphasis>
|
||
UTA
|
||
</taxonomicName>
|
||
95952;
|
||
<taxonomicName authority="FMNH" authorityName="FMNH" box="[435,632,289,311]" class="Reptilia" family="Anguidae" genus="Barisia" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="ciliaris">
|
||
<emphasis box="[435,546,289,311]" italics="true" pageId="9" pageNumber="761">B. ciliaris</emphasis>
|
||
FMNH
|
||
</taxonomicName>
|
||
30707).
|
||
</paragraph>
|
||
<paragraph blockId="9.[145,761,197,1200]" pageId="9" pageNumber="761">
|
||
<taxonomicName authorityName="Wilson" authorityYear="1968" box="[169,476,320,341]" class="Reptilia" family="Anguidae" genus="Gerrhonotus" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="mungerorum">
|
||
<emphasis box="[169,476,320,341]" italics="true" pageId="9" pageNumber="761">Gerrhonotus mungerorum</emphasis>
|
||
</taxonomicName>
|
||
was described based on a single, isolated frontal (
|
||
<bibRefCitation author="Wilson RL" box="[457,611,350,372]" pageId="9" pageNumber="761" pagination="75 - 126" refId="ref11909" refString="Wilson RL. 1968. Systematics and faunal analysis of a lower Pliocene vertebrate assemblage from Trego County, Kansas. Contributions from the Museum of Paleontology UniVersity of Michigan 22: 75 - 126." type="journal article" year="1968">Wilson, 1968</bibRefCitation>
|
||
). Additional fossils, including partial dentaries, maxillae and a parietal, were later attributed to the taxon from the type locality and other localities (e.g.
|
||
<bibRefCitation author="Holman JA" pageId="9" pageNumber="761" pagination="125 - 134" refId="ref10452" refString="Holman JA. 1973. Reptiles of the Egelhoff herpetofauna (Upper Miocene) of Nebraska. Contributions from the Museum of Paleontology UniVersity of Michigan 24: 125 - 134." type="journal article" year="1973">Holman, 1973</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Holman JA" box="[220,278,473,495]" pageId="9" pageNumber="761" pagination="49 - 66" refId="ref10484" refString="Holman JA. 1975. Herpetofauna of the WaKeeney local fauna (Lower Pliocene: Clarendonian) of Trego County, Kansas. UniVersity of Michigan Papers on Paleontology 12: 49 - 66." type="journal article" year="1975">1975</bibRefCitation>
|
||
), but those identifications were either explicitly tentative or based on features not unique to
|
||
<taxonomicName authorityName="Wilson" authorityYear="1968" box="[145,329,535,556]" class="Reptilia" family="Anguidae" genus="Gerrhonotus" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="mungerorum">
|
||
<emphasis box="[145,329,535,556]" italics="true" pageId="9" pageNumber="761">G. mungerorum</emphasis>
|
||
</taxonomicName>
|
||
with respect to other gerrhonotines.
|
||
<taxonomicName authorityName="Wilson" authorityYear="1968" box="[145,465,565,587]" class="Reptilia" family="Anguidae" genus="Gerrhonotus" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="mungerorum">
|
||
<emphasis box="[145,465,565,587]" italics="true" pageId="9" pageNumber="761">Gerrhonotus mungerorum</emphasis>
|
||
</taxonomicName>
|
||
requires further study, especially the tentatively referred fossils, and a systematic assessment of the species would benefit from the procurement of additional fossils from the type locality. We have not examined the
|
||
<typeStatus box="[628,728,688,710]" pageId="9" pageNumber="761">holotype</typeStatus>
|
||
or attributed specimens of
|
||
<taxonomicName authorityName="Wilson" authorityYear="1968" box="[442,631,719,740]" class="Reptilia" family="Anguidae" genus="Gerrhonotus" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="mungerorum">
|
||
<emphasis box="[442,631,719,740]" italics="true" pageId="9" pageNumber="761">G. mungerorum</emphasis>
|
||
</taxonomicName>
|
||
in person; therefore, based on the description of the
|
||
<typeStatus box="[614,711,749,771]" pageId="9" pageNumber="761">holotype</typeStatus>
|
||
and descriptions of referred fossils,
|
||
<taxonomicName authorityName="Wilson" authorityYear="1968" box="[521,707,780,801]" class="Reptilia" family="Anguidae" genus="Gerrhonotus" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="mungerorum">
|
||
<emphasis box="[521,707,780,801]" italics="true" pageId="9" pageNumber="761">G. mungerorum</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[145,269,811,832]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="cuyama">
|
||
<emphasis box="[145,269,811,832]" italics="true" pageId="9" pageNumber="761">A. cuyama</emphasis>
|
||
</taxonomicName>
|
||
share a heavily sculptured frontal, sharp and recurved teeth, and the presence of near bicuspid crowns on the distal teeth. Sharp and recurved teeth and near bicuspid distal crowns are common to many gerrhonotines (
|
||
<bibRefCitation author="Scarpetta SG & Ledesma DT & Bell CJ" box="[321,565,933,955]" pageId="9" pageNumber="761" pagination="1 - 58" refId="ref11478" refString="Scarpetta SG, Ledesma DT, Bell CJ. 2021. A new extinct species of alligator lizard (Squamata: Elgaria) and an expanded perspective on the osteology and phylogeny of Gerrhonotinae. BMC Ecology and EVolution 21: 1 - 58." type="journal article" year="2021">
|
||
Scarpetta
|
||
<emphasis box="[440,497,933,955]" italics="true" pageId="9" pageNumber="761">et al.</emphasis>
|
||
, 2021
|
||
</bibRefCitation>
|
||
), and the former was not used as a phylogenetic character.
