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<document ID-DOI="http://doi.org/10.5281/zenodo.5013967" ID-GBIF-Dataset="a0723a89-d4df-4e36-9da9-cc6fd438c7c3" ID-GBIF-Taxon="183116066" ID-Zenodo-Dep="5013967" approvedBy="poa" checkinTime="1624379932173" checkinUser="tatiana" docAuthor="Bolet, Arnau, Stanley, Edward L., Daza, Juan D., Arias, J. Salvador, Cernanský, Andrej, Vidal-García, Marta, Bauer, Aaron M., Bevitt, Joseph J., Peretti, Adolf &amp; Evans, Susan E." docDate="2021" docId="714E87AE6E11FFB49E66FC95FC8FF875" docLanguage="en" docName="CurrBiol.31.1-12.pdf" docOrigin="Current Biology 31" docTitle="Oculudentavis naga Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans 2021, new species" docType="treatment" docVersion="6" lastPageId="8" lastPageNumber="8" masterDocId="8D77FFD66E15FFBC9D5BFF95FF90FFB6" masterDocTitle="Unusual morphology in the mid-Cretaceous lizard Oculudentavis" masterLastPageNumber="12" masterPageNumber="1" pageId="4" pageNumber="4" updateTime="1643430857924" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-ND-4.0" zenodo-license-figures="CC-BY-NC-ND-4.0">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Unusual morphology in the mid-Cretaceous lizard Oculudentavis</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bolet, Arnau</mods:namePart>
<mods:affiliation>Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Barcelona, Spain &amp; School of Earth Sciences, University of Bristol, Bristol, UK</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Stanley, Edward L.</mods:namePart>
<mods:affiliation>Department of Natural History, Florida Museum of Natural History, Gainesville, FL, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Daza, Juan D.</mods:namePart>
<mods:affiliation>Department of Biological Sciences, Sam Houston State University, Huntsville, TX, USA</mods:affiliation>
<mods:nameIdentifier type="email">juand.daza@gmail.com</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Arias, J. Salvador</mods:namePart>
<mods:affiliation>Unidad Ejecutora Lillo (CONICET - Fundación Miguel Lillo), San Miguel, de Tucumán, Tucumán, Argentina</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Cernanský, Andrej</mods:namePart>
<mods:affiliation>Department of Ecology, Laboratory of Evolutionary Biology, Faculty of Natural Sciences, Comenius University in Bratislava, Bratislava, Slovakia</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Vidal-García, Marta</mods:namePart>
<mods:affiliation>Department of Cell Biology &amp; Anatomy, University of Calgary, Calgary, AB, Canada</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bauer, Aaron M.</mods:namePart>
<mods:affiliation>Department of Biology and Center for Biodiversity and Ecosystem Stewardship, Villanova University, Villanova, PA, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Bevitt, Joseph J.</mods:namePart>
<mods:affiliation>Australian Centre for Neutron Scattering, Australian Nuclear Science and Technology Organisation, Sydney, NSW, Australia</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Peretti, Adolf</mods:namePart>
<mods:affiliation>GRS Gemresearch Swisslab AG and Peretti Museum Foundation, Meggen, Switzerland</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Evans, Susan E.</mods:namePart>
<mods:affiliation>Department of Cell and Developmental Biology, University College London, London, UK</mods:affiliation>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>Current Biology</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2021</mods:date>
<mods:detail type="pubDate">
<mods:number>2021-06-14</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>31</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>1</mods:start>
<mods:end>12</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1016/j.cub.2021.05.040</mods:identifier>
<mods:identifier type="GBIF-Dataset">a0723a89-d4df-4e36-9da9-cc6fd438c7c3</mods:identifier>
<mods:identifier type="Zenodo-Dep">5013953</mods:identifier>
</mods:mods>
<treatment ID-DOI="http://doi.org/10.5281/zenodo.5013967" ID-GBIF-Taxon="183116066" ID-Zenodo-Dep="5013967" LSID="urn:lsid:plazi:treatment:714E87AE6E11FFB49E66FC95FC8FF875" httpUri="http://treatment.plazi.org/id/714E87AE6E11FFB49E66FC95FC8FF875" lastPageId="8" lastPageNumber="8" pageId="4" pageNumber="4">
<subSubSection box="[829,1197,768,789]" pageId="4" pageNumber="4" type="nomenclature">
<paragraph blockId="4.[829,1468,768,1870]" box="[829,1197,768,789]" pageId="4" pageNumber="4">
<heading bold="true" box="[829,1197,768,789]" fontSize="8" level="8" pageId="4" pageNumber="4" reason="6">
<emphasis bold="true" box="[829,1197,768,789]" pageId="4" pageNumber="4">
<taxonomicName authority="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans, 2021" authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[829,1051,768,789]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="naga" status="new species">
<emphasis bold="true" box="[829,1051,768,789]" italics="true" pageId="4" pageNumber="4">Oculudentavis naga</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[1057,1197,768,789]" pageId="4" pageNumber="4" rank="species">new species</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="4" type="materials_examined">
<paragraph blockId="4.[829,1468,768,1870]" pageId="4" pageNumber="4">
<materialsCitation ID-GBIF-Occurrence="3307551302" collectionCode="A" country="Afghanistan" pageId="4" pageNumber="4" specimenCode="GRS-Ref-28627" specimenCount="1" stateProvince="Three-dimensional Model Of New Specimen Available At Https" typeStatus="holotype">
<heading bold="true" box="[829,929,797,818]" fontSize="8" level="8" pageId="4" pageNumber="4" reason="6">
<emphasis bold="true" box="[829,929,797,818]" italics="true" pageId="4" pageNumber="4">
<typeStatus box="[829,929,797,818]" pageId="4" pageNumber="4">Holotype</typeStatus>
</emphasis>
</heading>
Peretti Museum Foundation,
<specimenCode box="[1123,1288,826,847]" pageId="4" pageNumber="4">GRS-Ref-28627</specimenCode>
, a skull and anterior postcranial skeleton (
<figureCitation box="[1095,1192,856,877]" captionStart="Figure 1" captionStartId="2.[1053,1113,271,290]" captionTargetBox="[159,1016,271,1338]" captionTargetId="figure-191@2.[159,1016,271,1338]" captionTargetPageId="2" captionText="Figure 1. Oculudentavis naga (GRS-Ref- 286278) displaying the superb preservation of bone and soft tissue (A) Dorsal; (B) lateral; (C) ventral views. Scale bar represents 10 mm. Diptera associated with the lizard skeleton were identified as Phoridae, Platypezidae, Ceratopogonidae, or Brachycera (Empidoidea). See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5013955" httpUri="https://zenodo.org/record/5013955/files/figure.png" pageId="4" pageNumber="4">Figures 1</figureCitation>
,
<figureCitation box="[1210,1275,856,877]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="4" pageNumber="4">2A2J</figureCitation>
,
<figureCitation box="[1289,1365,856,877]" captionStart="Figure 3" captionStartId="5.[1038,1098,271,290]" captionTargetBox="[142,1003,271,943]" captionTargetId="figure-327@5.[142,1003,271,943]" captionTargetPageId="5" captionText="Figure 3. Comparison of the two specimens of Oculudentavis, details of the lacrimal and dentary Right side view of the orbit showing the ring-like lacrimal bone (salmon color) in situ in (A) O. naga and (B) O. khaungraae (orbit view here is posterolateral to show the orbital elements more clearly). Dorsolingual view of the dentary in (C) O. naga and (D) O. khaungraae is shown. Scale bar represents 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5013959" httpUri="https://zenodo.org/record/5013959/files/figure.png" pageId="4" pageNumber="4">3A, 3C</figureCitation>
, and S
<figureCitation box="[1446,1454,856,877]" captionStart="Figure 1" captionStartId="2.[1053,1113,271,290]" captionTargetBox="[159,1016,271,1338]" captionTargetId="figure-191@2.[159,1016,271,1338]" captionTargetPageId="2" captionText="Figure 1. Oculudentavis naga (GRS-Ref- 286278) displaying the superb preservation of bone and soft tissue (A) Dorsal; (B) lateral; (C) ventral views. Scale bar represents 10 mm. Diptera associated with the lizard skeleton were identified as Phoridae, Platypezidae, Ceratopogonidae, or Brachycera (Empidoidea). See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5013955" httpUri="https://zenodo.org/record/5013955/files/figure.png" pageId="4" pageNumber="4">1</figureCitation>
). Three-dimensional model of new specimen available at https:// tinyurl.com/Oculudentavis-L-10420.
</materialsCitation>
</paragraph>
<paragraph blockId="4.[829,1468,768,1870]" box="[829,971,943,964]" pageId="4" pageNumber="4">
<emphasis bold="true" box="[829,971,943,964]" italics="true" pageId="4" pageNumber="4">
<typeStatus box="[829,883,943,964]" pageId="4" pageNumber="4">Type</typeStatus>
locality
</emphasis>
</paragraph>
<paragraph blockId="4.[829,1468,768,1870]" pageId="4" pageNumber="4">
<materialsCitation ID-GBIF-Occurrence="3307551304" collectionCode="O" country="Iceland" location="Aung Bar mine" longitude="-34.0" pageId="4" pageNumber="4" specimenCode="GRS-Ref-28627" specimenCount="1" typeStatus="holotype">
The holotype specimen of
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[1092,1288,972,993]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1092,1288,972,993]" italics="true" pageId="4" pageNumber="4">Oculudentavis naga</emphasis>
</taxonomicName>
(
<specimenCode box="[1298,1461,972,993]" pageId="4" pageNumber="4">GRS-Ref-28627</specimenCode>
) and the holotype of
<taxonomicName authorityName="Xing et al." box="[1033,1180,1002,1023]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1033,1180,1002,1023]" italics="true" pageId="4" pageNumber="4">O. khaungraae</emphasis>
</taxonomicName>
(HPG-15-3) were recovered from the same mine (
<location LSID="urn:lsid:plazi:treatment:714E87AE6E11FFB49E66FC95FC8FF875:FC3860636E11FFB89949FB92FB3CFBAA" box="[1042,1196,1031,1052]" country="Iceland" longitude="-34.0" name="Aung Bar mine" pageId="4" pageNumber="4">Aung Bar mine</location>
,
<geoCoordinate box="[1204,1301,1028,1052]" pageId="4" pageNumber="4">
26
<emphasis bold="true" box="[1229,1237,1028,1042]" pageId="4" pageNumber="4">
<superScript attach="left" box="[1229,1237,1028,1042]" fontSize="6" pageId="4" pageNumber="4">O</superScript>
</emphasis>
09
<emphasis bold="true" box="[1270,1274,1028,1042]" pageId="4" pageNumber="4">
<superScript attach="left" box="[1270,1274,1028,1042]" fontSize="6" pageId="4" pageNumber="4">0</superScript>
</emphasis>
N
</geoCoordinate>
,
<geoCoordinate box="[1310,1404,1028,1052]" pageId="4" pageNumber="4">
96
<emphasis bold="true" box="[1335,1343,1028,1042]" pageId="4" pageNumber="4">
<superScript attach="left" box="[1335,1343,1028,1042]" fontSize="6" pageId="4" pageNumber="4">O</superScript>
</emphasis>
34
<emphasis bold="true" box="[1376,1380,1028,1042]" pageId="4" pageNumber="4">
<superScript attach="left" box="[1376,1380,1028,1042]" fontSize="6" pageId="4" pageNumber="4">0</superScript>
</emphasis>
E
</geoCoordinate>
)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="4" type="etymology">
<paragraph blockId="4.[829,1468,768,1870]" box="[829,947,1060,1081]" pageId="4" pageNumber="4">
<emphasis bold="true" box="[829,947,1060,1081]" italics="true" pageId="4" pageNumber="4">Etymology</emphasis>
</paragraph>
<paragraph blockId="4.[829,1468,768,1870]" pageId="4" pageNumber="4">
Combination of
<taxonomicName box="[986,1129,1089,1110]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[986,1129,1089,1110]" italics="true" pageId="4" pageNumber="4">Oculudentavis</emphasis>
</taxonomicName>
(oculus = eye, dentes = teeth, and avis = bird)
<superScript attach="left" box="[937,946,1114,1128]" fontSize="6" pageId="4" pageNumber="4">1</superScript>
and Naga, thename of oneof the many ethnic tribes living in the Burmese amber mines area. The Naga are mentioned in historical chronicles for their prominent role in amber trading. Divided into many sub-groups scattered across the hills and jungle of India (in Nagaland and other states) and in the Tiger valley region of Burma (where amber deposits are found), the Naga tribes are also reputed for their rich and fascinating culture.
</paragraph>
</subSubSection>
<subSubSection pageId="4" pageNumber="4" type="diagnosis">
<paragraph blockId="4.[829,1468,768,1870]" box="[829,939,1352,1373]" pageId="4" pageNumber="4">
<emphasis bold="true" box="[829,939,1352,1373]" italics="true" pageId="4" pageNumber="4">Diagnosis</emphasis>
</paragraph>
<paragraph blockId="4.[829,1468,768,1870]" pageId="4" pageNumber="4">
The
<typeStatus box="[874,961,1382,1403]" pageId="4" pageNumber="4">holotype</typeStatus>
of
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[996,1077,1382,1403]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="naga">
<emphasis box="[996,1077,1382,1403]" italics="true" pageId="4" pageNumber="4">O. naga</emphasis>
</taxonomicName>
(skull length =
<quantity box="[1234,1325,1382,1403]" metricMagnitude="-2" metricUnit="m" metricValue="1.42" pageId="4" pageNumber="4" unit="mm" value="14.2">14.2 mm</quantity>
) is somewhat smaller than that of
<taxonomicName authorityName="Xing et al." box="[1044,1194,1411,1432]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1044,1194,1411,1432]" italics="true" pageId="4" pageNumber="4">O. khaungraae</emphasis>
</taxonomicName>
(skull length =
<quantity box="[1362,1458,1411,1432]" metricMagnitude="-2" metricUnit="m" metricValue="1.73" pageId="4" pageNumber="4" unit="mm" value="17.3">17.3 mm</quantity>
).
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[829,1027,1440,1461]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="naga">
<emphasis box="[829,1027,1440,1461]" italics="true" pageId="4" pageNumber="4">Oculudentavis naga</emphasis>
</taxonomicName>
differs from
<taxonomicName authorityName="Xing et al." box="[1153,1299,1440,1461]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1153,1299,1440,1461]" italics="true" pageId="4" pageNumber="4">O. khaungraae</emphasis>
</taxonomicName>
in having a jugal process of the maxilla that reaches caudally to less than 25% of orbit length; in having a long squamosal process of the postorbital; in having a relatively smaller braincase, with short, distally expanded basipterygoid processes (versus longer, unexpanded processes); and in having anterior palatal rami of pterygoids parallel, diverging posteriorly just behind the fossa columellae, interpterygoid vacuity nearly rectangular (versus divergent pterygoids, heart-shaped vacuity), rostral part of premaxilla shorter and proportionally wider than that of
<taxonomicName authorityName="Xing et al." box="[1216,1364,1703,1724]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1216,1364,1703,1724]" italics="true" pageId="4" pageNumber="4">O. khaungraae</emphasis>
</taxonomicName>
, and less conspicuous platform on the dorsolabial surface of the posterior third of the dentary.
</paragraph>
</subSubSection>
<subSubSection lastPageId="5" lastPageNumber="5" pageId="4" pageNumber="4" type="discussion">
<paragraph blockId="4.[829,1468,768,1870]" box="[829,894,1791,1812]" pageId="4" pageNumber="4">
<emphasis bold="true" box="[829,894,1791,1812]" italics="true" pageId="4" pageNumber="4">Notes</emphasis>
</paragraph>
<paragraph blockId="4.[829,1468,768,1870]" lastBlockId="5.[142,781,1031,1987]" lastPageId="5" lastPageNumber="5" pageId="4" pageNumber="4">
There are also differences between the
<specimenCount box="[1221,1372,1820,1841]" pageId="4" pageNumber="4" type="generic">two specimens</specimenCount>
in the robusticity of the postorbital (greater in
<taxonomicName authorityName="Xing et al." box="[1203,1348,1849,1870]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1203,1348,1849,1870]" italics="true" pageId="4" pageNumber="4">O. khaungraae</emphasis>
</taxonomicName>
); the height of the premaxillary crest (greater in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[509,589,1031,1052]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[509,589,1031,1052]" italics="true" pageId="5" pageNumber="5">O. naga</emphasis>
</taxonomicName>
); the extent of the nasal emargination of the frontal (greater in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[587,667,1060,1081]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[587,667,1060,1081]" italics="true" pageId="5" pageNumber="5">O. naga</emphasis>
</taxonomicName>
); the presence of a large anterior palatine fenestra (
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[575,656,1090,1111]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[575,656,1090,1111]" italics="true" pageId="5" pageNumber="5">O. naga</emphasis>
</taxonomicName>
); the length and height of the coronoid process (larger and taller in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[142,225,1148,1169]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[142,225,1148,1169]" italics="true" pageId="5" pageNumber="5">O. naga</emphasis>
</taxonomicName>
); the shape of the quadrate conch (more angular in
<taxonomicName authorityName="Xing et al." box="[142,289,1177,1198]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[142,289,1177,1198]" italics="true" pageId="5" pageNumber="5">O. khaungraae</emphasis>
</taxonomicName>
); and in the overall shape of the rostrum (more pointed in
<taxonomicName authorityName="Xing et al." box="[248,394,1206,1227]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[248,394,1206,1227]" italics="true" pageId="5" pageNumber="5">O. khaungraae</emphasis>
</taxonomicName>
) and postorbital skull (more vaulted in
<taxonomicName authorityName="Xing et al." box="[142,289,1236,1257]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[142,289,1236,1257]" italics="true" pageId="5" pageNumber="5">O. khaungraae</emphasis>
</taxonomicName>
).
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[310,510,1236,1257]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[310,510,1236,1257]" italics="true" pageId="5" pageNumber="5">Oculudentavis naga</emphasis>
</taxonomicName>
also displays a very large palatal fenestra between the vomers and palatines. This region is poorly preserved in the
<typeStatus box="[414,501,1294,1315]" pageId="5" pageNumber="5">holotype</typeStatus>
of
<taxonomicName authorityName="Xing et al." box="[538,686,1294,1315]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[538,686,1294,1315]" italics="true" pageId="5" pageNumber="5">O. khaungraae</emphasis>
</taxonomicName>
, and the presence or absence of the fenestra cannot be determined. However, it is possible that at least some of these differences between the
<specimenCount box="[246,398,1382,1403]" pageId="5" pageNumber="5" type="generic">two specimens</specimenCount>
are due to a combination of individual variation, taphonomical deformation (also rendering some elements difficult to segment precisely), and perhaps sexual dimorphism (comparing a male of one species with a female of another could exaggerate interspecific differences like the premaxillary crest height). With only a single specimen of each species, individual variation is impossible to assess.Also note that the skull of
<taxonomicName authorityName="Xing et al." box="[142,290,1586,1607]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[142,159,1586,1607]" italics="true" pageId="5" pageNumber="5">O</emphasis>
.
<emphasis box="[174,290,1586,1607]" italics="true" pageId="5" pageNumber="5">khaungraae</emphasis>
</taxonomicName>
was reported as measuring
<quantity box="[593,666,1586,1607]" metricMagnitude="-2" metricUnit="m" metricValue="1.4" pageId="5" pageNumber="5" unit="mm" value="14.0">14 mm</quantity>
in length,
<superScript attach="left" box="[770,779,1582,1596]" fontSize="6" pageId="5" pageNumber="5">1</superScript>
whereas our own measurement of the specimen gives a length of
<quantity box="[168,260,1645,1666]" metricMagnitude="-2" metricUnit="m" metricValue="1.73" pageId="5" pageNumber="5" unit="mm" value="17.3">17.3 mm</quantity>
.
</paragraph>
</subSubSection>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5014013" ID-Zenodo-Dep="5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="4" pageNumber="4" startId="4.[159,219,1918,1937]" targetBox="[163,1427,271,1897]" targetPageId="3">
<paragraph blockId="4.[159,1467,1918,1986]" box="[159,1049,1918,1937]" pageId="4" pageNumber="4">
<emphasis bold="true" box="[159,1049,1918,1937]" pageId="4" pageNumber="4">
Figure 2. Comparison of the two specimens of
<taxonomicName box="[609,746,1918,1937]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis bold="true" box="[609,746,1918,1937]" italics="true" pageId="4" pageNumber="4">Oculudentavis</emphasis>
</taxonomicName>
, each bone digitally segmented
</emphasis>
</paragraph>
<paragraph blockId="4.[159,1467,1918,1986]" pageId="4" pageNumber="4">
Synchrotron HRCT of
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[340,403,1944,1961]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="naga">
<emphasis box="[340,403,1944,1961]" italics="true" pageId="4" pageNumber="4">O.naga</emphasis>
</taxonomicName>
(AJ) and
<taxonomicName authorityName="Xing et al." box="[488,608,1944,1961]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[488,502,1944,1961]" italics="true" pageId="4" pageNumber="4">O</emphasis>
.
<emphasis box="[512,608,1944,1961]" italics="true" pageId="4" pageNumber="4">khaungraae</emphasis>
</taxonomicName>
(KT).(Aand K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1.
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5013959" ID-Zenodo-Dep="5013959" httpUri="https://zenodo.org/record/5013959/files/figure.png" pageId="5" pageNumber="5" startId="5.[1038,1098,271,290]" targetBox="[142,1003,271,943]" targetPageId="5">
<paragraph blockId="5.[1038,1449,271,516]" pageId="5" pageNumber="5">
<emphasis bold="true" pageId="5" pageNumber="5">
Figure 3. Comparison of the two specimens of
<taxonomicName box="[1062,1199,296,315]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis bold="true" box="[1062,1199,296,315]" italics="true" pageId="5" pageNumber="5">Oculudentavis</emphasis>
</taxonomicName>
, details of the lacrimal and dentary
</emphasis>
</paragraph>
<paragraph blockId="5.[1038,1449,271,516]" pageId="5" pageNumber="5">
Right side view of the orbit showing the ring-like lacrimal bone (salmon color)
<emphasis box="[1280,1329,373,390]" italics="true" pageId="5" pageNumber="5">in situ</emphasis>
in (A)
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[1383,1448,373,390]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1383,1397,373,390]" italics="true" pageId="5" pageNumber="5">O</emphasis>
.
<emphasis box="[1408,1448,373,390]" italics="true" pageId="5" pageNumber="5">naga</emphasis>
</taxonomicName>
and (B)
<taxonomicName authorityName="Xing et al." box="[1124,1255,398,415]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1124,1138,398,415]" italics="true" pageId="5" pageNumber="5">O</emphasis>
.
<emphasis box="[1159,1255,398,415]" italics="true" pageId="5" pageNumber="5">khaungraae</emphasis>
</taxonomicName>
(orbit view here is posterolateral to show the orbital elements more clearly). Dorsolingual view of the dentary in (C)
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1429,1443,449,466]" italics="true" pageId="5" pageNumber="5">O</emphasis>
.
<emphasis box="[1038,1078,474,491]" italics="true" pageId="5" pageNumber="5">naga</emphasis>
</taxonomicName>
and (D)
<taxonomicName authorityName="Xing et al." box="[1153,1275,474,491]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1153,1167,474,491]" italics="true" pageId="5" pageNumber="5">O</emphasis>
.
<emphasis box="[1179,1275,474,491]" italics="true" pageId="5" pageNumber="5">khaungraae</emphasis>
</taxonomicName>
is shown. Scale bar represents 1 mm.
</paragraph>
</caption>
<subSubSection lastPageId="8" lastPageNumber="8" pageId="5" pageNumber="5" type="description">
<paragraph blockId="5.[142,781,1031,1987]" box="[142,272,1674,1695]" pageId="5" pageNumber="5">
<heading box="[142,272,1674,1695]" pageId="5" pageNumber="5">
<emphasis bold="true" box="[142,272,1674,1695]" italics="true" pageId="5" pageNumber="5">Description</emphasis>
</heading>
</paragraph>
<paragraph blockId="5.[142,781,1031,1987]" pageId="5" pageNumber="5">
In its bird-like shape (vaulted cranium and tapering rostrum), the skull of
<taxonomicName authorityName="Xing et al." box="[226,495,1733,1754]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[226,495,1733,1754]" italics="true" pageId="5" pageNumber="5">Oculudentavis khaungraae</emphasis>
</taxonomicName>
appears strikingly different from that of any known lizard (
<figureCitation box="[435,514,1762,1783]" captionStart="Figure 4" captionStartId="6.[159,219,657,676]" captionTargetBox="[175,1450,272,617]" captionTargetId="figure-807@6.[175,1450,271,617]" captionTargetPageId="6" captionText="Figure 4. Cranial disparity of typical lepidosaurs to demonstrate the atypical skulls of Oculudentavis (A) Sphenodon punctatus, Rhynchocephalia UF11978;(B) Anelytropsis papillosus UF-H-86708,Dibamidae;(C) Sphaerodactylus caicosensis UF95971,Gekkota; (D) Smaug swazicus NMB-R9201, Cordyliformes; (E) Eugongylus albofasciolatus CAS159825, Scincidae; (F) Varanus sp. UF71411, Varanidae; (G) Rieppeleon brevicaudatus CAS168891, Iguania;(H) Boaedon fuliginosus CAS85747, Serpentes;(I) Oculudentavis naga GRS-Ref-28627;(J) O. khaungraae HPG-15-3.Scale bar represents 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5013961" httpUri="https://zenodo.org/record/5013961/files/figure.png" pageId="5" pageNumber="5">Figure 4</figureCitation>
). The bird-like appearance is less striking in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[310,388,1791,1812]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[310,388,1791,1812]" italics="true" pageId="5" pageNumber="5">O. naga</emphasis>
</taxonomicName>
, which has a less compressed rostrum (
<figureCitation box="[148,229,1820,1841]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="5" pageNumber="5">Figure 2</figureCitation>
).
</paragraph>
<paragraph blockId="5.[142,781,1031,1987]" lastBlockId="5.[811,1450,622,1987]" pageId="5" pageNumber="5">
Despite thecompression of the rostrum,the two speciesshare many characters that distinguish them from other lizards. The nares are bounded by the premaxilla anterodorsally, the maxilla posteroventrally, and by the nasals posteriorly. The location of the nares is also the same, being placed at mid-length of the antorbital region and in having an elongated oval shape. The orbit is more intact in
<taxonomicName authorityName="Xing et al." box="[1061,1206,680,701]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1061,1206,680,701]" italics="true" pageId="5" pageNumber="5">O. khaungraae</emphasis>
</taxonomicName>
, being nearly circular. In both species, the longest axis of the orbit is about 1/3 the total length of the skull and the orbit is complete and separated from the temporal fenestrae by a complete postorbital bar. The parietal supratemporal processes are aligned with the long and slender vertical supratemporals and fail to meet the squamosals ventrally. There is a complete circumorbital series in both specimens—jugal (ventrally), lacrimal (anteriorly), prefrontal (anterodorsally), frontal (dorsal), postfrontal (posterodorsally), and postorbital (posteriorly).
</paragraph>
<paragraph blockId="5.[811,1450,622,1987]" pageId="5" pageNumber="5">
<emphasis box="[834,937,1089,1110]" italics="true" pageId="5" pageNumber="5">Premaxilla</emphasis>
(
<figureCitation box="[948,1291,1089,1110]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="5" pageNumber="5">
Figures 2, S2AS2B, S
<date box="[1178,1198,1089,1110]" pageId="5" pageNumber="5">2I</date>
, and S2J
</figureCitation>
). The upper jaw comprises an unpaired median premaxilla with slender, pointed teeth (
<quantity box="[873,907,1148,1169]" metricMagnitude="-1" metricUnit="m" metricValue="2.286" pageId="5" pageNumber="5" unit="in" value="9.0">9 in</quantity>
<taxonomicName authorityName="Xing et al." box="[912,1056,1148,1169]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[912,1056,1148,1169]" italics="true" pageId="5" pageNumber="5">O. khaungraae</emphasis>
</taxonomicName>
and
<emphasis bold="true" box="[1104,1121,1149,1169]" pageId="5" pageNumber="5">~</emphasis>
<quantity box="[1122,1169,1148,1169]" metricMagnitude="-1" metricUnit="m" metricValue="2.54" pageId="5" pageNumber="5" unit="in" value="10.0">10 in</quantity>
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[1174,1251,1148,1169]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1174,1251,1148,1169]" italics="true" pageId="5" pageNumber="5">O. naga</emphasis>
</taxonomicName>
; count refers to one side of the element). The more anterior premaxillary teeth appear recurved in
<taxonomicName authorityName="Xing et al." box="[933,1080,1206,1227]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[933,1080,1206,1227]" italics="true" pageId="5" pageNumber="5">O. khaungraae</emphasis>
</taxonomicName>
, but the equivalent teeth in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[1369,1449,1206,1227]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1369,1449,1206,1227]" italics="true" pageId="5" pageNumber="5">O. naga</emphasis>
</taxonomicName>
are partially obscured. Both species have a long crest along the premaxillary nasal process (the crest was considered taphonomic in
<taxonomicName authorityName="Xing et al." box="[902,1048,1294,1315]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[902,1048,1294,1315]" italics="true" pageId="5" pageNumber="5">O. khaungraae</emphasis>
</taxonomicName>
),
<superScript attach="left" box="[1060,1069,1289,1303]" fontSize="6" pageId="5" pageNumber="5">1</superScript>
which continues onto the nasals. The palatal shelf is broad and flat and has two narrow palatal processes that bound a large premaxilla-vomer fenestra in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[811,893,1382,1403]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[811,893,1382,1403]" italics="true" pageId="5" pageNumber="5">O. naga</emphasis>
</taxonomicName>
, which has a less deformed palate. The palatal processes of the premaxilla are also visible in
<taxonomicName authorityName="Xing et al." box="[1254,1401,1411,1432]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1254,1401,1411,1432]" italics="true" pageId="5" pageNumber="5">O. khaungraae</emphasis>
</taxonomicName>
, but not the intervening fenestra (see below).
</paragraph>
<paragraph blockId="5.[811,1450,622,1987]" pageId="5" pageNumber="5">
<emphasis box="[834,903,1469,1490]" italics="true" pageId="5" pageNumber="5">Maxilla</emphasis>
(
<figureCitation box="[917,1287,1469,1490]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="5" pageNumber="5">Figures 2, S2CS2E, and S2KS2M</figureCitation>
). The maxilla of both species has a low, medially curved facial process, a long rostral component, and a suborbital ramus that does not reach the posterior margin of the orbit—extending up to one-quarter of the orbit length in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[1062,1149,1586,1607]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1062,1149,1586,1607]" italics="true" pageId="5" pageNumber="5">O. naga</emphasis>
</taxonomicName>
and to about mid-orbit in
<taxonomicName authorityName="Xing et al." box="[811,959,1616,1637]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[811,959,1616,1637]" italics="true" pageId="5" pageNumber="5">O. khaungraae</emphasis>
</taxonomicName>
. It is excluded from the orbital rim by the jugal. The maxillary teeth are conical and pointed. There are 24 to 25 tooth maxillary loci in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[1031,1110,1674,1695]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1031,1110,1674,1695]" italics="true" pageId="5" pageNumber="5">O. naga</emphasis>
</taxonomicName>
and
<quantity box="[1161,1245,1674,1695]" metricMagnitude="-1" metricUnit="m" metricValue="7.112" metricValueMax="7.366" metricValueMin="6.858" pageId="5" pageNumber="5" unit="in" value="28.0" valueMax="29.0" valueMin="27.0">2729 in</quantity>
<taxonomicName authorityName="Xing et al." box="[1252,1398,1674,1695]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1252,1398,1674,1695]" italics="true" pageId="5" pageNumber="5">O. khaungraae</emphasis>
</taxonomicName>
. The maxilla has two horizontal facets: one to support the prefrontal and another for the jugal.
</paragraph>
<paragraph blockId="5.[811,1450,622,1987]" pageId="5" pageNumber="5">
<emphasis box="[834,889,1762,1783]" italics="true" pageId="5" pageNumber="5">Nasal</emphasis>
(
<figureCitation box="[905,1167,1762,1783]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="5" pageNumber="5">Figures 2, S2F, and S2N</figureCitation>
). The paired nasals form a rhomboid plate, and combined with the maxilla, they define a long tapering rostrum with retracted narial openings. The nasals are paired, but they exhibit partial fusion along the crest and remain separated posterior to the crest.
</paragraph>
<paragraph blockId="5.[811,1450,622,1987]" lastBlockId="6.[159,799,827,1987]" lastPageId="6" lastPageNumber="6" pageId="5" pageNumber="5">
<emphasis box="[834,930,1908,1929]" italics="true" pageId="5" pageNumber="5">Prefrontal</emphasis>
(
<figureCitation box="[938,1015,1908,1929]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="5" pageNumber="5">Figure 2</figureCitation>
).The prefrontalscomprisea flatanterodorsal plate and a weakly concave orbital plate, contacting the ringshaped lacrimal ventrally. The anterodorsal plate seems less developed in
<taxonomicName authorityName="Xing et al." box="[292,435,827,848]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[292,435,827,848]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
than
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[489,566,827,848]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="naga">
<emphasis box="[489,566,827,848]" italics="true" pageId="6" pageNumber="6">O. naga</emphasis>
</taxonomicName>
. The lateral edge of the anterodorsal plate projects as a short angular (
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[622,700,856,877]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="naga">
<emphasis box="[622,700,856,877]" italics="true" pageId="6" pageNumber="6">O. naga</emphasis>
</taxonomicName>
) or ridgelike (
<taxonomicName authorityName="Xing et al." box="[203,346,885,906]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[203,346,885,906]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
) shelf that overhangs the lacrimal and maxilla. This shelf is autapomorphic among lizards, with a ridge, crest, or boss in this position variably present (e.g., some iguanians, including chameleons; some
<taxonomicName authorityName="Wiegmann" authorityYear="1828" box="[446,569,973,994]" class="Reptilia" family="Phrynosomatidae" genus="Phrynosoma" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[446,569,973,994]" italics="true" pageId="6" pageNumber="6">Phrynosoma</emphasis>
</taxonomicName>
; and some
<taxonomicName authorityName="Daudin" authorityYear="1802" box="[684,744,973,994]" class="Reptilia" family="Dactyloidae" genus="Anolis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis box="[684,744,973,994]" italics="true" pageId="6" pageNumber="6">Anolis</emphasis>
</taxonomicName>
).
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5013961" ID-Zenodo-Dep="5013961" httpUri="https://zenodo.org/record/5013961/files/figure.png" pageId="6" pageNumber="6" startId="6.[159,219,657,676]" targetBox="[175,1450,272,617]" targetPageId="6">
<paragraph blockId="6.[159,1468,657,776]" box="[159,1113,657,676]" pageId="6" pageNumber="6">
Figure 4. Cranial disparity of typical lepidosaurs to demonstrate the atypical skulls of
<taxonomicName box="[976,1113,657,676]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="23" phylum="Chordata" rank="genus">Oculudentavis</taxonomicName>
</paragraph>
<paragraph blockId="6.[159,1468,657,776]" pageId="6" pageNumber="6">
(A)
<taxonomicName baseAuthorityName="Gray" baseAuthorityYear="1842" box="[185,367,683,700]" class="Reptilia" family="Sphenodontidae" genus="Sphenodon" kingdom="Animalia" order="Rhynchocephalia" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="punctatus">
<emphasis box="[185,367,683,700]" italics="true" pageId="6" pageNumber="6">Sphenodon punctatus</emphasis>
</taxonomicName>
,
<taxonomicName authority="UF" authorityName="UF" box="[376,549,683,700]" class="Reptilia" kingdom="Animalia" order="Rhynchocephalia" pageId="6" pageNumber="6" phylum="Chordata" rank="order">Rhynchocephalia UF</taxonomicName>
11978;(B)
<taxonomicName authorityName="Cope" authorityYear="1885" box="[635,823,683,700]" class="Reptilia" family="Dibamidae" genus="Anelytropsis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="papillosus">
<emphasis box="[635,823,683,700]" italics="true" pageId="6" pageNumber="6">Anelytropsis papillosus</emphasis>
</taxonomicName>
UF-H-86708,
<taxonomicName authorityName="Boulenger" authorityYear="1884" box="[939,1032,683,700]" class="Reptilia" family="Dibamidae" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="family">Dibamidae</taxonomicName>
;(C)
<taxonomicName authority="UF" authorityName="UF" box="[1064,1332,683,700]" class="Reptilia" family="Sphaerodactylidae" genus="Sphaerodactylus" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="caicosensis">
<emphasis box="[1064,1303,683,700]" italics="true" pageId="6" pageNumber="6">Sphaerodactylus caicosensis</emphasis>
UF
</taxonomicName>
95971,Gekkota; (D)
<taxonomicName authorityName="Bates &amp; Stanley" authorityYear="2020" box="[188,326,708,725]" class="Reptilia" family="Cordylidae" genus="Smaug" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="swazicus">
<emphasis box="[188,326,708,725]" italics="true" pageId="6" pageNumber="6">Smaug swazicus</emphasis>
</taxonomicName>
NMB-R9201, Cordyliformes; (E)
<taxonomicName authority="CAS" authorityName="CAS" box="[600,870,708,725]" class="Reptilia" family="Scincidae" genus="Eugongylus" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="albofasciolatus">
<emphasis box="[600,825,708,725]" italics="true" pageId="6" pageNumber="6">Eugongylus albofasciolatus</emphasis>
CAS
</taxonomicName>
159825,
<taxonomicName authorityName="Oppel" authorityYear="1811" box="[941,1026,708,725]" class="Reptilia" family="Scincidae" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="family">Scincidae</taxonomicName>
; (F)
<taxonomicName box="[1059,1158,708,725]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="undetermined">
<emphasis box="[1059,1125,708,725]" italics="true" pageId="6" pageNumber="6">Varanus</emphasis>
sp.
</taxonomicName>
UF71411,
<taxonomicName authorityName="Merrem" authorityYear="1820" box="[1249,1335,708,725]" class="Reptilia" family="Varanidae" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="family">Varanidae</taxonomicName>
; (G)
<taxonomicName authority="CAS" authorityName="CAS" class="Reptilia" family="Chamaeleonidae" genus="Rieppeleon" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="brevicaudatus">
<emphasis italics="true" pageId="6" pageNumber="6">Rieppeleon brevicaudatus</emphasis>
CAS
</taxonomicName>
168891, Iguania;(H)
<taxonomicName authority="CAS" authorityName="CAS" box="[486,696,734,751]" class="Reptilia" family="Lamprophiidae" genus="Boaedon" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="fuliginosus">
<emphasis box="[486,652,734,751]" italics="true" pageId="6" pageNumber="6">Boaedon fuliginosus</emphasis>
CAS
</taxonomicName>
85747, Serpentes;(I)
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[869,1031,734,751]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="naga">
<emphasis box="[869,1031,734,751]" italics="true" pageId="6" pageNumber="6">Oculudentavis naga</emphasis>
</taxonomicName>
GRS-Ref-28627;(J)
<taxonomicName authorityName="Xing et al." box="[1202,1322,734,751]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1202,1216,734,751]" italics="true" pageId="6" pageNumber="6">O</emphasis>
.
<emphasis box="[1226,1322,734,751]" italics="true" pageId="6" pageNumber="6">khaungraae</emphasis>
</taxonomicName>
HPG-15-3.Scale bar represents 10 mm.
</paragraph>
</caption>
<paragraph blockId="6.[159,799,827,1987]" pageId="6" pageNumber="6">
<emphasis box="[182,267,1002,1023]" italics="true" pageId="6" pageNumber="6">Lacrimal</emphasis>
(
<figureCitation box="[280,470,1002,1023]" captionStart="Figure 3" captionStartId="5.[1038,1098,271,290]" captionTargetBox="[142,1003,271,943]" captionTargetId="figure-327@5.[142,1003,271,943]" captionTargetPageId="5" captionText="Figure 3. Comparison of the two specimens of Oculudentavis, details of the lacrimal and dentary Right side view of the orbit showing the ring-like lacrimal bone (salmon color) in situ in (A) O. naga and (B) O. khaungraae (orbit view here is posterolateral to show the orbital elements more clearly). Dorsolingual view of the dentary in (C) O. naga and (D) O. khaungraae is shown. Scale bar represents 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5013959" httpUri="https://zenodo.org/record/5013959/files/figure.png" pageId="6" pageNumber="6">Figures 3A and 3B</figureCitation>
). The lacrimal of both species is unique among lizards and is one of several distinctive features that demonstrates their close relationship. It forms a ring, completely enclosing a large lacrimal foramen.
</paragraph>
<paragraph blockId="6.[159,799,827,1987]" pageId="6" pageNumber="6">
<emphasis box="[182,236,1119,1140]" italics="true" pageId="6" pageNumber="6">Jugal</emphasis>
(
<figureCitation box="[249,618,1119,1140]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="6" pageNumber="6">Figures 2, S2G, S2H, S2O, and S2P</figureCitation>
). In both species, the jugal forms a dorsomedially expanded flange that provides ventral support to the large eye. The orientation of the jugal is unusual for squamates, being dorsomedially inclined. The postorbital process of the jugal is short (distorted on the right side of
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[159,238,1265,1286]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="naga">
<emphasis box="[159,238,1265,1286]" italics="true" pageId="6" pageNumber="6">O. naga</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph blockId="6.[159,799,827,1987]" pageId="6" pageNumber="6">
<emphasis box="[182,252,1294,1315]" italics="true" pageId="6" pageNumber="6">Frontal</emphasis>
(
<figureCitation box="[267,363,1294,1315]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="6" pageNumber="6">Figures 2</figureCitation>
, S
<figureCitation box="[393,528,1294,1315]" captionStart="Figure 3" captionStartId="5.[1038,1098,271,290]" captionTargetBox="[142,1003,271,943]" captionTargetId="figure-327@5.[142,1003,271,943]" captionTargetPageId="5" captionText="Figure 3. Comparison of the two specimens of Oculudentavis, details of the lacrimal and dentary Right side view of the orbit showing the ring-like lacrimal bone (salmon color) in situ in (A) O. naga and (B) O. khaungraae (orbit view here is posterolateral to show the orbital elements more clearly). Dorsolingual view of the dentary in (C) O. naga and (D) O. khaungraae is shown. Scale bar represents 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5013959" httpUri="https://zenodo.org/record/5013959/files/figure.png" pageId="6" pageNumber="6">3A, and S3E</figureCitation>
). In both species, the unpaired median frontal has weak sub-olfactory processes and a deep V-shaped anterior emargination that receives the nasals. The frontal is overlapped extensively by the nasals, reaching the level of the mid-orbit in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[467,551,1411,1432]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="naga">
<emphasis box="[467,551,1411,1432]" italics="true" pageId="6" pageNumber="6">O. naga</emphasis>
</taxonomicName>
and somewhat less in
<taxonomicName authorityName="Xing et al." box="[159,309,1440,1461]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[159,309,1440,1461]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
. The supraorbital margins are subparallel and diverge posterolaterally, establishing a broad contact with the anterior margin of the parietal. The structure of the posteromedial margin of the fronto-parietal suture is unclear in both specimens (
<figureCitation box="[227,308,1557,1578]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="6" pageNumber="6">Figure 2</figureCitation>
, dashed lines).
</paragraph>
<paragraph blockId="6.[159,799,827,1987]" pageId="6" pageNumber="6">
<emphasis box="[182,257,1586,1607]" italics="true" pageId="6" pageNumber="6">Parietal</emphasis>
(
<figureCitation box="[268,360,1586,1607]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="6" pageNumber="6">Figures 2</figureCitation>
, S
<figureCitation box="[388,515,1586,1607]" captionStart="Figure 3" captionStartId="5.[1038,1098,271,290]" captionTargetBox="[142,1003,271,943]" captionTargetId="figure-327@5.[142,1003,271,943]" captionTargetPageId="5" captionText="Figure 3. Comparison of the two specimens of Oculudentavis, details of the lacrimal and dentary Right side view of the orbit showing the ring-like lacrimal bone (salmon color) in situ in (A) O. naga and (B) O. khaungraae (orbit view here is posterolateral to show the orbital elements more clearly). Dorsolingual view of the dentary in (C) O. naga and (D) O. khaungraae is shown. Scale bar represents 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5013959" httpUri="https://zenodo.org/record/5013959/files/figure.png" pageId="6" pageNumber="6">3B, and S3F</figureCitation>
). The parietals are short and partially fused (separated posteriorly). They have a rounded lateral profile, lack a parietal foramen, and have short supratemporal processes that curve ventrally rather than posteriorly to meet the supratemporals. This portion of the skull contacts the short paroccipital processes of the otoccipital. Li et al.
<superScript attach="left" box="[710,719,1728,1742]" fontSize="6" pageId="6" pageNumber="6">3</superScript>
argued that the small opening in the midline of the parietals in the
<typeStatus pageId="6" pageNumber="6">holotype</typeStatus>
of
<taxonomicName authorityName="Xing et al." box="[232,376,1791,1812]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[232,376,1791,1812]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
corresponds to a parietal foramen, but it is irregular and appears to be an artifact of breakage.
</paragraph>
<paragraph blockId="6.[159,799,827,1987]" pageId="6" pageNumber="6">
<emphasis box="[182,291,1849,1870]" italics="true" pageId="6" pageNumber="6">Postfrontal</emphasis>
(
<figureCitation box="[307,392,1849,1870]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="6" pageNumber="6">Figure 2</figureCitation>
). The postfrontal is a small, splint-like bone, lateral to the frontal and the parietal in
<taxonomicName authorityName="Xing et al." box="[647,797,1879,1900]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[647,797,1879,1900]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
but of uncertain structure and position in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[571,649,1908,1929]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="naga">
<emphasis box="[571,649,1908,1929]" italics="true" pageId="6" pageNumber="6">O. naga</emphasis>
</taxonomicName>
. The postfrontals are very reduced in both species and were not noticed in the original description of
<taxonomicName authorityName="Xing et al." box="[383,529,1966,1987]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[383,529,1966,1987]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="6.[829,1467,827,1987]" pageId="6" pageNumber="6">
<emphasis box="[851,959,827,848]" italics="true" pageId="6" pageNumber="6">Postorbital</emphasis>
(
<figureCitation box="[975,1072,827,848]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="6" pageNumber="6">Figures 2</figureCitation>
, S
<figureCitation box="[1103,1243,827,848]" captionStart="Figure 3" captionStartId="5.[1038,1098,271,290]" captionTargetBox="[142,1003,271,943]" captionTargetId="figure-327@5.[142,1003,271,943]" captionTargetPageId="5" captionText="Figure 3. Comparison of the two specimens of Oculudentavis, details of the lacrimal and dentary Right side view of the orbit showing the ring-like lacrimal bone (salmon color) in situ in (A) O. naga and (B) O. khaungraae (orbit view here is posterolateral to show the orbital elements more clearly). Dorsolingual view of the dentary in (C) O. naga and (D) O. khaungraae is shown. Scale bar represents 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5013959" httpUri="https://zenodo.org/record/5013959/files/figure.png" pageId="6" pageNumber="6">3C, and S3G</figureCitation>
). The postorbital is a strongly triradiate bone with a long (
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[1262,1352,856,877]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1262,1352,856,877]" italics="true" pageId="6" pageNumber="6">O. naga</emphasis>
</taxonomicName>
) or short (
<taxonomicName authorityName="Xing et al." box="[835,981,885,906]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[835,981,885,906]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
) posterior process that contacts the squamosal posteriorly. The postorbital differs in the two species: the postorbital squamosal process tapers gradually in the
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[1325,1406,943,964]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1325,1406,943,964]" italics="true" pageId="6" pageNumber="6">O. naga</emphasis>
</taxonomicName>
<typeStatus pageId="6" pageNumber="6">holotype</typeStatus>
, while the tapering appears more abrupt in the
<taxonomicName authorityName="Xing et al." box="[829,974,1002,1023]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[829,974,1002,1023]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
specimen. Due to the proportionally thicker postorbital, the right side of
<taxonomicName authorityName="Xing et al." box="[1069,1215,1031,1052]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1069,1215,1031,1052]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
shows a more extensive contact between the postorbital and the descending process of the parietal, entirely covering the braincase laterally and almost completely closing the upper temporal fenestra. However, on the left side, it is clear that this fenestra remained open. In
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[829,908,1177,1198]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="naga">
<emphasis box="[829,908,1177,1198]" italics="true" pageId="6" pageNumber="6">O. naga</emphasis>
</taxonomicName>
, the upper temporal fenestra looks larger, but the skull table of this specimen is very depressed and the postorbital is more gracile, so the differences in configuration of the upper temporal bar may be exaggerated by taphonomic deformation.
</paragraph>
<paragraph blockId="6.[829,1467,827,1987]" pageId="6" pageNumber="6">
<emphasis box="[851,963,1323,1344]" italics="true" pageId="6" pageNumber="6">Squamosal</emphasis>
(
<figureCitation box="[978,1074,1323,1344]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="6" pageNumber="6">Figures 2</figureCitation>
, S
<figureCitation box="[1104,1241,1323,1344]" captionStart="Figure 3" captionStartId="5.[1038,1098,271,290]" captionTargetBox="[142,1003,271,943]" captionTargetId="figure-327@5.[142,1003,271,943]" captionTargetPageId="5" captionText="Figure 3. Comparison of the two specimens of Oculudentavis, details of the lacrimal and dentary Right side view of the orbit showing the ring-like lacrimal bone (salmon color) in situ in (A) O. naga and (B) O. khaungraae (orbit view here is posterolateral to show the orbital elements more clearly). Dorsolingual view of the dentary in (C) O. naga and (D) O. khaungraae is shown. Scale bar represents 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5013959" httpUri="https://zenodo.org/record/5013959/files/figure.png" pageId="6" pageNumber="6">3D, and S3H</figureCitation>
). In both species, the typically squamate hockey-stick-shaped squamosal lacks an ascending process and lies between the supratemporal, the postorbital, and the quadrate.
</paragraph>
<paragraph blockId="6.[829,1467,827,1987]" pageId="6" pageNumber="6">
<emphasis box="[851,1000,1440,1461]" italics="true" pageId="6" pageNumber="6">Supratemporal</emphasis>
(
<figureCitation box="[1013,1106,1440,1461]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="6" pageNumber="6">Figures 2</figureCitation>
, S
<figureCitation box="[1134,1264,1440,1461]" captionStart="Figure 3" captionStartId="5.[1038,1098,271,290]" captionTargetBox="[142,1003,271,943]" captionTargetId="figure-327@5.[142,1003,271,943]" captionTargetPageId="5" captionText="Figure 3. Comparison of the two specimens of Oculudentavis, details of the lacrimal and dentary Right side view of the orbit showing the ring-like lacrimal bone (salmon color) in situ in (A) O. naga and (B) O. khaungraae (orbit view here is posterolateral to show the orbital elements more clearly). Dorsolingual view of the dentary in (C) O. naga and (D) O. khaungraae is shown. Scale bar represents 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5013959" httpUri="https://zenodo.org/record/5013959/files/figure.png" pageId="6" pageNumber="6">3B, and S3F</figureCitation>
, articulated with the parietal). The supratemporal is also reduced to a slender vertical splint of bone that contacts the lateral margin of the parietal supratemporal process, separating it from the squamosal.
</paragraph>
<paragraph blockId="6.[829,1467,827,1987]" lastBlockId="7.[142,780,271,1987]" lastPageId="7" lastPageNumber="7" pageId="6" pageNumber="6">
<emphasis box="[851,913,1557,1578]" italics="true" pageId="6" pageNumber="6">Palate</emphasis>
(
<figureCitation box="[940,1046,1557,1578]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="6" pageNumber="6">Figures 2</figureCitation>
, S
<figureCitation box="[1088,1245,1557,1578]" captionStart="Figure 4" captionStartId="6.[159,219,657,676]" captionTargetBox="[175,1450,272,617]" captionTargetId="figure-807@6.[175,1450,271,617]" captionTargetPageId="6" captionText="Figure 4. Cranial disparity of typical lepidosaurs to demonstrate the atypical skulls of Oculudentavis (A) Sphenodon punctatus, Rhynchocephalia UF11978;(B) Anelytropsis papillosus UF-H-86708,Dibamidae;(C) Sphaerodactylus caicosensis UF95971,Gekkota; (D) Smaug swazicus NMB-R9201, Cordyliformes; (E) Eugongylus albofasciolatus CAS159825, Scincidae; (F) Varanus sp. UF71411, Varanidae; (G) Rieppeleon brevicaudatus CAS168891, Iguania;(H) Boaedon fuliginosus CAS85747, Serpentes;(I) Oculudentavis naga GRS-Ref-28627;(J) O. khaungraae HPG-15-3.Scale bar represents 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5013961" httpUri="https://zenodo.org/record/5013961/files/figure.png" pageId="6" pageNumber="6">4A, and S4D</figureCitation>
). In the palate of
<taxonomicName authorityName="Xing et al." box="[829,976,1586,1607]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[829,976,1586,1607]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
, the premaxilla-vomer fenestra is totally obliterated (due to compression). In this respect,
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[1266,1345,1616,1637]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1266,1345,1616,1637]" italics="true" pageId="6" pageNumber="6">O. naga</emphasis>
</taxonomicName>
has a more intact rostrum, more clearly exhibiting thepremaxilla-vomer fenestra and the very large fenestra exochoanalis. The suborbital fenestra is oval in both specimens and is bounded by the same bones: palatines anteromedially, ectopterygoids laterally, and pterygoids posteriorly, although the sutures between these bones are not easy to identify. It also looks as if the ectopterygoid barely contacts the palatine in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[1129,1206,1820,1841]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1129,1206,1820,1841]" italics="true" pageId="6" pageNumber="6">O. naga</emphasis>
</taxonomicName>
, but the degree of contact is ambiguous in
<taxonomicName authorityName="Xing et al." box="[993,1139,1849,1870]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[993,1139,1849,1870]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
. The shape of the interpterygoid vacuity differs between the two species. Pterygoid teeth are present and are arranged in a row on the anteromedial process of the pterygoid, just posterior to the inferred suture with the palatine. There are about 3 to 4 on each bone in
<taxonomicName authorityName="Xing et al." box="[1276,1422,1966,1987]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1276,1422,1966,1987]" italics="true" pageId="6" pageNumber="6">O. khaungraae</emphasis>
</taxonomicName>
; the same area is fragmented in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[434,515,271,292]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="naga">
<emphasis box="[434,515,271,292]" italics="true" pageId="7" pageNumber="7">O. naga</emphasis>
</taxonomicName>
, but small projections on both pterygoids can be interpreted as pterygoid teeth.
</paragraph>
<paragraph blockId="7.[142,780,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[164,257,330,351]" italics="true" pageId="7" pageNumber="7">Quadrate</emphasis>
(
<figureCitation box="[270,364,330,351]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="7" pageNumber="7">Figures 2</figureCitation>
, S
<figureCitation box="[392,523,330,351]" captionStart="Figure 4" captionStartId="6.[159,219,657,676]" captionTargetBox="[175,1450,272,617]" captionTargetId="figure-807@6.[175,1450,271,617]" captionTargetPageId="6" captionText="Figure 4. Cranial disparity of typical lepidosaurs to demonstrate the atypical skulls of Oculudentavis (A) Sphenodon punctatus, Rhynchocephalia UF11978;(B) Anelytropsis papillosus UF-H-86708,Dibamidae;(C) Sphaerodactylus caicosensis UF95971,Gekkota; (D) Smaug swazicus NMB-R9201, Cordyliformes; (E) Eugongylus albofasciolatus CAS159825, Scincidae; (F) Varanus sp. UF71411, Varanidae; (G) Rieppeleon brevicaudatus CAS168891, Iguania;(H) Boaedon fuliginosus CAS85747, Serpentes;(I) Oculudentavis naga GRS-Ref-28627;(J) O. khaungraae HPG-15-3.Scale bar represents 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5013961" httpUri="https://zenodo.org/record/5013961/files/figure.png" pageId="7" pageNumber="7">4B, and S4E</figureCitation>
). The quadrate is distinctively low in position and small in size in both species. The quadrate is stouter in
<taxonomicName authorityName="Xing et al." box="[319,467,388,409]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[319,467,388,409]" italics="true" pageId="7" pageNumber="7">O. khaungraae</emphasis>
</taxonomicName>
(with a more prominent head) than in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[219,300,417,438]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="naga">
<emphasis box="[219,300,417,438]" italics="true" pageId="7" pageNumber="7">O. naga</emphasis>
</taxonomicName>
, but the overall shape is similar in both specimens, with a shallow conch, a slightly curved medial pillar, and a lateral tympanic crest that has a 90-degree angulation along its length. The quadrate suspension in both species is characteristically squamate.
</paragraph>
<paragraph blockId="7.[142,780,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[164,263,563,584]" italics="true" pageId="7" pageNumber="7">Braincase</emphasis>
(
<figureCitation box="[280,378,563,584]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="7" pageNumber="7">Figures 2</figureCitation>
and S
<figureCitation box="[453,465,563,584]" captionStart="Figure 5" captionStartId="8.[159,219,1316,1335]" captionTargetBox="[176,1449,273,1275]" captionTargetId="figure-278@8.[175,1450,271,1276]" captionTargetPageId="8" captionText="Figure 5. Phylogenetic trees showing the position of Oculudentavis (A) Phylogenetic tree showing the position of Oculudentavis using an amniote dataset;6 (B) photograph of the holotype of O. naga; (C) simplified phylogenetic tree showing the position of Oculudentavis using a squamate dataset7 combined with a molecular dataset15 treating some morphological characters as ordered; (D) simplified phylogenetic tree showing the position of Oculudentavis using the same dataset as in (C) but with all characters unordered;(E) simplified phylogenetic tree showing the position of Oculudentavis using the same dataset as (C) but removing molecular data. In (C)(E), crown groups were collapsed and are represented by silhouettes. Sphenodon punctatus, Anelytropsis papillosus (Dibamidae), Sphaerodactylus klauberi (Gekkota), Smaug giganteus (Cordylidae), Xantusia vigilis (Xantusiidae), Tribolonotus gracilis (Scincidae), Bachia flavescens (Lacertoidea), Lacerta bilineata (Lacertoidea), Blanus cinereous (Amphisbaenia), Varanus komodoensis (Anguimorpha), Physignathus cocincinus (Iguania), and Ophiophagus hannah (Serpentes) are shown. Node values indicate Bremer support; nodes with no support were recovered in the implied weights analyses. See also Data S1." figureDoi="http://doi.org/10.5281/zenodo.5013965" httpUri="https://zenodo.org/record/5013965/files/figure.png" pageId="7" pageNumber="7">5</figureCitation>
). By comparison with that of
<taxonomicName authorityName="Xing et al." box="[142,288,593,614]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[142,288,593,614]" italics="true" pageId="7" pageNumber="7">O. khaungraae</emphasis>
</taxonomicName>
, the braincase of
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[471,551,593,614]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="naga">
<emphasis box="[471,551,593,614]" italics="true" pageId="7" pageNumber="7">O. naga</emphasis>
</taxonomicName>
is unevenly dorsoventrally compressed, so that the right side is more damaged than the left and the posteroventral margin is abnormally low. Nonetheless, comparison of the two braincases shows more similarities than differences, notably the well-developed crista prootica, short alar processes, slender basipterygoid processes, short basisphenoid, enclosed vidian canals opening posteriorly within the basisphenoid, robust parasphenoid rostrum (base only preserved in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[240,318,826,847]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="naga">
<emphasis box="[240,318,826,847]" italics="true" pageId="7" pageNumber="7">O. naga</emphasis>
</taxonomicName>
), short uncrested supraoccipital with a visible processus ascendens (mineralization uncertain), and short paroccipital processes. The parasphenoid rostrum is well preserved in
<taxonomicName authorityName="Xing et al." box="[245,394,914,935]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[245,394,914,935]" italics="true" pageId="7" pageNumber="7">O. khaungraae</emphasis>
</taxonomicName>
, being longer than the basipterygoid processes, and almost entirely divides the interpterygoid vacuity. In
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[208,290,972,993]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="naga">
<emphasis box="[208,290,972,993]" italics="true" pageId="7" pageNumber="7">O. naga</emphasis>
</taxonomicName>
, the parasphenoid rostrum is represented only by its base, possibly due a fracture or weak mineralization. However, there are differences in the orientation, length, and distal shape of the basipterygoid processes in the two species.
</paragraph>
<paragraph blockId="7.[142,780,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[164,291,1089,1110]" italics="true" pageId="7" pageNumber="7">Epipterygoid</emphasis>
(
<figureCitation box="[304,385,1089,1110]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="7" pageNumber="7">Figure 2</figureCitation>
). These elements are poorly preserved and displaced in both species. They are columnar and still in articulation within the fossa columellae of the pterygoid, this articulation being another uniquely squamate character. In the
<typeStatus box="[142,229,1206,1227]" pageId="7" pageNumber="7">holotype</typeStatus>
of
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[264,345,1206,1227]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="naga">
<emphasis box="[264,345,1206,1227]" italics="true" pageId="7" pageNumber="7">O. naga</emphasis>
</taxonomicName>
, a portion of the left epipterygoid remains attached to the alar process.
</paragraph>
<paragraph blockId="7.[142,780,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[164,317,1265,1286]" italics="true" pageId="7" pageNumber="7">Scleral ossicles</emphasis>
(
<figureCitation box="[328,419,1265,1286]" captionStart="Figure 1" captionStartId="2.[1053,1113,271,290]" captionTargetBox="[159,1016,271,1338]" captionTargetId="figure-191@2.[159,1016,271,1338]" captionTargetPageId="2" captionText="Figure 1. Oculudentavis naga (GRS-Ref- 286278) displaying the superb preservation of bone and soft tissue (A) Dorsal; (B) lateral; (C) ventral views. Scale bar represents 10 mm. Diptera associated with the lizard skeleton were identified as Phoridae, Platypezidae, Ceratopogonidae, or Brachycera (Empidoidea). See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5013955" httpUri="https://zenodo.org/record/5013955/files/figure.png" pageId="7" pageNumber="7">Figures 1</figureCitation>
, S
<figureCitation box="[445,571,1265,1286]" captionStart="Figure 4" captionStartId="6.[159,219,657,676]" captionTargetBox="[175,1450,272,617]" captionTargetId="figure-807@6.[175,1450,271,617]" captionTargetPageId="6" captionText="Figure 4. Cranial disparity of typical lepidosaurs to demonstrate the atypical skulls of Oculudentavis (A) Sphenodon punctatus, Rhynchocephalia UF11978;(B) Anelytropsis papillosus UF-H-86708,Dibamidae;(C) Sphaerodactylus caicosensis UF95971,Gekkota; (D) Smaug swazicus NMB-R9201, Cordyliformes; (E) Eugongylus albofasciolatus CAS159825, Scincidae; (F) Varanus sp. UF71411, Varanidae; (G) Rieppeleon brevicaudatus CAS168891, Iguania;(H) Boaedon fuliginosus CAS85747, Serpentes;(I) Oculudentavis naga GRS-Ref-28627;(J) O. khaungraae HPG-15-3.Scale bar represents 10 mm." figureDoi="http://doi.org/10.5281/zenodo.5013961" httpUri="https://zenodo.org/record/5013961/files/figure.png" pageId="7" pageNumber="7">4C, and S4F</figureCitation>
). In both species, the orbit contains a large ring of spoon-shaped scleral ossicles that supported a large eye. The ossicle count is
<quantity box="[617,664,1323,1344]" metricMagnitude="-1" metricUnit="m" metricValue="3.556" pageId="7" pageNumber="7" unit="in" value="14.0">14 in</quantity>
both specimens. Due to the distinctive shape of the ossicles, they overlap at both their inner edges (which would have surrounded the iris and the pupil) and the outer edges, leaving oval gaps between ossicles in the middle of the sclerotic ring. Although the skull of
<taxonomicName authorityName="Xing et al." box="[142,286,1469,1490]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[142,286,1469,1490]" italics="true" pageId="7" pageNumber="7">O. khaungraae</emphasis>
</taxonomicName>
is 1.2X longer than that of
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[551,628,1469,1490]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="naga">
<emphasis box="[551,628,1469,1490]" italics="true" pageId="7" pageNumber="7">O. naga</emphasis>
</taxonomicName>
, the scleral ossicles are proportionally larger in
<emphasis box="[496,653,1499,1520]" italics="true" pageId="7" pageNumber="7">
<taxonomicName authorityName="Xing et al." box="[496,649,1499,1520]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="khaungraae">O. khaungraae</taxonomicName>
,
</emphasis>
being 1.5X larger than those of
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[344,423,1528,1549]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="naga">
<emphasis box="[344,423,1528,1549]" italics="true" pageId="7" pageNumber="7">O. naga</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="7.[142,780,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[164,242,1557,1578]" italics="true" pageId="7" pageNumber="7">Dentary</emphasis>
(
<figureCitation box="[254,345,1557,1578]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="7" pageNumber="7">Figures 2</figureCitation>
,
<figureCitation box="[358,430,1557,1578]" captionStart="Figure 3" captionStartId="5.[1038,1098,271,290]" captionTargetBox="[142,1003,271,943]" captionTargetId="figure-327@5.[142,1003,271,943]" captionTargetPageId="5" captionText="Figure 3. Comparison of the two specimens of Oculudentavis, details of the lacrimal and dentary Right side view of the orbit showing the ring-like lacrimal bone (salmon color) in situ in (A) O. naga and (B) O. khaungraae (orbit view here is posterolateral to show the orbital elements more clearly). Dorsolingual view of the dentary in (C) O. naga and (D) O. khaungraae is shown. Scale bar represents 1 mm." figureDoi="http://doi.org/10.5281/zenodo.5013959" httpUri="https://zenodo.org/record/5013959/files/figure.png" pageId="7" pageNumber="7">3C, 3D</figureCitation>
, and S
<figureCitation box="[497,509,1557,1578]" captionStart="Figure 6" captionStartId="9.[1038,1098,271,290]" captionTargetBox="[143,1003,271,671]" captionTargetId="figure-838@9.[142,1003,271,671]" captionTargetPageId="9" captionText="Figure 6. Heatmaps displaying the possible degree of deformation in Oculudentavis species O. khaungraae HPG-15-3 (A and B) and O. naga GRS-Ref-28627 (C and D). The shape variation (degree of deformation) between the original and retrodeformed 3D surface is color coded, with yellow and dark blue representing no deformation and high deformation, respectively. Heatmaps indicating the amount of retrodeformation provide a good visual aid to display the regions where a specimen would have been deformed and what its overall shape would have looked like prior to deformation. All changes are based on observed distortions in the specimens themselves. See also Data S1." figureDoi="http://doi.org/10.5281/zenodo.5013969" httpUri="https://zenodo.org/record/5013969/files/figure.png" pageId="7" pageNumber="7">6</figureCitation>
). Both species have a long shallow mandible of which the straight dentary forms the major part (
<emphasis bold="true" box="[197,214,1616,1636]" pageId="7" pageNumber="7">~</emphasis>
75%) and a large number of sharp, weakly pleurodont teeth (
<quantity box="[202,302,1645,1666]" metricMagnitude="-1" metricUnit="m" metricValue="7.493" metricValueMax="7.62" metricValueMin="7.366" pageId="7" pageNumber="7" unit="in" value="29.5" valueMax="30.0" valueMin="29.0">29 to 30 in</quantity>
both specimens). Both speciesalso have a large number of lateral neurovascular foramina (1012), and the dentary in each specimen has parallel upper and lower margins. Thesymphysealregiondoes not extendbeyondthe second tooth locus in either specimen. The lower margin of the dentary curves dorsomedially and closely approaches the subdental shelf, thus restricting the Meckelian fossa but without fusion.The Meckelian fossa remains open posteriorly, where it is overlapped by the splenial. The dorsolabial surface of the posterior one-third of the dentary bears a flattened, shelf-like surface.
</paragraph>
<paragraph blockId="7.[142,780,271,1987]" lastBlockId="7.[811,1450,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[164,243,1937,1958]" italics="true" pageId="7" pageNumber="7">Splenial</emphasis>
(
<figureCitation box="[257,352,1937,1958]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="7" pageNumber="7">Figures 2</figureCitation>
and S
<figureCitation box="[421,433,1937,1958]" captionStart="Figure 6" captionStartId="9.[1038,1098,271,290]" captionTargetBox="[143,1003,271,671]" captionTargetId="figure-838@9.[142,1003,271,671]" captionTargetPageId="9" captionText="Figure 6. Heatmaps displaying the possible degree of deformation in Oculudentavis species O. khaungraae HPG-15-3 (A and B) and O. naga GRS-Ref-28627 (C and D). The shape variation (degree of deformation) between the original and retrodeformed 3D surface is color coded, with yellow and dark blue representing no deformation and high deformation, respectively. Heatmaps indicating the amount of retrodeformation provide a good visual aid to display the regions where a specimen would have been deformed and what its overall shape would have looked like prior to deformation. All changes are based on observed distortions in the specimens themselves. See also Data S1." figureDoi="http://doi.org/10.5281/zenodo.5013969" httpUri="https://zenodo.org/record/5013969/files/figure.png" pageId="7" pageNumber="7">6</figureCitation>
). The splenial is very slender and does not extend anteriorly beyond the posterior one-third of the dentary, closing only the posterior part of the Meckelian fossa in both species. Posteriorly, the splenial does not extend beyond the level of the coronoid eminence.
</paragraph>
<paragraph blockId="7.[811,1450,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[834,927,359,380]" italics="true" pageId="7" pageNumber="7">Coronoid</emphasis>
(
<figureCitation box="[939,1032,359,380]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="7" pageNumber="7">Figures 2</figureCitation>
and S
<figureCitation box="[1098,1110,359,380]" captionStart="Figure 6" captionStartId="9.[1038,1098,271,290]" captionTargetBox="[143,1003,271,671]" captionTargetId="figure-838@9.[142,1003,271,671]" captionTargetPageId="9" captionText="Figure 6. Heatmaps displaying the possible degree of deformation in Oculudentavis species O. khaungraae HPG-15-3 (A and B) and O. naga GRS-Ref-28627 (C and D). The shape variation (degree of deformation) between the original and retrodeformed 3D surface is color coded, with yellow and dark blue representing no deformation and high deformation, respectively. Heatmaps indicating the amount of retrodeformation provide a good visual aid to display the regions where a specimen would have been deformed and what its overall shape would have looked like prior to deformation. All changes are based on observed distortions in the specimens themselves. See also Data S1." figureDoi="http://doi.org/10.5281/zenodo.5013969" httpUri="https://zenodo.org/record/5013969/files/figure.png" pageId="7" pageNumber="7">6</figureCitation>
). The postdentary region is short, including a coronoid with a low, posteriorly set, coronoid eminence. The coronoid looks significantly larger in the
<typeStatus box="[1361,1448,417,438]" pageId="7" pageNumber="7">holotype</typeStatus>
of
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[835,911,446,467]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="17" phylum="Chordata" rank="species" species="naga">
O.
<emphasis box="[862,911,446,467]" italics="true" pageId="7" pageNumber="7">naga</emphasis>
</taxonomicName>
than in the
<typeStatus box="[1022,1109,446,467]" pageId="7" pageNumber="7">holotype</typeStatus>
of
<taxonomicName authorityName="Xing et al." box="[1137,1281,446,467]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="khaungraae">
<emphasis box="[1137,1281,446,467]" italics="true" pageId="7" pageNumber="7">O. khaungraae</emphasis>
</taxonomicName>
, especially in the development of the anterolateral and anteromedial processes. However, these differences could be due to damage during deformation, making it difficult to establish clear bone boundaries (e.g., between surangular and coronoid), as this was one of the most problematic regions to segment in both specimens.
</paragraph>
<paragraph blockId="7.[811,1450,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[834,911,622,643]" italics="true" pageId="7" pageNumber="7">Angular</emphasis>
(
<figureCitation box="[922,1014,622,643]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="7" pageNumber="7">Figures 2</figureCitation>
and S
<figureCitation box="[1078,1090,622,643]" captionStart="Figure 6" captionStartId="9.[1038,1098,271,290]" captionTargetBox="[143,1003,271,671]" captionTargetId="figure-838@9.[142,1003,271,671]" captionTargetPageId="9" captionText="Figure 6. Heatmaps displaying the possible degree of deformation in Oculudentavis species O. khaungraae HPG-15-3 (A and B) and O. naga GRS-Ref-28627 (C and D). The shape variation (degree of deformation) between the original and retrodeformed 3D surface is color coded, with yellow and dark blue representing no deformation and high deformation, respectively. Heatmaps indicating the amount of retrodeformation provide a good visual aid to display the regions where a specimen would have been deformed and what its overall shape would have looked like prior to deformation. All changes are based on observed distortions in the specimens themselves. See also Data S1." figureDoi="http://doi.org/10.5281/zenodo.5013969" httpUri="https://zenodo.org/record/5013969/files/figure.png" pageId="7" pageNumber="7">6</figureCitation>
). This is a very reduced and slender bone, limited to the posteroventral side of the jaw.
</paragraph>
<paragraph blockId="7.[811,1450,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[834,1005,680,701]" italics="true" pageId="7" pageNumber="7">Compound bone</emphasis>
(
<figureCitation box="[1018,1111,680,701]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="7" pageNumber="7">Figures 2</figureCitation>
and S
<figureCitation box="[1178,1190,680,701]" captionStart="Figure 6" captionStartId="9.[1038,1098,271,290]" captionTargetBox="[143,1003,271,671]" captionTargetId="figure-838@9.[142,1003,271,671]" captionTargetPageId="9" captionText="Figure 6. Heatmaps displaying the possible degree of deformation in Oculudentavis species O. khaungraae HPG-15-3 (A and B) and O. naga GRS-Ref-28627 (C and D). The shape variation (degree of deformation) between the original and retrodeformed 3D surface is color coded, with yellow and dark blue representing no deformation and high deformation, respectively. Heatmaps indicating the amount of retrodeformation provide a good visual aid to display the regions where a specimen would have been deformed and what its overall shape would have looked like prior to deformation. All changes are based on observed distortions in the specimens themselves. See also Data S1." figureDoi="http://doi.org/10.5281/zenodo.5013969" httpUri="https://zenodo.org/record/5013969/files/figure.png" pageId="7" pageNumber="7">6</figureCitation>
). There is no obvious suture between the surangular and the articular or prearticular in either specimen. Both specimens have a long retroarticular process and a short, deep adductor fossa. It is uncertain whether the coronoid reached the anterior margin of the adductor fossa.
</paragraph>
<paragraph blockId="7.[811,1450,271,1987]" pageId="7" pageNumber="7">
Although only part of the postcranial skeleton is preserved in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[811,894,885,906]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="7" pageNumber="7" phylum="Chordata" rank="species" species="naga">
<emphasis box="[811,894,885,906]" italics="true" pageId="7" pageNumber="7">O. naga</emphasis>
</taxonomicName>
, it shows a short neck with eight cervical vertebrae that are amphicoelous, atlantal arches bearing posterior zygapophyses, and a pectoral region comprising a T-shaped interclavicle, medially expanded clavicles, and a typically squamate scapulocoracoid with scapular, scapulocoracoid, and primary coracoid fenestrae.
</paragraph>
<paragraph blockId="7.[811,1450,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[834,932,1060,1081]" italics="true" pageId="7" pageNumber="7">Vertebrae</emphasis>
(
<figureCitation box="[943,1160,1060,1081]" captionStart="Figure 7" captionStartId="10.[159,219,1131,1150]" captionTargetBox="[223,733,271,1091]" captionTargetId="figure-771@10.[223,733,271,1091]" captionTargetPageId="10" captionText="Figure 7. Oculudentavis naga prior to being trapped in tree resin Scientific illustration by Stephanie Abramowicz." figureDoi="http://doi.org/10.5281/zenodo.5013971" httpUri="https://zenodo.org/record/5013971/files/figure.png" pageId="7" pageNumber="7">Figures S7Aand S7B</figureCitation>
). Eight cervical vertebrae are preserved, including the atlas and the axis, as well as a small number of dorsal vertebrae (using the traditional anatomical definition whereby the first dorsal vertebra is that with a rib that meets the sternum, contra Gauthier et al.
<superScript attach="none" box="[1228,1237,1172,1186]" fontSize="6" pageId="7" pageNumber="7">7</superScript>
). The atlantal arches are not fused, and they have well-developed postzygapophyses. The axis preserves the dens, which is already fused in place. The vertebrae are amphicoelous and notochordal, with low neural spines. There are simple semicircular intercentra visible in the anterior part of the neck, with only a weak ventromedian keel (
<figureCitation box="[817,914,1352,1373]" captionStart="Figure 7" captionStartId="10.[159,219,1131,1150]" captionTargetBox="[223,733,271,1091]" captionTargetId="figure-771@10.[223,733,271,1091]" captionTargetPageId="10" captionText="Figure 7. Oculudentavis naga prior to being trapped in tree resin Scientific illustration by Stephanie Abramowicz." figureDoi="http://doi.org/10.5281/zenodo.5013971" httpUri="https://zenodo.org/record/5013971/files/figure.png" pageId="7" pageNumber="7">Figure S7</figureCitation>
). As in living gekkotans, these elements are free and intercentral in position. The first visible cervical rib is on cervical six, but there may have been ribs more anteriorly. There are no gastralia.
</paragraph>
<paragraph blockId="7.[811,1450,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[834,912,1469,1490]" italics="true" pageId="7" pageNumber="7">Clavicle</emphasis>
(
<figureCitation box="[924,1092,1469,1490]" captionStart="Figure 7" captionStartId="10.[159,219,1131,1150]" captionTargetBox="[223,733,271,1091]" captionTargetId="figure-771@10.[223,733,271,1091]" captionTargetPageId="10" captionText="Figure 7. Oculudentavis naga prior to being trapped in tree resin Scientific illustration by Stephanie Abramowicz." figureDoi="http://doi.org/10.5281/zenodo.5013971" httpUri="https://zenodo.org/record/5013971/files/figure.png" pageId="7" pageNumber="7">Figure S7CS7E</figureCitation>
). The clavicles are expanded medially and have a well-defined clavicular fenestra completely enclosed by bone. The clavicles are separated at the ventral midline by tip of the T-shaped interclavicle. Dorsally, the clavicles appear to extend above the level of the scapula blade, possibly meeting a suprascapular cartilage.
</paragraph>
<paragraph blockId="7.[811,1450,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[834,1004,1645,1666]" italics="true" pageId="7" pageNumber="7">Scapulocoracoid</emphasis>
(
<figureCitation box="[1017,1197,1645,1666]" captionStart="Figure 7" captionStartId="10.[159,219,1131,1150]" captionTargetBox="[223,733,271,1091]" captionTargetId="figure-771@10.[223,733,271,1091]" captionTargetPageId="10" captionText="Figure 7. Oculudentavis naga prior to being trapped in tree resin Scientific illustration by Stephanie Abramowicz." figureDoi="http://doi.org/10.5281/zenodo.5013971" httpUri="https://zenodo.org/record/5013971/files/figure.png" pageId="7" pageNumber="7">Figures S7CS7E</figureCitation>
). Both scapulocoracoids are preserved and display an anterior primary coracoid emargination, anemarginated scapular blade, and a large circular scapulocoracoid emargination. Dorsal to the scapula, there is an irregular mass that may represent the suprascapular cartilage.
</paragraph>
<paragraph blockId="7.[811,1450,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[834,920,1791,1812]" italics="true" pageId="7" pageNumber="7">Sternum</emphasis>
(
<figureCitation box="[930,1043,1791,1812]" captionStart="Figure 7" captionStartId="10.[159,219,1131,1150]" captionTargetBox="[223,733,271,1091]" captionTargetId="figure-771@10.[223,733,271,1091]" captionTargetPageId="10" captionText="Figure 7. Oculudentavis naga prior to being trapped in tree resin Scientific illustration by Stephanie Abramowicz." figureDoi="http://doi.org/10.5281/zenodo.5013971" httpUri="https://zenodo.org/record/5013971/files/figure.png" pageId="7" pageNumber="7">Figure S7D</figureCitation>
). Only the anterior border of the cartilage sternum is preserved, suggesting it was rhomboid.
</paragraph>
<paragraph blockId="7.[811,1450,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[834,953,1849,1870]" italics="true" pageId="7" pageNumber="7">Interclavicle</emphasis>
(
<figureCitation box="[968,1085,1849,1870]" captionStart="Figure 7" captionStartId="10.[159,219,1131,1150]" captionTargetBox="[223,733,271,1091]" captionTargetId="figure-771@10.[223,733,271,1091]" captionTargetPageId="10" captionText="Figure 7. Oculudentavis naga prior to being trapped in tree resin Scientific illustration by Stephanie Abramowicz." figureDoi="http://doi.org/10.5281/zenodo.5013971" httpUri="https://zenodo.org/record/5013971/files/figure.png" pageId="7" pageNumber="7">Figure S7D</figureCitation>
). The interclavicle is T-shaped and quite robust.
</paragraph>
<paragraph blockId="7.[811,1450,271,1987]" pageId="7" pageNumber="7">
<emphasis box="[834,925,1908,1929]" italics="true" pageId="7" pageNumber="7">Humerus</emphasis>
(
<figureCitation box="[936,1050,1908,1929]" captionStart="Figure 7" captionStartId="10.[159,219,1131,1150]" captionTargetBox="[223,733,271,1091]" captionTargetId="figure-771@10.[223,733,271,1091]" captionTargetPageId="10" captionText="Figure 7. Oculudentavis naga prior to being trapped in tree resin Scientific illustration by Stephanie Abramowicz." figureDoi="http://doi.org/10.5281/zenodo.5013971" httpUri="https://zenodo.org/record/5013971/files/figure.png" pageId="7" pageNumber="7">Figure S7D</figureCitation>
). The proximal portion of the left humerus is present, preserving the humeral head and the lateral tuberosity.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.5013965" ID-Zenodo-Dep="5013965" httpUri="https://zenodo.org/record/5013965/files/figure.png" pageId="8" pageNumber="8" startId="8.[159,219,1316,1335]" targetBox="[176,1449,273,1275]" targetPageId="8">
<paragraph blockId="8.[159,1468,1316,1536]" box="[159,801,1316,1335]" pageId="8" pageNumber="8">
Figure 5. Phylogenetic trees showing the position of
<taxonomicName box="[664,801,1316,1335]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="23" phylum="Chordata" rank="genus">Oculudentavis</taxonomicName>
</paragraph>
<paragraph blockId="8.[159,1468,1316,1536]" pageId="8" pageNumber="8">
(A) Phylogenetic tree showing the position of
<taxonomicName box="[525,643,1343,1360]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[525,643,1343,1360]" italics="true" pageId="8" pageNumber="8">Oculudentavis</emphasis>
</taxonomicName>
usingan amniote dataset;
<superScript attach="left" box="[857,865,1339,1351]" fontSize="5" pageId="8" pageNumber="8">6</superScript>
(B) photograph of the holotypeof
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[1141,1205,1343,1360]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="naga">
<emphasis box="[1141,1155,1343,1360]" italics="true" pageId="8" pageNumber="8">O</emphasis>
.
<emphasis box="[1165,1205,1343,1360]" italics="true" pageId="8" pageNumber="8">naga</emphasis>
</taxonomicName>
; (C) simplified phylogenetic tree showing the position of
<taxonomicName box="[355,473,1368,1385]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[355,473,1368,1385]" italics="true" pageId="8" pageNumber="8">Oculudentavis</emphasis>
</taxonomicName>
using a squamate dataset
<superScript attach="left" box="[690,698,1364,1376]" fontSize="5" pageId="8" pageNumber="8">7</superScript>
combined with a molecular dataset
<superScript attach="left" box="[991,1006,1364,1376]" fontSize="5" pageId="8" pageNumber="8">15</superScript>
treating some morphological characters as ordered; (D) simplified phylogenetic tree showing the position of
<taxonomicName box="[584,702,1393,1410]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[584,702,1393,1410]" italics="true" pageId="8" pageNumber="8">Oculudentavis</emphasis>
</taxonomicName>
using the same dataset as in (C) but with all characters unordered;(E) simplified phylogenetic tree showing the position of
<taxonomicName box="[397,515,1418,1435]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[397,515,1418,1435]" italics="true" pageId="8" pageNumber="8">Oculudentavis</emphasis>
</taxonomicName>
using the same dataset as (C) but removing molecular data. In (C)(E), crown groups were collapsed and are represented by silhouettes.
<taxonomicName baseAuthorityName="Gray" baseAuthorityYear="1842" box="[364,547,1443,1460]" class="Reptilia" family="Sphenodontidae" genus="Sphenodon" kingdom="Animalia" order="Rhynchocephalia" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="punctatus">
<emphasis box="[364,547,1443,1460]" italics="true" pageId="8" pageNumber="8">Sphenodon punctatus</emphasis>
</taxonomicName>
,
<taxonomicName authorityName="Cope" authorityYear="1885" box="[558,748,1443,1460]" class="Reptilia" family="Dibamidae" genus="Anelytropsis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="papillosus">
<emphasis box="[558,748,1443,1460]" italics="true" pageId="8" pageNumber="8">Anelytropsis papillosus</emphasis>
</taxonomicName>
(
<taxonomicName authorityName="Boulenger" authorityYear="1884" box="[758,851,1443,1460]" class="Reptilia" family="Dibamidae" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="family">Dibamidae</taxonomicName>
),
<taxonomicName authority="(Gekkota)" baseAuthorityName="Gekkota" box="[863,1160,1443,1460]" class="Reptilia" family="Sphaerodactylidae" genus="Sphaerodactylus" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="klauberi">
<emphasis box="[863,1072,1443,1460]" italics="true" pageId="8" pageNumber="8">Sphaerodactylus klauberi</emphasis>
(Gekkota)
</taxonomicName>
,
<taxonomicName baseAuthorityName="A.Smith" baseAuthorityYear="1844" box="[1168,1313,1443,1460]" class="Reptilia" family="Cordylidae" genus="Smaug" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="giganteus">
<emphasis box="[1168,1313,1443,1460]" italics="true" pageId="8" pageNumber="8">Smaug giganteus</emphasis>
</taxonomicName>
(
<taxonomicName authorityName="Fitzinger" authorityYear="1826" box="[1322,1416,1443,1460]" class="Reptilia" family="Cordylidae" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="family">Cordylidae</taxonomicName>
),
<taxonomicName authorityName="Baird" authorityYear="1858" class="Reptilia" family="Xantusiidae" genus="Xantusia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="vigilis">
<emphasis italics="true" pageId="8" pageNumber="8">Xantusia vigilis</emphasis>
</taxonomicName>
(
<taxonomicName authorityName="Baird" authorityYear="1858" box="[259,360,1469,1486]" class="Reptilia" family="Xantusiidae" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="family">Xantusiidae</taxonomicName>
),
<taxonomicName authorityName="De Rooij" authorityYear="1909" box="[372,540,1469,1486]" class="Reptilia" family="Scincidae" genus="Tribolonotus" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="gracilis">
<emphasis box="[372,540,1469,1486]" italics="true" pageId="8" pageNumber="8">Tribolonotus gracilis</emphasis>
</taxonomicName>
(
<taxonomicName authorityName="Oppel" authorityYear="1811" box="[550,636,1469,1486]" class="Reptilia" family="Scincidae" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="family">Scincidae</taxonomicName>
),
<taxonomicName authority="(Lacertoidea)" baseAuthorityName="Lacertoidea" box="[648,914,1469,1486]" class="Reptilia" family="Gymnophthalmidae" genus="Bachia" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="flavescens">
<emphasis box="[648,798,1469,1486]" italics="true" pageId="8" pageNumber="8">Bachia flavescens</emphasis>
(Lacertoidea)
</taxonomicName>
,
<taxonomicName authority="(Lacertoidea)" baseAuthorityName="Lacertoidea" box="[923,1176,1469,1486]" class="Reptilia" family="Lacertidae" genus="Lacerta" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="bilineata">
<emphasis box="[923,1059,1469,1486]" italics="true" pageId="8" pageNumber="8">Lacerta bilineata</emphasis>
(Lacertoidea)
</taxonomicName>
,
<taxonomicName authority="(Amphisbaenia)" baseAuthorityName="Amphisbaenia" box="[1185,1464,1469,1486]" class="Reptilia" family="Blanidae" genus="Blanus" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="cinereous">
<emphasis box="[1185,1328,1469,1486]" italics="true" pageId="8" pageNumber="8">Blanus cinereous</emphasis>
(Amphisbaenia)
</taxonomicName>
,
<taxonomicName authority="(Anguimorpha)" baseAuthorityName="Anguimorpha" box="[159,476,1494,1511]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="komodoensis">
<emphasis box="[159,344,1494,1511]" italics="true" pageId="8" pageNumber="8">Varanus komodoensis</emphasis>
(Anguimorpha)
</taxonomicName>
,
<taxonomicName authority="(Iguania)" baseAuthorityName="Iguania" box="[487,775,1494,1511]" class="Reptilia" family="Agamidae" genus="Physignathus" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="cocincinus">
<emphasis box="[487,694,1494,1511]" italics="true" pageId="8" pageNumber="8">Physignathus cocincinus</emphasis>
(Iguania)
</taxonomicName>
, and
<taxonomicName authority="(Serpentes)" baseAuthorityName="Serpentes" box="[824,1107,1494,1511]" class="Reptilia" family="Elapidae" genus="Ophiophagus" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="hannah">
<emphasis box="[824,1004,1494,1511]" italics="true" pageId="8" pageNumber="8">Ophiophagus hannah</emphasis>
(Serpentes)
</taxonomicName>
are shown. Node values indicate Bremer support; nodes with no support were recovered in the implied weights analyses. See also Data S1.
</paragraph>
</caption>
<paragraph blockId="8.[159,799,1616,1987]" pageId="8" pageNumber="8">
<emphasis box="[182,287,1616,1637]" italics="true" pageId="8" pageNumber="8">Soft tissue</emphasis>
(
<figureCitation box="[300,381,1616,1637]" captionStart="Figure 1" captionStartId="2.[1053,1113,271,290]" captionTargetBox="[159,1016,271,1338]" captionTargetId="figure-191@2.[159,1016,271,1338]" captionTargetPageId="2" captionText="Figure 1. Oculudentavis naga (GRS-Ref- 286278) displaying the superb preservation of bone and soft tissue (A) Dorsal; (B) lateral; (C) ventral views. Scale bar represents 10 mm. Diptera associated with the lizard skeleton were identified as Phoridae, Platypezidae, Ceratopogonidae, or Brachycera (Empidoidea). See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5013955" httpUri="https://zenodo.org/record/5013955/files/figure.png" pageId="8" pageNumber="8">Figure 1</figureCitation>
; Data S1, Gular scales in
<taxonomicName box="[646,785,1616,1637]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="genus">
<emphasis box="[646,785,1616,1637]" italics="true" pageId="8" pageNumber="8">Oculudentavis</emphasis>
</taxonomicName>
). Both specimens also preserve soft tissue. The head and body are covered in small, granular scales, with large rectangular supralabial and infralabial scales, tiny scales covering the eyelid, and a nostril placed anterior to the midpoint of each retracted narial opening (
<figureCitation box="[252,344,1762,1783]" captionStart="Figure 1" captionStartId="2.[1053,1113,271,290]" captionTargetBox="[159,1016,271,1338]" captionTargetId="figure-191@2.[159,1016,271,1338]" captionTargetPageId="2" captionText="Figure 1. Oculudentavis naga (GRS-Ref- 286278) displaying the superb preservation of bone and soft tissue (A) Dorsal; (B) lateral; (C) ventral views. Scale bar represents 10 mm. Diptera associated with the lizard skeleton were identified as Phoridae, Platypezidae, Ceratopogonidae, or Brachycera (Empidoidea). See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5013955" httpUri="https://zenodo.org/record/5013955/files/figure.png" pageId="8" pageNumber="8">Figures 1</figureCitation>
and
<figureCitation box="[397,409,1762,1783]" captionStart="Figure 2" captionStartId="4.[159,219,1918,1937]" captionTargetId="figure-0@3.[163,1427,271,1897]" captionTargetPageId="3" captionText="Figure 2. Comparison of the two specimens of Oculudentavis, each bone digitally segmented Synchrotron HRCT of O.naga (AJ) and O. khaungraae (KT).(A and K) Anterior view, (B and L) posterior view, (C, M, I, and S) dorsal view,(D, N, J, and T) ventral view, (E and O) left lateral view, (F, G, P, and Q) right lateral view, and (H and R) medial view are shown. See also Figures S1S7 and Data S1." figureDoi="http://doi.org/10.5281/zenodo.5014013" httpUri="https://zenodo.org/record/5014013/files/figure.png" pageId="8" pageNumber="8">2</figureCitation>
) in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[447,525,1762,1783]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="naga">
<emphasis box="[447,525,1762,1783]" italics="true" pageId="8" pageNumber="8">O. naga</emphasis>
</taxonomicName>
. There are no osteoderms. On the ventral surface of the head in
<taxonomicName authorityName="Bolet &amp; Stanley &amp; Daza &amp; Arias &amp; Cernanský &amp; Vidal-García &amp; Bauer &amp; Bevitt &amp; Peretti &amp; Evans" authorityYear="2021" box="[530,609,1791,1812]" class="Reptilia" family="Ceratopogonidae" genus="Oculudentavis" kingdom="Animalia" order="Squamata" pageId="8" pageNumber="8" phylum="Chordata" rank="species" species="naga">
<emphasis box="[530,609,1791,1812]" italics="true" pageId="8" pageNumber="8">O. naga</emphasis>
</taxonomicName>
, along the midline, the epidermal scales are raised and form a line of evenly spaced short ridges. Posterior to this midventral line, the skin of the gular region is thrown into a series of narrow linear folds. This folded region underlies the hyoid ceratobranchials and may demonstrate the resting anatomy of loose gular skin that could be inflated, for example in territorial display,in associationwith hyoidmovements.
</paragraph>
</subSubSection>
</treatment>
</document>