|
||
<taxonomicName authorityName="Wilson" authorityYear="1968" class="Reptilia" family="Anguidae" genus="Gerrhonotus" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="mungerorum">
|
||
<emphasis italics="true" pageId="9" pageNumber="761">Gerrhonotus mungerorum</emphasis>
|
||
</taxonomicName>
|
||
differs from
|
||
<taxonomicName authorityName="Scarpetta & Ledesma" authorityYear="2023" box="[466,594,995,1016]" class="Reptilia" family="Anguidae" genus="Abronia" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="cuyama">
|
||
<emphasis box="[466,594,995,1016]" italics="true" pageId="9" pageNumber="761">A. cuyama</emphasis>
|
||
</taxonomicName>
|
||
in having an elongate frontal and in lacking keels on the frontal osteoderm crust.
|
||
</paragraph>
|
||
<paragraph blockId="9.[145,761,197,1200]" lastBlockId="9.[809,1425,197,525]" pageId="9" pageNumber="761">
|
||
Cranial osteoderm keels are present in
|
||
<taxonomicName authority="Blainville, 1835" authorityName="Blainville" authorityYear="1835" class="Reptilia" family="Anguidae" genus="Elgaria" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="multicarinata">
|
||
<emphasis italics="true" pageId="9" pageNumber="761">Elgaria multicarinata</emphasis>
|
||
Blainville, 1835
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authority="Fitch, 1934" authorityName="Fitch" authorityYear="1934" class="Reptilia" family="Anguidae" genus="Elgaria" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="nana">
|
||
<emphasis box="[539,690,1117,1139]" italics="true" pageId="9" pageNumber="761">Elgaria nana</emphasis>
|
||
Fitch, 1934
|
||
</taxonomicName>
|
||
, although neither of those species has osteoderm keels anterior to the frontal or on the frontal (
|
||
<bibRefCitation author="Ledesma DT & Scarpetta SG & Bell CJ" box="[658,948,197,1200]" pageId="9" pageNumber="761" pagination="11602" refId="ref10854" refString="Ledesma DT, Scarpetta SG, Bell CJ. 2021. Variation in the skulls of Elgaria and Gerrhonotus (Anguidae, Gerrhonotinae) and implications for phylogenetics and fossil identification. PeerJ 9: e 11602." type="journal article" year="2021">
|
||
Ledesma
|
||
<emphasis box="[809,872,197,219]" italics="true" pageId="9" pageNumber="761">et al.</emphasis>
|
||
, 2021
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Scarpetta SG & Ledesma DT & Bell CJ" box="[967,1237,197,219]" pageId="9" pageNumber="761" pagination="1 - 58" refId="ref11478" refString="Scarpetta SG, Ledesma DT, Bell CJ. 2021. A new extinct species of alligator lizard (Squamata: Elgaria) and an expanded perspective on the osteology and phylogeny of Gerrhonotinae. BMC Ecology and EVolution 21: 1 - 58." type="journal article" year="2021">
|
||
Scarpetta
|
||
<emphasis box="[1097,1161,197,219]" italics="true" pageId="9" pageNumber="761">et al.</emphasis>
|
||
, 2021
|
||
</bibRefCitation>
|
||
). Among other squamates, nuchal osteoderm sails were observed on
|
||
<taxonomicName authority="Stanley, Ceriaco, Bandeira, Valerio, Bates & Branch, 2016" authorityName="Stanley, Ceriaco, Bandeira, Valerio, Bates & Branch" authorityYear="2016" class="Reptilia" family="Cordylidae" genus="Cordylus" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="namakuiyus">
|
||
<emphasis box="[844,1101,258,279]" italics="true" pageId="9" pageNumber="761">Cordylus namakuiyus</emphasis>
|
||
Stanley, Ceríaco, Bandeira, Valerio, Bates & Branch, 2016
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authority="Bocage, 1895" authorityName="Bocage" authorityYear="1895" class="Reptilia" family="Cordylidae" genus="Cordylus" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="angolensis">
|
||
<emphasis box="[1198,1425,289,310]" italics="true" pageId="9" pageNumber="761">Cordylus angolensis</emphasis>
|
||
Bocage, 1895
|
||
</taxonomicName>
|
||
(CAS 254912 and AMNH 47333, respectively). Among other anguimorphs, postcranial osteoderm keels were observed in
|
||
<taxonomicName authority="Ahl, 1930" authorityName="Ahl" authorityYear="1930" class="Reptilia" family="Shinisauridae" genus="Shinisaurus" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="crocodilurus">
|
||
<emphasis italics="true" pageId="9" pageNumber="761">Shinisaurus crocodilurus</emphasis>
|
||
Ahl, 1930
|
||
</taxonomicName>
|
||
(UF 45615 and FMNH 215541), and keels are present on some cranial osteoderms and the frontal osteodermal crust in some
|
||
<taxonomicName box="[1240,1373,474,495]" class="Reptilia" family="Xenosauridae" genus="Xenosaurus" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1240,1373,474,495]" italics="true" pageId="9" pageNumber="761">Xenosaurus</emphasis>
|
||
</taxonomicName>
|
||
(e.g.
|
||
<taxonomicName authority="Gray, 1856" authorityName="Gray" authorityYear="1856" box="[809,1167,503,525]" class="Reptilia" family="Xenosauridae" genus="Xenosaurus" kingdom="Animalia" order="Squamata" pageId="9" pageNumber="761" phylum="Chordata" rank="species" species="grandis">
|
||
<emphasis box="[809,1036,504,525]" italics="true" pageId="9" pageNumber="761">Xenosaurus grandis</emphasis>
|
||
Gray, 1856
|
||
</taxonomicName>
|
||
; FMNH 123702).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |