treatments-xml/data/03/D8/DD/03D8DD41FFCAB446FC29FD1B0D38FB9E.xml

<document id="6B7AB96E19C8AF5A9EF0AC94397CB948" ID-CLB-Dataset="306628" ID-DOI="10.1093/zoolinnean/zlae133" ID-GBIF-Dataset="43347976-f064-4571-8add-a1f636695906" ID-ISSN="0024-4082" ID-Zenodo-Dep="14514549" ID-ZooBank="DF599A3-0A7B-4A76-AA28-81147F6733FF" IM.bibliography_approvedBy="guilherme" IM.illustrations_approvedBy="guilherme" IM.materialsCitations_approvedBy="guilherme" IM.metadata_approvedBy="guilherme" IM.tables_requiresApprovalFor="GgImagineBatch,operationResults" IM.taxonomicNames_approvedBy="guilherme" IM.treatments_approvedBy="guilherme" checkinTime="1734033366110" checkinUser="plazi" docAuthor="Georgalis, Georgios L, Smith, Krister T, Marivaux, Laurent, Herrel, Anthony, Essid, El Mabrouk, Khayati Ammar, Hayet, Marzougui, Wissem, Temani, Rim &amp; Tabuce, Rodolphe" docDate="2024" docId="03D8DD41FFCAB446FC29FD1B0D38FB9E" docLanguage="en" docName="zlae133.pdf" docOrigin="Zoological Journal of the Linnean Society (zlae 133) 202 (3)" docSource="https://doi.org/10.1093/zoolinnean/zlae133" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.9:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="9" docTitle="Terastiodontosaurus marcelosanchezi Georgalis &amp; Smith 2024, sp. nov." docType="treatment" docVersion="2" lastPageNumber="21" masterDocId="FFE1A539FFCEB45CFF82FE630C53FFED" masterDocTitle="The world’s largest worm lizard: a new giant trogonophid (Squamata: Amphisbaenia) with extreme dental adaptations from the Eocene of Chambi, Tunisia" masterLastPageNumber="44" masterPageNumber="1" pageNumber="5" updateTime="1734529714011" updateUser="ExternalLinkService">
<mods:mods id="CF82050533D608D868AC631B0A25969B" xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo id="A057424C31298C54B54B1041372DE1C4">
<mods:title id="01C1E2B1F151F0503FC8F2E46D9C7B45">The world’s largest worm lizard: a new giant trogonophid (Squamata: Amphisbaenia) with extreme dental adaptations from the Eocene of Chambi, Tunisia</mods:title>
</mods:titleInfo>
<mods:name id="9572AE189C1E818DC370CC71B017ED8D" type="personal">
<mods:role id="2858A8B6396F6149680AA94F5D90BC00">
<mods:roleTerm id="2FDC0CD8D714F8A09A369561B91DA275">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="18EBDE66D0D5D31756886E05D4639CAC">Georgalis, Georgios L</mods:namePart>
</mods:name>
<mods:name id="591C9E9188C11AD91275E488174A6C9C" type="personal">
<mods:role id="1D1F704A33B0B84CF7288F86CC3B175E">
<mods:roleTerm id="D6A70B25AECE3BE3AE02A5DF7C831309">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="D6921B56F5650D4096777068E60DB458">Smith, Krister T</mods:namePart>
</mods:name>
<mods:name id="13C72B0B804B14F106191BDB5FDA5451" type="personal">
<mods:role id="2A9D7DC469DEC5DC7A232A3877B79895">
<mods:roleTerm id="8D3A6AD5C434B9006B4E662423A05CFD">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="70705E6BD12F2553F970E5986D53B69E">Marivaux, Laurent</mods:namePart>
<mods:affiliation id="55782D9F10FB3920DC2BACD1913DBB75">Institut des Sciences de l’Évolution de Montpellier (ISE-M * UMR 5554 * CNRS / UM / IRD / EPHE) * Université de Montpellier * 34095 Montpellier Cedex 5 * France</mods:affiliation>
<mods:nameIdentifier id="0C7EA4A76E248BE62CD58BCA67167826" type="email">georgalis@isez.pan.krakow.pl</mods:nameIdentifier>
</mods:name>
<mods:name id="51D05C39C7A9A0B53373857790FEFF45" type="personal">
<mods:role id="322F835BAF2053CCD6A2DF23FE029350">
<mods:roleTerm id="38F991E54B3850F3250F45F56650E71E">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="2A02EFAB3ED4E80EDF61F8687D0E4540">Herrel, Anthony</mods:namePart>
<mods:affiliation id="65C0EA0E75E9C364B6B20A68A223304D">Mécanismes Adaptatifs et Evolution * UMR 7179 * Muséum national d’Histoire naturelle CNRS * Paris * France &amp; Department of Biology * Evolutionary Morphology of Vertebrates * Ghent University * Ghent * Belgium &amp; Department of Biology * University of Antwerp * Wilrijk * Belgium &amp; Naturhistorisches Museum Bern * Bern * Switzerland</mods:affiliation>
<mods:nameIdentifier id="81AC196C73860284AF73E9E2AD0A834F" type="email">georgalis@isez.pan.krakow.pl</mods:nameIdentifier>
</mods:name>
<mods:name id="C43EC375CAB704C09CA64276CFE46FAE" type="personal">
<mods:role id="0E074F8566FA17E98244EEE09AF79CEA">
<mods:roleTerm id="21CF738BDF1AC562A8CE95FB7A12438C">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="F1F8DCBF2EA7F383FFBCC0E3C1AB054A">Essid, El Mabrouk</mods:namePart>
<mods:affiliation id="B908C420EF8CA9FC3EB860FB1CB06F25">Office National des Mines (ONM) * 24 rue 8601 * 2035 La Charguia * Tunis * Tunisia</mods:affiliation>
</mods:name>
<mods:name id="2D8C4F021EECFA7861DAAE127940220D" type="personal">
<mods:role id="41B3333F8FBAB682947AA6367CAA82A4">
<mods:roleTerm id="BB55F0B5F01AD1938DFC048A6F8E248A">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="B8810E3A3B175E8DDF15D43690199987">Khayati Ammar, Hayet</mods:namePart>
</mods:name>
<mods:name id="872F42DBF56FC3C3EBBFE6EA340A1ACE" type="personal">
<mods:role id="D70FE66EFE495B4EA24DB080E5EC3AC3">
<mods:roleTerm id="1FED45E8AB919B3EBC6A0AC310328CEC">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="4308B1B0E902BCEF9F382550C043590E">Marzougui, Wissem</mods:namePart>
<mods:affiliation id="458052DB7383457E6E3E2B4DFB728FBC">Office National des Mines (ONM) * 24 rue 8601 * 2035 La Charguia * Tunis * Tunisia</mods:affiliation>
</mods:name>
<mods:name id="82EF1DB092B493C0B90777D269FB4D42" type="personal">
<mods:role id="87E5314D477868553277511E4CD02DAA">
<mods:roleTerm id="9E49FD93E85BE3DB0E713CDAB0BB98CB">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="3E43F8993652C5E027A8660B5AA6E823">Temani, Rim</mods:namePart>
<mods:affiliation id="D48256ED8FF796B882AB9D0361AE7ECC">Office National des Mines (ONM) * 24 rue 8601 * 2035 La Charguia * Tunis * Tunisia</mods:affiliation>
</mods:name>
<mods:name id="2D8522C6FAC02B11C2EF89264B47F46F" type="personal">
<mods:role id="52AE243FFA08E305FA7CADF608E02A92">
<mods:roleTerm id="EF2241735466D61BA9B3F5317692A71E">Author</mods:roleTerm>
</mods:role>
<mods:namePart id="1A15C8A2F8D31C690CE1490DA88DF126">Tabuce, Rodolphe</mods:namePart>
<mods:affiliation id="CAE148BFE408BBB0885375E36CC31657">Institut des Sciences de l’Évolution de Montpellier (ISE-M * UMR 5554 * CNRS / UM / IRD / EPHE) * Université de Montpellier * 34095 Montpellier Cedex 5 * France</mods:affiliation>
<mods:nameIdentifier id="4DCBA1F2C007E06AC46A2B456C7220B4" type="email">georgalis@isez.pan.krakow.pl</mods:nameIdentifier>
</mods:name>
<mods:typeOfResource id="64B130426ED0D0DF58CBBA1434F39A84">text</mods:typeOfResource>
<mods:relatedItem id="A491EA7EE165587F577ECA612FA140E0" type="host">
<mods:titleInfo id="2B8A52B5C06F6E6506312DC77EDE9A54">
<mods:title id="910919F84C23ED0ED63473E26FC9B353">Zoological Journal of the Linnean Society</mods:title>
</mods:titleInfo>
<mods:part id="8B5A09AC9641C758B55172B448BB3AF4">
<mods:date id="080F4E3EC6E07FDD2D4D920D1725587E">2024</mods:date>
<mods:detail id="249F7D22637526B1B27394B42ACA8B10" type="series">
<mods:title id="38CB17270F019751EFE079518918AA52">zlae 133</mods:title>
</mods:detail>
<mods:detail id="9C0374010C35A758A1219B2CFB3976FD" type="pubDate">
<mods:number id="F1D14221431FDE2CB93423F2DE46AAE4">2024-11-21</mods:number>
</mods:detail>
<mods:detail id="08506A755F35949F11550B7CC8D25C5B" type="volume">
<mods:number id="22B8C4A5395135F5A082EBCC06379F49">202</mods:number>
</mods:detail>
<mods:detail id="43F62369AC6379A2560E56EFA9E68339" type="issue">
<mods:number id="438085C366052EB5F4D22D49D5840FED">3</mods:number>
</mods:detail>
<mods:extent id="4CC7FBCC262789C94D15656A0FA9485F" unit="page">
<mods:start id="9DE943E7EA7777905744551A7D467140">1</mods:start>
<mods:end id="640B95F34E6D605B7587F04E74A78517">44</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location id="B84D9B6FEAF432465E96BE53B7EEC50D">
<mods:url id="78C24CCEFD7D8AA2A59B286CB1147E78">https://doi.org/10.1093/zoolinnean/zlae133</mods:url>
</mods:location>
<mods:classification id="36AFBFA55884ACCD8EC0474C3EFAF1DA">journal article</mods:classification>
<mods:identifier id="BD662A834C8C6A7A6A70A34787963AC6" type="CLB-Dataset">306628</mods:identifier>
<mods:identifier id="5F28B9416BBBB1D0ECF9F6D4D4DF5BDC" type="DOI">10.1093/zoolinnean/zlae133</mods:identifier>
<mods:identifier id="23C349A94CB09A04573570794FDA59AA" type="GBIF-Dataset">43347976-f064-4571-8add-a1f636695906</mods:identifier>
<mods:identifier id="BD263200F221CECE6FF022F6EBC15728" type="ISSN">0024-4082</mods:identifier>
<mods:identifier id="A16BE642F21D15BC6C910339B0D15A04" type="Zenodo-Dep">14514549</mods:identifier>
<mods:identifier id="323A85F87CA160F21DB4F9E7B473A9E4" type="ZooBank">DF599A3-0A7B-4A76-AA28-81147F6733FF</mods:identifier>
</mods:mods>
<treatment id="03D8DD41FFCAB446FC29FD1B0D38FB9E" ID-DOI="http://doi.org/10.5281/zenodo.14514557" ID-Zenodo-Dep="14514557" LSID="urn:lsid:plazi:treatment:03D8DD41FFCAB446FC29FD1B0D38FB9E" httpUri="http://treatment.plazi.org/id/03D8DD41FFCAB446FC29FD1B0D38FB9E" lastPageId="26" lastPageNumber="21" pageId="4" pageNumber="5">
<subSubSection id="C36B3FDCFFCAB458FC29FD1B095DFC5F" pageId="4" pageNumber="5" type="nomenclature">
<paragraph id="8BCE6C57FFCAB458FC29FD1B095DFC5F" blockId="4.[939,1360,888,946]" pageId="4" pageNumber="5">
<emphasis id="B905B045FFCAB458FC29FD1B095DFC5F" bold="true" pageId="4" pageNumber="5">
<taxonomicName id="4C7117D4FFCAB458FC29FD1B08E1FC5F" authority="Georgalis &amp; Smith" authorityName="Georgalis &amp; Smith" authorityYear="2024" class="Reptilia" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" order="Squamata" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="marcelosanchezi" status="sp. nov.">
<emphasis id="B905B045FFCAB458FC29FD1B0903FC7E" bold="true" box="[939,1360,888,915]" italics="true" pageId="4" pageNumber="5">Terastiodontosaurus marcelosanchezi</emphasis>
Georgalis &amp; Smith
</taxonomicName>
<taxonomicNameLabel id="A2360D3EFFCAB458FB35FDFB095DFC5F" box="[1207,1294,920,946]" pageId="4" pageNumber="5" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C36B3FDCFFCAB458FCDCFDB508E4FBA3" pageId="4" pageNumber="5" type="description">
<paragraph id="8BCE6C57FFCAB458FCDCFDB509CDFC1D" blockId="4.[862,1438,982,1009]" box="[862,1438,982,1009]" pageId="4" pageNumber="5">
<emphasis id="B905B045FFCAB458FCDCFDB509CDFC1D" bold="true" box="[862,1438,982,1009]" pageId="4" pageNumber="5">
(
<figureCitation id="134A70D2FFCAB458FCEAFDB40F82FC1C" box="[872,977,983,1009]" captionStart-0="Figure 2" captionStart-1="Figure 3" captionStart-10="Figure 12" captionStart-11="Figure 13" captionStart-12="Figure 14" captionStart-13="Figure 15" captionStart-14="Figure 16" captionStart-15="Figure 17" captionStart-2="Figure 4" captionStart-3="Figure 5" captionStart-4="Figure 6" captionStart-5="Figure 7" captionStart-6="Figure 8" captionStart-7="Figure 9" captionStart-8="Figure 10" captionStart-9="Figure 11" captionStartId-0="6.[129,194,1839,1863]" captionStartId-1="7.[113,178,1839,1863]" captionStartId-10="17.[113,178,1844,1868]" captionStartId-11="19.[113,178,1288,1312]" captionStartId-12="20.[129,194,1626,1650]" captionStartId-13="21.[113,178,1840,1864]" captionStartId-14="22.[129,194,1840,1864]" captionStartId-15="23.[113,178,1844,1868]" captionStartId-2="8.[129,194,1934,1958]" captionStartId-3="9.[113,178,1840,1864]" captionStartId-4="11.[113,178,1661,1685]" captionStartId-5="12.[129,194,1408,1432]" captionStartId-6="13.[113,178,1839,1863]" captionStartId-7="14.[129,194,1836,1860]" captionStartId-8="15.[113,178,1840,1864]" captionStartId-9="16.[129,194,1840,1864]" captionTargetBox-0="[155,1448,146,1809]" captionTargetBox-1="[166,1405,146,1809]" captionTargetBox-10="[165,1408,151,1814]" captionTargetBox-11="[116,1456,146,1258]" captionTargetBox-12="[136,1459,156,1585]" captionTargetBox-13="[162,1409,147,1810]" captionTargetBox-14="[180,1423,146,1810]" captionTargetBox-15="[166,1406,151,1814]" captionTargetBox-2="[131,1471,514,1904]" captionTargetBox-3="[174,1399,147,1810]" captionTargetBox-4="[116,1455,146,1632]" captionTargetBox-5="[131,1471,146,1378]" captionTargetBox-6="[165,1408,146,1809]" captionTargetBox-7="[162,1439,151,1799]" captionTargetBox-8="[150,1421,148,1810]" captionTargetBox-9="[182,1423,146,1809]" captionTargetId-0="figure-75@6.[153,1450,144,1811]" captionTargetId-1="figure-6@7.[164,1407,144,1811]" captionTargetId-10="figure-7@17.[163,1410,149,1816]" captionTargetId-11="figure-358@19.[114,1458,144,1260]" captionTargetId-12="figure-160@20.[129,1473,144,1599]" captionTargetId-13="figure-6@21.[160,1411,145,1812]" captionTargetId-14="figure-7@22.[178,1425,144,1812]" captionTargetId-15="figure-6@23.[164,1408,149,1816]" captionTargetId-2="figure-224@8.[129,1473,512,1906]" captionTargetId-3="figure-7@9.[172,1401,145,1812]" captionTargetId-4="figure-170@11.[114,1458,144,1634]" captionTargetId-5="figure-323@12.[129,1473,144,1380]" captionTargetId-6="figure-6@13.[163,1410,144,1811]" captionTargetId-7="figure-8@14.[156,1446,144,1808]" captionTargetId-8="figure-7@15.[148,1423,145,1812]" captionTargetId-9="figure-8@16.[177,1425,144,1811]" captionTargetPageId-0="6" captionTargetPageId-1="7" captionTargetPageId-10="17" captionTargetPageId-11="19" captionTargetPageId-12="20" captionTargetPageId-13="21" captionTargetPageId-14="22" captionTargetPageId-15="23" captionTargetPageId-2="8" captionTargetPageId-3="9" captionTargetPageId-4="11" captionTargetPageId-5="12" captionTargetPageId-6="13" captionTargetPageId-7="14" captionTargetPageId-8="15" captionTargetPageId-9="16" captionText-0="Figure 2. Holotype right maxilla (ONM CBI-1-645) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), dorsal (C), and ventral (D) views." captionText-1="Figure 3. Holotype right maxilla (ONM CBI-1-645) of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of the specimen in labial (A), medial (B), ventromedial (C), ventral (D), dorsal (E), dorsolateral (F), dorsomedial (G), anterodorsal (H), anterior (I), and posterior (J) views." captionText-10="Figure 12. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, posterior fragment of right maxilla ONM CBI-1-667 in labial (A), medial (B), and ventral (C) views; D–F, posterior fragment of right maxilla ONM CBI-1-1017 in labial (D), ventromedial (E), and ventral (E) views; G–I, anterior fragment of left maxilla ONM CBI-1-1012 in labial (G), medial (H), and ventral (I) views; J–L, anterior fragment of left maxilla ONM CBI-1-1016 in labial (J), medial (K), and ventral (L) views; M–O, anterior fragment of left maxilla ONM CBI-1-1018 in labial (M), medial (N), and ventral (O) views; and P–R, anterior fragment of left maxilla ONM CBI-1-1022 in labial (P), medial (Q), and ventral (R) views." captionText-11="Figure 13. Dentary of Terastiodontosaurus marcelosanchezi. Photographs of left dentary ONM CBI-1-647 in labial (A) and medial (B) views." captionText-12="Figure 14. Dentaries of Terastiodontosaurus marcelosanchezi. Photographs of left dentary ONM CBI-1-657 in medial view (A); left dentary ONM CBI-1-655 in labial view (B); and right dentary ONM CBI-1-656 in medial view (C)." captionText-13="Figure 15. Dentaries of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of: A–C, left dentary ONM CBI-1-657 in labial (A), medial (B), and dorsal (C) views; D–F, left dentary ONM CBI-1-659 in labial (D), medial (E), and dorsal (F) views; G–I, right dentary ONM CBI-1-660 in labial (G), medial (H), and dorsal (I) views; J–L, left dentary ONM CBI-1-661 in labial (J), medial (K), and dorsal (L) views; M–O, left dentary ONM CBI-1-668 in labial (M), medial (N), and dorsal (O) views; and P–S, left dentary ONM CBI-1-670 in labial (P), medial (Q), dorsal (R), and posterior (S) views." captionText-14="Figure 16. Dentaries of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left dentary ONM CBI-1-662 in labial (A), medial (B), and dorsal (C) views; D, E, right dentary ONM CBI-1-1013 in labial (D) and dorsal (E) views; F–H, right dentary ONM CBI-1-666 in labial (F), dorsolabial (G), and medial (H) views; I–K, anterior fragment of right dentary ONM CBI-1-1020 in labial (I), dorsal (J), and medial (K) views; L–N, anterior fragment of left dentary ONM CBI-1-1015 in labial (L), dorsal (M), and medial (N) views; and O–Q, anterior fragment of left dentary ONM CBI-1-1014 in dorsal (O), labial (P), and medial (Q) views." captionText-15="Figure 17. Vertebrae tentatively attributed to Terastiodontosaurus marcelosanchezi. A–E, presacral vertebra ONM CBI-1-833 in anterior (A), posterior (B), right lateral (C), dorsal (D), and ventral (E) views; F–J, presacral vertebra ONM CBI-1-860 in anterior (F), posterior (G), left lateral (H), dorsal (I), and ventral (J) views; K–O, presacral vertebra ONM CBI-1-820 in anterior (K), posterior (L), left lateral (M), dorsal (N), and ventral (O) views; P–R, presacral vertebra ONM CBI-1-682 in anterior (P), dorsal (Q), and ventral (R) views; S–U, presacral vertebra ONM CBI-1-687 in dorsal (S), ventral (T), and right lateral (U) views; V, W, anterior caudal vertebra ONM CBI-1-689 in left lateral (V) and ventral (W) views; X, Y, posterior caudal vertebra ONM CBI-1-686 in ventral (X) and left lateral (Y) views." captionText-2="Figure 4. Paratype left dentary (ONM CBI-1-646) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), and dorsal (C) views." captionText-3="Figure 5. Paratype left dentary (ONM CBI-1-646) of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of the specimen in labial (A), medial (B), ventromedial (C), dorsal (D), ventral (E), anterodorsal (F), and posterior (G) views. Note that the specimen was slightly damaged during micro-computed tomography scanning, hence the difference from the photographs in Figure 4." captionText-4="Figure 6. Premaxillae of Terastiodontosaurus marcelosanchezi. Photographs of the larger specimens:A–C, ONM CBI-1-711 in anterior (A), right lateral (B), and posterior (C) views; and D–G, ONM CBI-1-658 in anterior (D), posterior (E), dorsal (F), and ventral (G) views." captionText-5="Figure 7. Premaxillae of Terastiodontosaurus marcelosanchezi. Photographs of the smaller specimens:A–D, ONM CBI-1-672 in anterior (A), right lateral (B), posterior (C), and dorsal (D) views; and E–H, ONM CBI-1-1021 in anterior (E), right anterolateral (F), posterior (G), right dorsolateral (H) views." captionText-6="Figure 8. Premaxillae of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of: A–F, ONM CBI-1-711 in anterior (A), right lateral (B), left ventrolateral (C), posterior (D), ventral (E), and dorsal (F) views; G–L, ONM CBI-1-658 in anterior (G), right lateral (H), left lateral (I), posterior (J), dorsal (K), and ventral (L) views; and M–R, ONM CBI-1-672 in anterior (M), right lateral (N), left lateral (O), posterior (P), ventral (Q), and dorsal (R) views." captionText-7="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." captionText-8="Figure 10. Maxillae of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of: A–D, left maxilla ONM CBI-1-648 in labial (A), medial (B), dorsal (C), and ventral (D) views; E–J, right maxilla ONM CBI-1-651 in labial (E), ventral (F), dorsal (G), medial (H), ventromedial (I), and posteroventral (J) views; and K–M, fragment of right maxilla ONM CBI-1-650 in labial (K), medial (L), and ventral (M) views. Note that specimen ONM CBI-1-648 was slightly damaged during micro-computed tomography scanning, hence the difference from the photographs in Figure 9." captionText-9="Figure 11. Maxillae of Terastiodontosaurus marcelosanchezi, small-sized individuals. μCT 3D reconstructions of: A–I, right maxilla ONM CBI-1-654 in labial (A), medial (B), anteromedial (C), ventral (D), ventromedial (E), posteromedial (F), posteroventral (G), dorsal (H), and dorsomedial (I) views; J–Q, right maxilla ONM CBI-1-649 in labial (J), medial (K), posteromedial (L), ventral (M), anteroventral (N), ventromedial (O), dorsal (P), and anterior (Q) views; and R–U, posterior portion of left maxilla ONM CBI-1-653 in labial (R), medial (S), ventral (T), and posteromedial (U) views." figureDoi-0="http://doi.org/10.5281/zenodo.14514555" figureDoi-1="http://doi.org/10.5281/zenodo.14514559" figureDoi-10="http://doi.org/10.5281/zenodo.14514579" figureDoi-11="http://doi.org/10.5281/zenodo.14514581" figureDoi-12="http://doi.org/10.5281/zenodo.14514583" figureDoi-13="http://doi.org/10.5281/zenodo.14514585" figureDoi-14="http://doi.org/10.5281/zenodo.14514587" figureDoi-15="http://doi.org/10.5281/zenodo.14514589" figureDoi-2="http://doi.org/10.5281/zenodo.14514561" figureDoi-3="http://doi.org/10.5281/zenodo.14514563" figureDoi-4="http://doi.org/10.5281/zenodo.14514567" figureDoi-5="http://doi.org/10.5281/zenodo.14514569" figureDoi-6="http://doi.org/10.5281/zenodo.14514571" figureDoi-7="http://doi.org/10.5281/zenodo.14514573" figureDoi-8="http://doi.org/10.5281/zenodo.14514575" figureDoi-9="http://doi.org/10.5281/zenodo.14514577" httpUri-0="https://zenodo.org/record/14514555/files/figure.png" httpUri-1="https://zenodo.org/record/14514559/files/figure.png" httpUri-10="https://zenodo.org/record/14514579/files/figure.png" httpUri-11="https://zenodo.org/record/14514581/files/figure.png" httpUri-12="https://zenodo.org/record/14514583/files/figure.png" httpUri-13="https://zenodo.org/record/14514585/files/figure.png" httpUri-14="https://zenodo.org/record/14514587/files/figure.png" httpUri-15="https://zenodo.org/record/14514589/files/figure.png" httpUri-2="https://zenodo.org/record/14514561/files/figure.png" httpUri-3="https://zenodo.org/record/14514563/files/figure.png" httpUri-4="https://zenodo.org/record/14514567/files/figure.png" httpUri-5="https://zenodo.org/record/14514569/files/figure.png" httpUri-6="https://zenodo.org/record/14514571/files/figure.png" httpUri-7="https://zenodo.org/record/14514573/files/figure.png" httpUri-8="https://zenodo.org/record/14514575/files/figure.png" httpUri-9="https://zenodo.org/record/14514577/files/figure.png" pageId="4" pageNumber="5">Figs 2–17</figureCitation>
, 
<figureCitation id="134A70D2FFCAB458FC58FDB40855FC1C" box="[986,1030,983,1009]" captionStart="Figure 23" captionStartId="33.[115,180,1358,1382]" captionTargetBox="[114,1458,479,1331]" captionTargetId="figure-201@33.[114,1458,479,1331]" captionTargetPageId="33" captionText="Figure 23.The profound enamel thickness in the teeth of Trogonophidae in comparison to other amphisbaenians.A, paratype dentary (ONM CBI-1-646) of Terastiodontosaurus marcelosanchezi. B, holotype dentary (UM THR 407) of Todrasaurus gheerbranti. C, Trogonophis wiegmanni (YPM HERR 6903), transverse section of snout. D, Zygaspis quadrifrons (collection of A.H., uncatalogued), transverse section of snout. E, Monopeltis capensis (collection of A.H., uncatalogued), transverse section of snout." figureDoi="http://doi.org/10.5281/zenodo.14514607" httpUri="https://zenodo.org/record/14514607/files/figure.png" pageId="4" pageNumber="5">23A</figureCitation>
; Supporting Information, 
<figureCitation id="134A70D2FFCAB458FA9FFDB409D5FC1C" box="[1309,1414,983,1009]" captionStart="Figure 1" captionStartId="2.[129,194,1021,1045]" captionTargetBox="[162,1441,146,991]" captionTargetId="figure-409@2.[160,1443,144,993]" captionTargetPageId="2" captionText="Figure 1. A, map of northern Africa, indicating the position of the late early–early middle Eocene locality of Chambi-1 (CBI-1). Also shown are the sole other few Palaeogene African localities that have yielded fossil amphisbaenians (with the exception of the record from the middle Eocene of Black Crow, Namibia). Circle for Palaeocene, square for Eocene. Map adapted from d-maps (d-maps.com). B, photograph of the CBI-1 fossil-bearing locality; arrow indicates the position of the fossiliferous limestone. C, close-up photograph of the fossiliferous limestone." figureDoi="http://doi.org/10.5281/zenodo.14514553" httpUri="https://zenodo.org/record/14514553/files/figure.png" pageId="4" pageNumber="5">Figs S1–S</figureCitation>
<figureCitation id="134A70D2FFCAB458FA04FDB409C0FC1C" box="[1414,1427,983,1009]" captionStart="Figure 6" captionStartId="11.[113,178,1661,1685]" captionTargetBox="[116,1455,146,1632]" captionTargetId="figure-170@11.[114,1458,144,1634]" captionTargetPageId="11" captionText="Figure 6. Premaxillae of Terastiodontosaurus marcelosanchezi. Photographs of the larger specimens:A–C, ONM CBI-1-711 in anterior (A), right lateral (B), and posterior (C) views; and D–G, ONM CBI-1-658 in anterior (D), posterior (E), dorsal (F), and ventral (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514567" httpUri="https://zenodo.org/record/14514567/files/figure.png" pageId="4" pageNumber="5">6</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="8BCE6C57FFCAB458FCBBFA7408E4FBA3" blockId="4.[825,1474,1047,1102]" pageId="4" pageNumber="5">
<emphasis id="B905B045FFCAB458FCBBFA740875FBC2" box="[825,1062,1047,1071]" italics="true" pageId="4" pageNumber="5">Zoobank registration:</emphasis>
<uri id="FFE06055FFCAB458FBC5FA7408E4FBA3" pageId="4" pageNumber="5">
urn:lsid:zoobank.org:act: 
<uuid id="FFD75682FFCAB458FAC6FA7408E4FBA3" pageId="4" pageNumber="5">881978AE-4954-4D2C-8D6E-12CD56CB4C20</uuid>
</uri>
</paragraph>
</subSubSection>
<subSubSection id="C36B3FDCFFCAB458FCBBFA130860FAC8" pageId="4" pageNumber="5" type="etymology">
<paragraph id="8BCE6C57FFCAB458FCBBFA130860FAC8" blockId="4.[824,1476,1136,1317]" pageId="4" pageNumber="5">
<emphasis id="B905B045FFCAB458FCBBFA130FFBFB65" box="[825,936,1136,1160]" italics="true" pageId="4" pageNumber="5">Etymology:</emphasis>
The species epithet is named after Professor Marcelo Sánchez-Villagra, director of the Palaeontological Institute of the University of 
<collectingRegion id="49B5A2B5FFCAB458FC44FACC085BFB2A" box="[966,1032,1199,1223]" country="Switzerland" name="Zuerich" pageId="4" pageNumber="5">Zurich</collectingRegion>
, as an honour for his major contributions to palaeontology, zoology, and evolutionary biology, in addition to the kind friendship and his great support to the first author (G.L.G.) for many years.
</paragraph>
</subSubSection>
<subSubSection id="C36B3FDCFFCAB459FCBBFB250ED4FEEB" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5" type="materials_examined">
<paragraph id="8BCE6C57FFCAB458FCBBFB250924FAB3" blockId="4.[825,1399,1350,1374]" box="[825,1399,1350,1374]" pageId="4" pageNumber="5">
<materialsCitation id="3B19660AFFCAB458FCBBFB250950FAB3" box="[825,1283,1350,1374]" collectionCode="ONM" country="Tunisia" location="Djebel Chambi" pageId="4" pageNumber="5" specimenCode="ONM CBI-1-645" specimenCount="1" stateProvince="Kasserine region" typeStatus="holotype">
<emphasis id="B905B045FFCAB458FCBBFB250FCAFAB3" box="[825,921,1350,1374]" italics="true" pageId="4" pageNumber="5">
<typeStatus id="54CAD2F5FFCAB458FCBBFB250FC7FAB3" box="[825,916,1350,1374]" pageId="4" pageNumber="5" type="holotype">Holotype</typeStatus>
:
</emphasis>
A right maxilla (
<specimenCode id="DBD7C42CFFCAB458FBC0FB2508AFFAB3" box="[1090,1276,1350,1374]" pageId="4" pageNumber="5">ONM CBI-1-645</specimenCode>
)
</materialsCitation>
(
<figureCitation id="134A70D2FFCAB458FA91FB25091CFAB3" box="[1299,1359,1350,1374]" captionStart="Figure 2" captionStartId="6.[129,194,1839,1863]" captionTargetBox="[155,1448,146,1809]" captionTargetId="figure-75@6.[153,1450,144,1811]" captionTargetPageId="6" captionText="Figure 2. Holotype right maxilla (ONM CBI-1-645) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), dorsal (C), and ventral (D) views." figureDoi="http://doi.org/10.5281/zenodo.14514555" httpUri="https://zenodo.org/record/14514555/files/figure.png" pageId="4" pageNumber="5">Figs 2</figureCitation>
, 3).
</paragraph>
<paragraph id="8BCE6C57FFCAB458FCBBFBE309CCFA7A" blockId="4.[825,1439,1407,1432]" box="[825,1439,1407,1432]" pageId="4" pageNumber="5">
<materialsCitation id="3B19660AFFCAB458FCBBFBE308A5FA7A" box="[825,1270,1407,1432]" collectionCode="ONM" country="Tunisia" location="Djebel Chambi" pageId="4" pageNumber="5" specimenCode="ONM CBI-1-646" specimenCount="1" stateProvince="Kasserine region" typeStatus="paratype">
<emphasis id="B905B045FFCAB458FCBBFBE30FCBFA7A" box="[825,920,1408,1431]" italics="true" pageId="4" pageNumber="5">
<typeStatus id="54CAD2F5FFCAB458FCBBFBE30FC1FA7A" box="[825,914,1408,1431]" pageId="4" pageNumber="5" type="paratype">Paratype</typeStatus>
:
</emphasis>
A left dentary (
<specimenCode id="DBD7C42CFFCAB458FBB6FB1C08BEFA7A" box="[1076,1261,1407,1431]" pageId="4" pageNumber="5">ONM CBI-1-646</specimenCode>
)
</materialsCitation>
(
<figureCitation id="134A70D2FFCAB458FA87FB1C0912FA7A" box="[1285,1345,1407,1431]" captionStart="Figure 4" captionStartId="8.[129,194,1934,1958]" captionTargetBox="[131,1471,514,1904]" captionTargetId="figure-224@8.[129,1473,512,1906]" captionTargetPageId="8" captionText="Figure 4. Paratype left dentary (ONM CBI-1-646) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), and dorsal (C) views." figureDoi="http://doi.org/10.5281/zenodo.14514561" httpUri="https://zenodo.org/record/14514561/files/figure.png" pageId="4" pageNumber="5">Figs 4</figureCitation>
, 5, 23A).
</paragraph>
<paragraph id="8BCE6C57FFCAB459FCBBFBDA0ED4FEEB" blockId="4.[825,1476,1464,1990]" lastBlockId="5.[112,762,144,262]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
<emphasis id="B905B045FFCAB458FCBBFBDA0FABFA3D" box="[825,1016,1465,1489]" italics="true" pageId="4" pageNumber="5">Referred specimens:</emphasis>
Four premaxillae (ONM CBI-1-658, ONM CBI-1-672, ONM CBI-1-711, and ONM CBI-1-1021), five right maxillae (ONM CBI-1-649, ONM CBI-1-651, ONM CBI-1-654, ONM CBI-1-667, and ONM CBI-1-1017), six left maxillae (ONM CBI-1-648, ONM CBI-1-653, ONM CBI-1- 1012, ONM CBI-1-1016, ONM CBI-1-1018, and ONM CBI-1-1022), one right maxilla fragment (ONM CBI-1-650), five right dentaries (ONM CBI-1-656, ONM CBI-1-660, ONM CBI-1-666, ONM CBI-1-1013, and ONM CBI-1-1020), 10 left dentaries (ONM CBI-1-647, ONM CBI-1-655, ONM CBI-1- 657, ONM CBI-1-659, ONM CBI-1-661, ONM CBI-1-662, ONM CBI-1-668, ONM CBI-1-670, ONM CBI-1-1014, and ONM CBI-1-1015), a fragment of the coronoid process of a left dentary (ONM CBI-1-664), and 17 tooth-bearing bone fragments (ONM CBI-1-652 and ONM CBI-1-671 [16 elements]). Tentatively also: numerous presacral vertebrae (ONM CBI-1- 673, ONM CBI-1-682, ONM CBI-1-685, ONM CBI-1-687, ONM CBI-1-691 [‘lot / batch’ with numerous vertebrae], ONM CBI-1-706 [‘lot / batch’ with numerous vertebrae], ONM CBI-1-820, ONM CBI-1-833, and ONM CBI-1-860) and two caudal vertebrae (ONM CBI-1-686 and ONM CBI-1-689).
</paragraph>
</subSubSection>
<subSubSection id="C36B3FDCFFCBB459FFF3FF4B0D34FB21" pageId="5" pageNumber="6" type="diagnosis">
<paragraph id="8BCE6C57FFCBB459FFF3FF4B0DB0FD5A" blockId="5.[112,764,295,1228]" pageId="5" pageNumber="6">
<emphasis id="B905B045FFCBB459FFF3FF4B0E1DFED2" box="[113,590,295,319]" italics="true" pageId="5" pageNumber="6">
Diagnosis: 
<taxonomicName id="4C7117D4FFCBB459FF60FF440E1DFED2" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[226,590,295,319]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="marcelosanchezi">Terastiodontosaurus marcelosanchezi</taxonomicName>
</emphasis>
can be referred to Amphisbaenia based on the prominent and enlarged median premaxillary tooth, the large anterior premaxillary foramina, the low tooth count on the maxilla and dentary, the ventral extension of the mandibular symphysis below Meckel’s groove, the broad insertion fossa for mandibular adductor muscles on the posterolateral surface of the dentary, and the strong and elevated coronoid process of the dentary. 
<taxonomicName id="4C7117D4FFCBB459FDB5FC600D5CFDD7" authorityName="Georgalis &amp; Smith" authorityYear="2024" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFCBB459FDB5FC600D5CFDD7" italics="true" pageId="5" pageNumber="6">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
can be referred to 
<taxonomicName id="4C7117D4FFCBB459FE6CFC410EDFFDD7" authorityName="Bonaparte" authorityYear="1838" box="[494,652,546,570]" class="Squamata" family="Trogonophidae" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Trogonophidae</taxonomicName>
based on the presence of acrodont dentition, closely appressed (‘fused’) teeth, the interdigitating suture between the frontal and the facial process of the maxilla, and ectopterygoid abutting the posteromedial corner of the maxilla.
</paragraph>
<paragraph id="8BCE6C57FFCBB459FF0FFCDC0D34FB21" blockId="5.[112,764,295,1228]" pageId="5" pageNumber="6">
<taxonomicName id="4C7117D4FFCBB459FF0FFCDC0DA1FD3A" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[141,498,703,727]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFCBB459FF0FFCDC0DA1FD3A" box="[141,498,703,727]" italics="true" pageId="5" pageNumber="6">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
is united with 
<taxonomicName id="4C7117D4FFCBB459FD02FCDC0C88FD1B" authorityName="Kaup" authorityYear="1830" class="Squamata" family="Trogonophidae" genus="Trogonophis" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="wiegmanni">
<emphasis id="B905B045FFCBB459FD02FCDC0C88FD1B" italics="true" pageId="5" pageNumber="6">Trogonophis wiegmanni</emphasis>
</taxonomicName>
by: thick enamel on marginal teeth, and ‘twinning’ of paired premaxillary teeth, with median tooth separated by a diastema from paired teeth. 
<taxonomicName id="4C7117D4FFCBB459FEFBFD7E0EB1FCD8" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[377,738,797,821]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFCBB459FEFBFD7E0EB1FCD8" box="[377,738,797,821]" italics="true" pageId="5" pageNumber="6">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
is united with 
<taxonomicName id="4C7117D4FFCBB459FF6AFD5F0D80FCB9" box="[232,467,828,852]" genus="Todrasaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="gheerbranti">
<emphasis id="B905B045FFCBB459FF6AFD5F0D80FCB9" box="[232,467,828,852]" italics="true" pageId="5" pageNumber="6">Todrasaurus gheerbranti</emphasis>
</taxonomicName>
by: thick enamel on marginal teeth, extremely enlarged (&gt;60% longer than adjacent teeth) dentary tooth 3–4 positions from the rear, and small ‘hills’ on posterior dentary teeth. 
<taxonomicName id="4C7117D4FFCBB459FEE8FDF90E81FC5F" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[362,722,922,946]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFCBB459FEE8FDF90E81FC5F" box="[362,722,922,946]" italics="true" pageId="5" pageNumber="6">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
can be differentiated from all other amphisbaenians by the combination of the following features: very large size (with maximum known maxilla length&gt; 
<quantity id="4C89C1B2FFCBB459FEE7FD9B0DF8FBE2" box="[357,427,1016,1040]" metricMagnitude="-2" metricUnit="m" metricValue="1.6" pageId="5" pageNumber="6" unit="mm" value="16.0">16 mm</quantity>
and maximum dentary length of 
<quantity id="4C89C1B2FFCBB459FFF0FA740CEAFBC2" box="[114,185,1047,1071]" metricMagnitude="-2" metricUnit="m" metricValue="1.7" pageId="5" pageNumber="6" unit="mm" value="17.0">17 mm</quantity>
), the flat apical surface of the cheek teeth, the position of the largest tooth on the maxilla, the number of maxillary teeth (usually three maxillary teeth), the ratio of the largest maxillary tooth length to the total maxillary tooth row length (between 0.5 and 0.7), and the position of the largest tooth on the dentary (fourth from posterior).
</paragraph>
</subSubSection>
<subSubSection id="C36B3FDCFFCBB459FFF3FA820D1AFA6E" pageId="5" pageNumber="6" type="materials_examined">
<paragraph id="8BCE6C57FFCBB459FFF3FA820EB2FADA" blockId="5.[113,762,1249,1335]" pageId="5" pageNumber="6">
<emphasis id="B905B045FFCBB459FFF3FA820D2AFB14" box="[113,377,1249,1273]" italics="true" pageId="5" pageNumber="6">Type locality and horizon:</emphasis>
Chambi 1 (CBI-1), Djebel Chambi, 
<collectingRegion id="49B5A2B5FFCBB459FFF3FB630D48FAF5" box="[113,283,1280,1304]" country="Tunisia" name="Kasserine" pageId="5" pageNumber="6">Kassérine region</collectingRegion>
, western part of Central 
<collectingCountry id="F3662CC7FFCBB459FDA0FB630E38FAF5" box="[546,619,1280,1304]" name="Tunisia" pageId="5" pageNumber="6">Tunisia</collectingCountry>
, Tunisia; late early to early middle Eocene (late Ypresian to early Lutetian).
</paragraph>
<paragraph id="8BCE6C57FFCBB459FFF3FB2F0D1AFA6E" blockId="5.[113,763,1356,1412]" pageId="5" pageNumber="6">
<emphasis id="B905B045FFCBB459FFF3FB2F0DA0FA89" box="[113,499,1356,1380]" italics="true" pageId="5" pageNumber="6">Geographical and stratigraphical range:</emphasis>
Taxon known exclusively from its 
<typeStatus id="54CAD2F5FFCBB459FF45FB0F0CA6FA69" box="[199,245,1388,1412]" pageId="5" pageNumber="6">type</typeStatus>
locality.
</paragraph>
</subSubSection>
<subSubSection id="C36B3FDCFFCBB459FFF3FBFB0D56F966" pageId="5" pageNumber="6" type="discussion">
<paragraph id="8BCE6C57FFCBB459FFF3FBFB0D56F966" blockId="5.[113,763,1432,1675]" pageId="5" pageNumber="6">
<emphasis id="B905B045FFCBB459FFF3FBFB0E0AFA5D" box="[113,601,1432,1456]" italics="true" pageId="5" pageNumber="6">Nomenclatural remark on the new taxon:</emphasis>
The authorship of this new genus and species should be referred to as 
<taxonomicName id="4C7117D4FFCBB459FFF3FBB40DF3FA02" authority="Georgalis" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[113,416,1495,1519]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
<emphasis id="B905B045FFCBB459FFF3FBB40D66FA02" box="[113,309,1495,1519]" italics="true" pageId="5" pageNumber="6">Terastiodontosaurus</emphasis>
Georgalis
</taxonomicName>
&amp; Smith gen. nov. (for the new genus) and 
<taxonomicName id="4C7117D4FFCBB459FF7BFB950E8AF9E3" authority="Georgalis" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[249,729,1526,1550]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFCBB459FF7BFB950E3AF9E3" box="[249,617,1526,1550]" italics="true" pageId="5" pageNumber="6">Terastiodontosaurus marcelosanchezi</emphasis>
Georgalis
</taxonomicName>
&amp; Smith gen. et sp. nov. (for the new species), following Article 50.1 and the ‘recommendation 50A concerning multiple authors’ of the International Code of Zoological Nomenclature (ICZN 1999).
</paragraph>
</subSubSection>
<subSubSection id="C36B3FDCFFCBB446FEF4F8D50D38FB9E" lastPageId="26" lastPageNumber="27" pageId="5" pageNumber="6" type="description">
<paragraph id="8BCE6C57FFCBB459FEF4F8D50DA5F93D" blockId="5.[374,502,1718,1744]" box="[374,502,1718,1744]" pageId="5" pageNumber="6">
<emphasis id="B905B045FFCBB459FEF4F8D50DA5F93D" bold="true" box="[374,502,1718,1744]" pageId="5" pageNumber="6">Description</emphasis>
</paragraph>
<paragraph id="8BCE6C57FFCBB459FEC9F885081DFD93" blockId="5.[111,764,1765,1985]" lastBlockId="5.[809,1462,144,1985]" pageId="5" pageNumber="6">
<emphasis id="B905B045FFCBB459FEC9F8850E72F912" bold="true" box="[331,545,1765,1792]" pageId="5" pageNumber="6">
<typeStatus id="54CAD2F5FFCBB459FEC9F8850DE1F8ED" box="[331,434,1766,1792]" pageId="5" pageNumber="6" type="holotype">Holotype</typeStatus>
(
<figureCitation id="134A70D2FFCBB459FE43F8860DADF912" box="[449,510,1765,1791]" captionStart="Figure 2" captionStartId="6.[129,194,1839,1863]" captionTargetBox="[155,1448,146,1809]" captionTargetId="figure-75@6.[153,1450,144,1811]" captionTargetPageId="6" captionText="Figure 2. Holotype right maxilla (ONM CBI-1-645) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), dorsal (C), and ventral (D) views." figureDoi="http://doi.org/10.5281/zenodo.14514555" httpUri="https://zenodo.org/record/14514555/files/figure.png" pageId="5" pageNumber="6">Figs 2</figureCitation>
, 
<figureCitation id="134A70D2FFCBB459FD8AF8860E45F912" box="[520,534,1765,1791]" captionStart="Figure 3" captionStartId="7.[113,178,1839,1863]" captionTargetBox="[166,1405,146,1809]" captionTargetId="figure-6@7.[164,1407,144,1811]" captionTargetPageId="7" captionText="Figure 3. Holotype right maxilla (ONM CBI-1-645) of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of the specimen in labial (A), medial (B), ventromedial (C), ventral (D), dorsal (E), dorsolateral (F), dorsomedial (G), anterodorsal (H), anterior (I), and posterior (J) views." figureDoi="http://doi.org/10.5281/zenodo.14514559" httpUri="https://zenodo.org/record/14514559/files/figure.png" pageId="5" pageNumber="6">3</figureCitation>
)
</emphasis>
Theholotyperightmaxilla (ONMCBI-1-645)isalmostcomplete (
<figureCitation id="134A70D2FFCBB459FFFEF9480CE9F8AE" box="[124,186,1835,1859]" captionStart="Figure 2" captionStartId="6.[129,194,1839,1863]" captionTargetBox="[155,1448,146,1809]" captionTargetId="figure-75@6.[153,1450,144,1811]" captionTargetPageId="6" captionText="Figure 2. Holotype right maxilla (ONM CBI-1-645) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), dorsal (C), and ventral (D) views." figureDoi="http://doi.org/10.5281/zenodo.14514555" httpUri="https://zenodo.org/record/14514555/files/figure.png" pageId="5" pageNumber="6">Figs 2</figureCitation>
, 3). The premaxillary process curves up anterodorsally, forming part of the medial border of the external naris. The sharp rim weakens anteriorly, and the anterior portion of the process as a whole slopes ventrolaterally. The dorsal surface is slightly striated, probably where it was overlapped by the septomaxilla. The ventral surface of the premaxillary process curves upwards and is more coarsely striated where it overlaps the premaxilla. The superior alveolar canal opens anteriorly through two foramina: a dorsally directed foramen on the medial side of the facial process, and an anteriorly directed foramen immediately anterior to the facial process; between them the canal is roofed, but there appears to be a narrow groove, as if two growing folds of perichondral bone met over a channel but did not fuse fully. Anterior to the anterior opening of the superior alveolar canal the dorsal surface of the premaxillary process is coarsely striated; here it was probably overlain by the septomaxilla, into which the neurovascular structure(s) of the superior alveolar nerve continued. It is unclear whether the division of the anterior opening of the superior alveolar canal marks a separation of the course of the subnarial artery and superior alveolar nerve, as in 
<taxonomicName id="4C7117D4FFCBB459FACCFC2409E0FDB2" box="[1358,1459,583,607]" class="Squamata" family="Iguanidae" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Iguanidae</taxonomicName>
(
<bibRefCitation id="EFE011A6FFCBB459FCB6FC050FEFFD93" author="Oelrich TM" box="[820,956,614,638]" pageId="5" pageNumber="6" pagination="1 - 122" refId="ref28440" refString="Oelrich TM. The anatomy of the head of Ctenosaura pectinata (Iguanidae). Miscellaneous Publications of the Museum of Zoology of the University of Michigan 1956; 94: 1 - 122. https: // doi. org / 10.2307 / 1440299" type="journal article" year="1956">Oelrich 1956</bibRefCitation>
, Smith 2009).
</paragraph>
<paragraph id="8BCE6C57FFCBB459FCC7FCE60FEDFBF8" blockId="5.[809,1462,144,1985]" pageId="5" pageNumber="6">
On the medial side of the premaxillary process is a strong, dorsomedially facing facet for the vomer. Behind this the palatal shelf is restricted and its dorsal surface hollows out, where Jacobson’s organ sat; together with the vomer, here the maxilla formed the fenestra vomeronasalis externa. Posterior to the cavity for Jacobson’s organ the maxilla possesses a distinct medial process that contacts the horizontal wing of the vomer. The process continues laterally across the palatal shelf and extends posterodorsally as a weak ridge on the medial surface of the facial process of the maxilla. In other squamates, Smith and 
<bibRefCitation id="EFE011A6FFCBB459FB9EFDDC089CFC3A" author="Gauthier JA" box="[1052,1231,959,983]" pageId="5" pageNumber="6" pagination="135 - 230" refId="ref30393" refString="ºmith KT * Gauthier JA. Early Eocene lizards of the Wasatch Formation near Bitter Creek * Wyoming: diversity and paleoenvironment during an interval of global warming. Bulletin of the Peabody Museum of Natural History 2013; 54: 135 - 230. https: // doi. org / 10.3374 / 014.054.0205" type="journal article" year="2013">Gauthier (2013)</bibRefCitation>
identified this as the ‘nasolacrimal ridge’ after ascertaining a relationship with the lacrimal duct.
</paragraph>
<paragraph id="8BCE6C57FFCBB459FCC7FA7E0824FB1C" blockId="5.[809,1462,144,1985]" pageId="5" pageNumber="6">Behind this process the palatal shelf is medially extensive. A distinct palatine process is absent, but an extensive, striated facet for the palatine articulation is present. The (posterior) superior alveolar foramen opens at the level of the anterior end of the palatine facet. A distinct facet for the ectopterygoid cannot be discerned, but it must have been an abutting one, and it might be confluent with the palatine facet.</paragraph>
<paragraph id="8BCE6C57FFCBB459FCC7FA9B0F03FA06" blockId="5.[809,1462,144,1985]" pageId="5" pageNumber="6">
The posterior process of the maxilla is broad and flares sharply outwards (or laterad 
<emphasis id="B905B045FFCBB459FB8BFB74087FFAC2" box="[1033,1068,1303,1327]" italics="true" pageId="5" pageNumber="6">fide</emphasis>
Gans and Montero 2008). Although striated surfaces of this maxilla commonly show tiny holes, the entire posterior surface of the posterior process is heavily porous and coarsely striated. Dorsally, the posterior process is thick and also coarsely longitudinally striated; there is no evidence that a distinct element articulated on this suborbital part of the maxilla.
</paragraph>
<paragraph id="8BCE6C57FFCBB459FCC7FB900FF3F82C" blockId="5.[809,1462,144,1985]" pageId="5" pageNumber="6">
The facial process is thick. A rough facet for the nasal bone is found on the anterior margin of the facial process and turns posterodorsally as it ascends the process. There is a distinct change in angulation below the peak of the facial process, and the facet behind it (now more on the medial than the anterior surface of the facial process) is smoother; probably, the change marks the boundary between the nasal and part of the frontal facet. Immediately behind its apex the facial process is deeply notched, almost certainly to receive a tongue-like process of the frontal bone. Additionally, there is a number of rough, anteroventrally trending ridges and grooves posterior to the frontal notch, which possibly indicate a more extensive overlap of the maxilla on the prefrontal (like in lizards) than is observed today in 
<taxonomicName id="4C7117D4FFCBB459FAB6F9E90FCAF82C" authorityName="Kaup" authorityYear="1830" class="Squamata" family="Trogonophidae" genus="Trogonophis" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="wiegmanni">
<emphasis id="B905B045FFCBB459FAB6F9E90FCAF82C" italics="true" pageId="5" pageNumber="6">Trogonophis wiegmanni</emphasis>
</taxonomicName>
.
</paragraph>
<caption id="DF0E3CDFFFC8B45AFF03F94C0E61F88E" ID-DOI="http://doi.org/10.5281/zenodo.14514555" ID-Zenodo-Dep="14514555" httpUri="https://zenodo.org/record/14514555/files/figure.png" pageId="6" pageNumber="7" startId="6.[129,194,1839,1863]" targetBox="[155,1448,146,1809]" targetPageId="6" targetType="figure">
<paragraph id="8BCE6C57FFC8B45AFF03F94C0E61F88E" blockId="6.[129,1428,1839,1892]" pageId="6" pageNumber="7">
<emphasis id="B905B045FFC8B45AFF03F94C0C8BF8AA" bold="true" box="[129,216,1839,1863]" pageId="6" pageNumber="7">Figure 2.</emphasis>
Holotype right maxilla (ONM CBI-1-645) of 
<taxonomicName id="4C7117D4FFC8B45AFD0DF94C0FA8F8AA" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[655,1019,1839,1863]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFC8B45AFD0DF94C0FA8F8AA" bold="true" box="[655,1019,1839,1863]" italics="true" pageId="6" pageNumber="7">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. Photographs of the specimen in labial (A), medial (B), dorsal (C), and ventral (D) views.
</paragraph>
</caption>
<paragraph id="8BCE6C57FFC8B454FF1EF91D0E2CFEA8" blockId="6.[129,778,1918,2005]" lastBlockId="8.[129,779,144,481]" lastPageId="8" lastPageNumber="9" pageId="6" pageNumber="7">
The lateral surface of the maxilla is porous and weakly rugose in its dorsal part, with some longitudinal grooves and ridges on the posterior process. There are three distinct labial foramina, all of them more or less above the level of the largest tooth. The two anterior ones are more closely spaced, while there is a considerable distance between the middle and posterior one, the latter also being the largest. These foramina communicate internally, as expected, with the superior alveolar canal, which runs longitudinally through the maxilla immediately above the tooth row. It conveys the superior alveolar nerve, which passes from the orbit into the maxillary canal through the superior alveolar foramen, in addition to the maxillary artery (
<bibRefCitation id="EFE011A6FFC6B454FE65FF4E0E3CFEA9" author="Oelrich TM" box="[487,623,300,325]" pageId="8" pageNumber="9" pagination="1 - 122" refId="ref28440" refString="Oelrich TM. The anatomy of the head of Ctenosaura pectinata (Iguanidae). Miscellaneous Publications of the Museum of Zoology of the University of Michigan 1956; 94: 1 - 122. https: // doi. org / 10.2307 / 1440299" type="journal article" year="1956">Oelrich 1956</bibRefCitation>
).
</paragraph>
<caption id="DF0E3CDFFFC9B45BFFF3F94C0D7FF892" ID-DOI="http://doi.org/10.5281/zenodo.14514559" ID-Zenodo-Dep="14514559" httpUri="https://zenodo.org/record/14514559/files/figure.png" pageId="7" pageNumber="8" startId="7.[113,178,1839,1863]" targetBox="[166,1405,146,1809]" targetPageId="7" targetType="figure">
<paragraph id="8BCE6C57FFC9B45BFFF3F94C0D7FF892" blockId="7.[113,1407,1838,1919]" pageId="7" pageNumber="8">
<emphasis id="B905B045FFC9B45BFFF3F94C0C9AF8AA" bold="true" box="[113,201,1839,1863]" pageId="7" pageNumber="8">Figure 3.</emphasis>
Holotype right maxilla (ONM CBI-1-645) of 
<taxonomicName id="4C7117D4FFC9B45BFD02F94D0FBFF8AB" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[640,1004,1838,1862]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFC9B45BFD02F94D0FBFF8AB" bold="true" box="[640,1004,1838,1862]" italics="true" pageId="7" pageNumber="8">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. μCT 3D reconstructions of the specimen in labial (A), medial (B), ventromedial (C), ventral (D), dorsal (E), dorsolateral (F), dorsomedial (G), anterodorsal (H), anterior (I), and posterior (J) views.
</paragraph>
</caption>
<paragraph id="8BCE6C57FFC6B454FF1EFF2F096FFEA8" blockId="8.[129,779,144,481]" lastBlockId="8.[825,1474,144,325]" pageId="8" pageNumber="9">The dentition is acrodont. The maxilla possesses only three teeth, of which the second is by far the largest (some 75% longer than the next-largest tooth) and, in fact, covers much of the ventral portion of the bone; this tooth is followed in size by the anteriormost tooth, while the posteriormost tooth is considerably tiny. The teeth are closely appressed, with no interdental gaps between them. All three teeth are much flattened, particularly the two largest ones: their dorsoventral height is extremely low. Moreover, the μCT scan reveals that the enamel is extremely thick (Supporting Information, Fig. S1A). Several nutritive foramina are present at the base of the teeth.</paragraph>
<paragraph id="8BCE6C57FFC6B454FB95FF0708B6FE93" blockId="8.[1047,1253,356,382]" box="[1047,1253,356,382]" pageId="8" pageNumber="9">
<emphasis id="B905B045FFC6B454FB95FF0708B6FE93" bold="true" box="[1047,1253,356,382]" pageId="8" pageNumber="9">
<typeStatus id="54CAD2F5FFC6B454FB95FF070824FE93" box="[1047,1143,356,382]" pageId="8" pageNumber="9" type="paratype">Paratype</typeStatus>
(
<figureCitation id="134A70D2FFC6B454FB06FF070891FE93" box="[1156,1218,356,382]" captionStart="Figure 4" captionStartId="8.[129,194,1934,1958]" captionTargetBox="[131,1471,514,1904]" captionTargetId="figure-224@8.[129,1473,512,1906]" captionTargetPageId="8" captionText="Figure 4. Paratype left dentary (ONM CBI-1-646) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), and dorsal (C) views." figureDoi="http://doi.org/10.5281/zenodo.14514561" httpUri="https://zenodo.org/record/14514561/files/figure.png" pageId="8" pageNumber="9">Figs 4</figureCitation>
, 
<figureCitation id="134A70D2FFC6B454FB4EFF070889FE93" box="[1228,1242,356,382]" captionStart="Figure 5" captionStartId="9.[113,178,1840,1864]" captionTargetBox="[174,1399,147,1810]" captionTargetId="figure-7@9.[172,1401,145,1812]" captionTargetPageId="9" captionText="Figure 5. Paratype left dentary (ONM CBI-1-646) of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of the specimen in labial (A), medial (B), ventromedial (C), dorsal (D), ventral (E), anterodorsal (F), and posterior (G) views. Note that the specimen was slightly damaged during micro-computed tomography scanning, hence the difference from the photographs in Figure 4." figureDoi="http://doi.org/10.5281/zenodo.14514563" httpUri="https://zenodo.org/record/14514563/files/figure.png" pageId="8" pageNumber="9">5</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="8BCE6C57FFC6B456FCBBFFE80C82FEEB" blockId="8.[825,1476,395,481]" lastBlockId="10.[127,780,144,1954]" lastPageId="10" lastPageNumber="11" pageId="8" pageNumber="9">
The 
<typeStatus id="54CAD2F5FFC6B454FCE5FFE80F93FE4E" box="[871,960,395,419]" pageId="8" pageNumber="9" type="paratype">paratype</typeStatus>
left dentary (ONM CBI-1-646) is relatively complete (
<figureCitation id="134A70D2FFC6B454FCFFFFC90FE8FE2F" box="[893,955,426,450]" captionStart="Figure 4" captionStartId="8.[129,194,1934,1958]" captionTargetBox="[131,1471,514,1904]" captionTargetId="figure-224@8.[129,1473,512,1906]" captionTargetPageId="8" captionText="Figure 4. Paratype left dentary (ONM CBI-1-646) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), and dorsal (C) views." figureDoi="http://doi.org/10.5281/zenodo.14514561" httpUri="https://zenodo.org/record/14514561/files/figure.png" pageId="8" pageNumber="9">Figs 4</figureCitation>
, 5). It is referred to the same species on the basis of co-occurrence and the following derived morphological features: great enamel thickness, reduced tooth row length and tooth count, presence of a greatly enlarged tooth, the ‘hill’ on the small, posteriormost teeth, and the apically flat teeth in the middle.
</paragraph>
<caption id="DF0E3CDFFFC6B454FF03F9ED0DEFF82F" ID-DOI="http://doi.org/10.5281/zenodo.14514561" ID-Zenodo-Dep="14514561" httpUri="https://zenodo.org/record/14514561/files/figure.png" pageId="8" pageNumber="9" startId="8.[129,194,1934,1958]" targetBox="[131,1471,514,1904]" targetPageId="8" targetType="figure">
<paragraph id="8BCE6C57FFC6B454FF03F9ED0DEFF82F" blockId="8.[129,1409,1934,1986]" pageId="8" pageNumber="9">
<emphasis id="B905B045FFC6B454FF03F9ED0C8BF84B" bold="true" box="[129,216,1934,1958]" pageId="8" pageNumber="9">Figure 4.</emphasis>
Paratype left dentary (ONM CBI-1-646) of 
<taxonomicName id="4C7117D4FFC6B454FDFEF9ED0FBBF84B" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[636,1000,1934,1958]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFC6B454FDFEF9ED0FBBF84B" bold="true" box="[636,1000,1934,1958]" italics="true" pageId="8" pageNumber="9">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. Photographs of the specimen in labial (A), medial (B), and dorsal (C) views.
</paragraph>
</caption>
<caption id="DF0E3CDFFFC7B455FFF3F9530885F86D" ID-DOI="http://doi.org/10.5281/zenodo.14514563" ID-Zenodo-Dep="14514563" httpUri="https://zenodo.org/record/14514563/files/figure.png" pageId="9" pageNumber="10" startId="9.[113,178,1840,1864]" targetBox="[174,1399,147,1810]" targetPageId="9" targetType="figure">
<paragraph id="8BCE6C57FFC7B455FFF3F9530885F86D" blockId="9.[113,1405,1840,1921]" pageId="9" pageNumber="10">
<emphasis id="B905B045FFC7B455FFF3F9530C9AF8A4" bold="true" box="[113,201,1840,1865]" pageId="9" pageNumber="10">Figure 5.</emphasis>
Paratype left dentary (ONM CBI-1-646) of 
<taxonomicName id="4C7117D4FFC7B455FDEFF9530F8AF8A5" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[621,985,1840,1864]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFC7B455FDEFF9530F8AF8A5" bold="true" box="[621,985,1840,1864]" italics="true" pageId="9" pageNumber="10">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. μCT 3D reconstructions of the specimen in labial (A), medial (B), ventromedial (C), dorsal (D), ventral (E), anterodorsal (F), and posterior (G) views. Note that the specimen was slightly damaged during micro-computed tomography scanning, hence the difference from the photographs in Figure 4.
</paragraph>
</caption>
<paragraph id="8BCE6C57FFC4B456FF1EFF6E0E7EFDCD" blockId="10.[127,780,144,1954]" pageId="10" pageNumber="11">
The symphysis is broad and anterodorsally inclined. As in other amphisbaenians (
<bibRefCitation id="EFE011A6FFC4B456FEF0FF4E0E6CFEA9" author="Longrich NR" box="[370,575,300,325]" pageId="10" pageNumber="11" refId="ref27285" refString="Longrich NR * Vinther J * Pyron RA et al. Biogeography of worm lizards (Amphisbaenia) driven by end-Cretaceous mass extinction. Proceedings Biological Sciences 2015; 282: 20143034. https: // doi. org / 10.1098 / rspb. 2014.3034" type="journal volume" year="2015">
Longrich 
<emphasis id="B905B045FFC4B456FE54FF4E0E55FEA8" box="[470,518,301,325]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
2015
</bibRefCitation>
), it is not restricted to the area above the Meckelian groove but instead curves anteroventrally around it and terminates posteroventrally in a sharp corner. It appears as though the Meckelian groove is closed and fused immediately behind the symphysis and that the Meckelian cartilage itself might be ossified as an anterodorsally– posteroventrally extensive wedge in the centre of the symphysis, but this interpretation is not entirely clear.
</paragraph>
<paragraph id="8BCE6C57FFC4B456FF1EFC440C86FAFD" blockId="10.[127,780,144,1954]" pageId="10" pageNumber="11">
Meckel’s groove is open (
<figureCitation id="134A70D2FFC4B456FE22FC440D8FFDD2" box="[416,476,551,575]" captionStart="Figure 4" captionStartId="8.[129,194,1934,1958]" captionTargetBox="[131,1471,514,1904]" captionTargetId="figure-224@8.[129,1473,512,1906]" captionTargetPageId="8" captionText="Figure 4. Paratype left dentary (ONM CBI-1-646) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), and dorsal (C) views." figureDoi="http://doi.org/10.5281/zenodo.14514561" httpUri="https://zenodo.org/record/14514561/files/figure.png" pageId="10" pageNumber="11">Figs 4</figureCitation>
, 5B, C). Above it is the indistinct supra-Meckelian lip (of Bhullar and Smith 2008), above which is the subdental shelf (
<emphasis id="B905B045FFC4B456FE24FC040D89FD93" box="[422,474,615,638]" italics="true" pageId="10" pageNumber="11">sensu</emphasis>
Rage and Augé 2010) that extends medial to the tooth row. The subdental shelf commences below the posterior portion of the first tooth. It is narrow anteriorly and becomes gradually wider in the posterior half of the dentary, and behind the tooth row it grades into the coronoid process. The part of the groove in which Meckel’s cartilage was situated is relatively narrow throughout most of its length, but it widens a bit posteriorly, immediately in front of the mandibular foramen. That groove is straight for most of its length but turns up strongly near the symphysis, giving it a hockey-stick shape, as in other amphisbaenians (
<bibRefCitation id="EFE011A6FFC4B456FE2CFDFC0ED3FC5A" author="Longrich NR" box="[430,640,927,951]" pageId="10" pageNumber="11" refId="ref27285" refString="Longrich NR * Vinther J * Pyron RA et al. Biogeography of worm lizards (Amphisbaenia) driven by end-Cretaceous mass extinction. Proceedings Biological Sciences 2015; 282: 20143034. https: // doi. org / 10.1098 / rspb. 2014.3034" type="journal volume" year="2015">
Longrich 
<emphasis id="B905B045FFC4B456FD97FDC30E15FC5A" box="[533,582,927,951]" italics="true" pageId="10" pageNumber="11">et al.</emphasis>
2015
</bibRefCitation>
). Posteriorly in the dentary there are two strong, elongate facets below the groove for Meckel’s cartilage, for the angular and the splenial. The presence of two distinct facets strongly suggests that those mandibular elements were discrete, not fused, as in 
<taxonomicName id="4C7117D4FFC4B456FD0DFA7E0CBFFBB9" authorityName="Kaup" authorityYear="1830" class="Squamata" family="Trogonophidae" genus="Trogonophis" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="wiegmanni">
<emphasis id="B905B045FFC4B456FD0DFA7E0CBFFBB9" italics="true" pageId="10" pageNumber="11">Trogonophis wiegmanni</emphasis>
</taxonomicName>
, where only one facet is present; of course, without the remainder of the mandible this cannot be taken as certain. The angular facet extends as far anteriorly as the anterior end of the enlarged tooth (fourth from the back), whereas the splenial facet only just passes the posterior end of the enlarged tooth. The ventral margin of the dentary behind the symphysis is straight.
</paragraph>
<paragraph id="8BCE6C57FFC4B456FF1EFB740D17F9A4" blockId="10.[127,780,144,1954]" pageId="10" pageNumber="11">The dentary possesses a posterodorsally ascending coronoid process immediately behind the posteriormost tooth. The dorsal tip of the process is incomplete, but even the preserved portion extends high above the tooth row, as in other amphisbaenians. The process is prominent and thick. There is a strong diagonal ridge on its lateral surface that delineates the adductor fossa dorsally. The ridge is narrower at its base but grows thick and porous as it curves dorsally and then diminishes; it might have served as an attachment site for other jaw adductors, probably via a bodenaponeurosis.</paragraph>
<paragraph id="8BCE6C57FFC4B456FF1EF8320EFFF8A9" blockId="10.[127,780,144,1954]" pageId="10" pageNumber="11">
The mandibular canal runs across the length of the dentary, transmitting the inferior alveolar nerve and mandibular artery; it communicates with the labial foramina present in the labial surface of the dentary and with the nutritive foramina at the bases of the teeth. Its entrance is the mandibular foramen, which is located well behind the tooth row at the level of the anterior edge of the coronoid process. In labial view (
<figureCitation id="134A70D2FFC4B456FD96F96E0E31F8C8" box="[532,610,1805,1829]" captionStart="Figure 4" captionStartId="8.[129,194,1934,1958]" captionTargetBox="[131,1471,514,1904]" captionTargetId="figure-224@8.[129,1473,512,1906]" captionTargetPageId="8" captionText="Figure 4. Paratype left dentary (ONM CBI-1-646) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), and dorsal (C) views." figureDoi="http://doi.org/10.5281/zenodo.14514561" httpUri="https://zenodo.org/record/14514561/files/figure.png" pageId="10" pageNumber="11">Figs 4A</figureCitation>
, 5A), four labial foramina are present in the anterior half of the dentary.
</paragraph>
<paragraph id="8BCE6C57FFC4B456FF1EF9280FE0FE2F" blockId="10.[127,780,144,1954]" lastBlockId="10.[825,1476,144,450]" pageId="10" pageNumber="11">
The dentary bears eight acrodont teeth, of which the largest is the fourth from posterior. The posteriormost tooth is the smallest one. The first two teeth are bulbous, and the anteriormost tooth is procumbent, extending beyond the anterior margin of the symphysis. As in the 
<typeStatus id="54CAD2F5FFC4B456FB89FECC0834FF2A" box="[1035,1127,175,199]" pageId="10" pageNumber="11" type="holotype">holotype</typeStatus>
maxilla described above, the teeth are closely appressed, with almost no interdental gaps between them (at maximum, only tiny empty spaces between some but not all teeth). All teeth except for the first two are much flattened (particularly the two largest ones), with the exception of the two anteriormost ones, which are bulbous and tall. The posterior teeth have tiny central cusps. Multiple nutritive foramina are situated above the subdental shelf ventrally to the each of the tooth bases.
</paragraph>
<paragraph id="8BCE6C57FFC4B456FB96FF9408B4FDFC" blockId="10.[1044,1255,503,529]" box="[1044,1255,503,529]" pageId="10" pageNumber="11">
<emphasis id="B905B045FFC4B456FB96FF9408B4FDFC" bold="true" box="[1044,1255,503,529]" pageId="10" pageNumber="11">Referred specimens</emphasis>
</paragraph>
<paragraph id="8BCE6C57FFC4B456FB8DFC4208BFFDD1" blockId="10.[824,1476,545,1517]" box="[1039,1260,545,572]" pageId="10" pageNumber="11">
<emphasis id="B905B045FFC4B456FB8DFC4208BFFDD1" box="[1039,1260,545,572]" italics="true" pageId="10" pageNumber="11">
Premaxillae (
<figureCitation id="134A70D2FFC4B456FB0DFC4108B1FDD1" box="[1167,1250,545,572]" captionStart-0="Figure 6" captionStart-1="Figure 7" captionStart-2="Figure 8" captionStartId-0="11.[113,178,1661,1685]" captionStartId-1="12.[129,194,1408,1432]" captionStartId-2="13.[113,178,1839,1863]" captionTargetBox-0="[116,1455,146,1632]" captionTargetBox-1="[131,1471,146,1378]" captionTargetBox-2="[165,1408,146,1809]" captionTargetId-0="figure-170@11.[114,1458,144,1634]" captionTargetId-1="figure-323@12.[129,1473,144,1380]" captionTargetId-2="figure-6@13.[163,1410,144,1811]" captionTargetPageId-0="11" captionTargetPageId-1="12" captionTargetPageId-2="13" captionText-0="Figure 6. Premaxillae of Terastiodontosaurus marcelosanchezi. Photographs of the larger specimens:A–C, ONM CBI-1-711 in anterior (A), right lateral (B), and posterior (C) views; and D–G, ONM CBI-1-658 in anterior (D), posterior (E), dorsal (F), and ventral (G) views." captionText-1="Figure 7. Premaxillae of Terastiodontosaurus marcelosanchezi. Photographs of the smaller specimens:A–D, ONM CBI-1-672 in anterior (A), right lateral (B), posterior (C), and dorsal (D) views; and E–H, ONM CBI-1-1021 in anterior (E), right anterolateral (F), posterior (G), right dorsolateral (H) views." captionText-2="Figure 8. Premaxillae of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of: A–F, ONM CBI-1-711 in anterior (A), right lateral (B), left ventrolateral (C), posterior (D), ventral (E), and dorsal (F) views; G–L, ONM CBI-1-658 in anterior (G), right lateral (H), left lateral (I), posterior (J), dorsal (K), and ventral (L) views; and M–R, ONM CBI-1-672 in anterior (M), right lateral (N), left lateral (O), posterior (P), ventral (Q), and dorsal (R) views." figureDoi-0="http://doi.org/10.5281/zenodo.14514567" figureDoi-1="http://doi.org/10.5281/zenodo.14514569" figureDoi-2="http://doi.org/10.5281/zenodo.14514571" httpUri-0="https://zenodo.org/record/14514567/files/figure.png" httpUri-1="https://zenodo.org/record/14514569/files/figure.png" httpUri-2="https://zenodo.org/record/14514571/files/figure.png" pageId="10" pageNumber="11">Figs 6–8</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="8BCE6C57FFC4B456FCBBFC2B0FC4FB98" blockId="10.[824,1476,545,1517]" pageId="10" pageNumber="11">
The most complete premaxillae are ONM CBI-1-672 and ONM CBI-1-711, which include a large portion of the nasal process, whereas this structure is mostly broken in ONM CBI-1-658 and ONM CBI-1-1021 (
<figureCitation id="134A70D2FFC4B456FB8BFCC5080CFD53" box="[1033,1119,678,702]" captionStart-0="Figure 6" captionStart-1="Figure 7" captionStart-2="Figure 8" captionStartId-0="11.[113,178,1661,1685]" captionStartId-1="12.[129,194,1408,1432]" captionStartId-2="13.[113,178,1839,1863]" captionTargetBox-0="[116,1455,146,1632]" captionTargetBox-1="[131,1471,146,1378]" captionTargetBox-2="[165,1408,146,1809]" captionTargetId-0="figure-170@11.[114,1458,144,1634]" captionTargetId-1="figure-323@12.[129,1473,144,1380]" captionTargetId-2="figure-6@13.[163,1410,144,1811]" captionTargetPageId-0="11" captionTargetPageId-1="12" captionTargetPageId-2="13" captionText-0="Figure 6. Premaxillae of Terastiodontosaurus marcelosanchezi. Photographs of the larger specimens:A–C, ONM CBI-1-711 in anterior (A), right lateral (B), and posterior (C) views; and D–G, ONM CBI-1-658 in anterior (D), posterior (E), dorsal (F), and ventral (G) views." captionText-1="Figure 7. Premaxillae of Terastiodontosaurus marcelosanchezi. Photographs of the smaller specimens:A–D, ONM CBI-1-672 in anterior (A), right lateral (B), posterior (C), and dorsal (D) views; and E–H, ONM CBI-1-1021 in anterior (E), right anterolateral (F), posterior (G), right dorsolateral (H) views." captionText-2="Figure 8. Premaxillae of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of: A–F, ONM CBI-1-711 in anterior (A), right lateral (B), left ventrolateral (C), posterior (D), ventral (E), and dorsal (F) views; G–L, ONM CBI-1-658 in anterior (G), right lateral (H), left lateral (I), posterior (J), dorsal (K), and ventral (L) views; and M–R, ONM CBI-1-672 in anterior (M), right lateral (N), left lateral (O), posterior (P), ventral (Q), and dorsal (R) views." figureDoi-0="http://doi.org/10.5281/zenodo.14514567" figureDoi-1="http://doi.org/10.5281/zenodo.14514569" figureDoi-2="http://doi.org/10.5281/zenodo.14514571" httpUri-0="https://zenodo.org/record/14514567/files/figure.png" httpUri-1="https://zenodo.org/record/14514569/files/figure.png" httpUri-2="https://zenodo.org/record/14514571/files/figure.png" pageId="10" pageNumber="11">Figs 6–8</figureCitation>
; Supporting Information, Fig. S2). The nasal process is large, dorsoventrally elongated and moderately wide; it gradually narrows in width apically. Given its preserved extent, it is likely that the nasal process reached the frontals, a condition unique to 
<taxonomicName id="4C7117D4FFC4B456FBFFFD400948FCD6" authorityName="Bonaparte" authorityYear="1838" box="[1149,1307,803,827]" class="Squamata" family="Trogonophidae" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="family">Trogonophidae</taxonomicName>
among amphisbaenians (
<bibRefCitation id="EFE011A6FFC4B456FC2AFD20086FFCB6" author="Kearney M" box="[936,1084,835,859]" pageId="10" pageNumber="11" pagination="1 - 74" refId="ref26695" refString="Kearney M. ºystematics of the Amphisbaenia (Lepidosauria: ºquamata) based on morphological evidence from recent and fossil forms. Herpetological Monographs 2003; 17: 1 - 74. https: // doi. org / 10.1655 / 0733 - 1347 (2003) 017 [0001: sotalb] 2.0. co; 2" type="journal article" year="2003">Kearney 2003</bibRefCitation>
), but more complete specimens are needed to verify this. A pair of large anterior premaxillary foramina is developed at the base of the nasal process. These are the anterior openings for the ethmoidal nerve that enters the premaxilla through the posterior premaxillary foramina on the posterior side of the nasal process. No rostral process or rostral blade is present. The palatal shelf or alveolar plate is most complete in ONM CBI-1-672 (
<figureCitation id="134A70D2FFC4B456FBDAFA7D08FAFBDB" box="[1112,1193,1054,1078]" captionStart="Figure 7" captionStartId="12.[129,194,1408,1432]" captionTargetBox="[131,1471,146,1378]" captionTargetId="figure-323@12.[129,1473,144,1380]" captionTargetPageId="12" captionText="Figure 7. Premaxillae of Terastiodontosaurus marcelosanchezi. Photographs of the smaller specimens:A–D, ONM CBI-1-672 in anterior (A), right lateral (B), posterior (C), and dorsal (D) views; and E–H, ONM CBI-1-1021 in anterior (E), right anterolateral (F), posterior (G), right dorsolateral (H) views." figureDoi="http://doi.org/10.5281/zenodo.14514569" httpUri="https://zenodo.org/record/14514569/files/figure.png" pageId="10" pageNumber="11">Figs 7C</figureCitation>
, D, 8P– 
<collectionCode id="ED60F492FFC4B456FB7CFA7D0943FBDB" box="[1278,1296,1054,1078]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="10" pageNumber="11">R</collectionCode>
), but it is ONM CBI-1-658 (
<figureCitation id="134A70D2FFC4B456FC38FA5E0856FBB8" box="[954,1029,1085,1109]" captionStart="Figure 6" captionStartId="11.[113,178,1661,1685]" captionTargetBox="[116,1455,146,1632]" captionTargetId="figure-170@11.[114,1458,144,1634]" captionTargetPageId="11" captionText="Figure 6. Premaxillae of Terastiodontosaurus marcelosanchezi. Photographs of the larger specimens:A–C, ONM CBI-1-711 in anterior (A), right lateral (B), and posterior (C) views; and D–G, ONM CBI-1-658 in anterior (D), posterior (E), dorsal (F), and ventral (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514567" httpUri="https://zenodo.org/record/14514567/files/figure.png" pageId="10" pageNumber="11">Figs 6F</figureCitation>
, G, 8J–L) that most clearly shows it to be bipartite.
</paragraph>
<paragraph id="8BCE6C57FFC4B456FCD7FA1F091FFA00" blockId="10.[824,1476,545,1517]" pageId="10" pageNumber="11">
The dentition is acrodont. On the alveolar plate, there are five teeth, all of bulbous morphology. The central median tooth is the most robust and prominent (a synapomorphy of amphisbaenians; see 
<bibRefCitation id="EFE011A6FFC4B456FC49FAB9086EFB1F" author="Gans C" box="[971,1085,1242,1266]" pageId="10" pageNumber="11" pagination="347 - 416" refId="ref24277" refString="Gans C. The characteristics and affinities of the Amphisbaenia. Transactions of the Zoological Society of London 1978; 34: 347 - 416. https: // doi. org / 10.1111 / j. 1096 - 3642.1978. tb 00376. x" type="journal article" year="1978">Gans 1978</bibRefCitation>
, Smith 2009). The μCT scans reveal that the median tooth possesses great apical enamel thickness (Supporting Information, Fig. S1C). There is a diastema between the median tooth and the lateral ones that contrasts strongly with the otherwise close appression of the teeth generally and gives the impression of the lateral two teeth being ‘twinned’. In fact, in the two pairs of lateral teeth, the dental gaps between the teeth are almost absent. A slight vertical striation is observable on all teeth, being more distinct in the central tooth.
</paragraph>
<paragraph id="8BCE6C57FFC4B456FB95F86F08B6F9CA" blockId="10.[825,1475,1548,1955]" box="[1047,1253,1548,1575]" pageId="10" pageNumber="11">
<emphasis id="B905B045FFC4B456FB95F86F08B6F9CA" box="[1047,1253,1548,1575]" italics="true" pageId="10" pageNumber="11">
Maxillae (
<figureCitation id="134A70D2FFC4B456FBF9F86E0889F9CB" box="[1147,1242,1548,1574]" captionStart-0="Figure 9" captionStart-1="Figure 10" captionStart-2="Figure 11" captionStart-3="Figure 12" captionStartId-0="14.[129,194,1836,1860]" captionStartId-1="15.[113,178,1840,1864]" captionStartId-2="16.[129,194,1840,1864]" captionStartId-3="17.[113,178,1844,1868]" captionTargetBox-0="[162,1439,151,1799]" captionTargetBox-1="[150,1421,148,1810]" captionTargetBox-2="[182,1423,146,1809]" captionTargetBox-3="[165,1408,151,1814]" captionTargetId-0="figure-8@14.[156,1446,144,1808]" captionTargetId-1="figure-7@15.[148,1423,145,1812]" captionTargetId-2="figure-8@16.[177,1425,144,1811]" captionTargetId-3="figure-7@17.[163,1410,149,1816]" captionTargetPageId-0="14" captionTargetPageId-1="15" captionTargetPageId-2="16" captionTargetPageId-3="17" captionText-0="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." captionText-1="Figure 10. Maxillae of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of: A–D, left maxilla ONM CBI-1-648 in labial (A), medial (B), dorsal (C), and ventral (D) views; E–J, right maxilla ONM CBI-1-651 in labial (E), ventral (F), dorsal (G), medial (H), ventromedial (I), and posteroventral (J) views; and K–M, fragment of right maxilla ONM CBI-1-650 in labial (K), medial (L), and ventral (M) views. Note that specimen ONM CBI-1-648 was slightly damaged during micro-computed tomography scanning, hence the difference from the photographs in Figure 9." captionText-2="Figure 11. Maxillae of Terastiodontosaurus marcelosanchezi, small-sized individuals. μCT 3D reconstructions of: A–I, right maxilla ONM CBI-1-654 in labial (A), medial (B), anteromedial (C), ventral (D), ventromedial (E), posteromedial (F), posteroventral (G), dorsal (H), and dorsomedial (I) views; J–Q, right maxilla ONM CBI-1-649 in labial (J), medial (K), posteromedial (L), ventral (M), anteroventral (N), ventromedial (O), dorsal (P), and anterior (Q) views; and R–U, posterior portion of left maxilla ONM CBI-1-653 in labial (R), medial (S), ventral (T), and posteromedial (U) views." captionText-3="Figure 12. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, posterior fragment of right maxilla ONM CBI-1-667 in labial (A), medial (B), and ventral (C) views; D–F, posterior fragment of right maxilla ONM CBI-1-1017 in labial (D), ventromedial (E), and ventral (E) views; G–I, anterior fragment of left maxilla ONM CBI-1-1012 in labial (G), medial (H), and ventral (I) views; J–L, anterior fragment of left maxilla ONM CBI-1-1016 in labial (J), medial (K), and ventral (L) views; M–O, anterior fragment of left maxilla ONM CBI-1-1018 in labial (M), medial (N), and ventral (O) views; and P–R, anterior fragment of left maxilla ONM CBI-1-1022 in labial (P), medial (Q), and ventral (R) views." figureDoi-0="http://doi.org/10.5281/zenodo.14514573" figureDoi-1="http://doi.org/10.5281/zenodo.14514575" figureDoi-2="http://doi.org/10.5281/zenodo.14514577" figureDoi-3="http://doi.org/10.5281/zenodo.14514579" httpUri-0="https://zenodo.org/record/14514573/files/figure.png" httpUri-1="https://zenodo.org/record/14514575/files/figure.png" httpUri-2="https://zenodo.org/record/14514577/files/figure.png" httpUri-3="https://zenodo.org/record/14514579/files/figure.png" pageId="10" pageNumber="11">Figs 9–12</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="8BCE6C57FFC4B456FCBBF8500F3EF84E" blockId="10.[825,1475,1548,1955]" pageId="10" pageNumber="11">
The available maxillae pertain to different-sized individuals, as can be attested by the drastic size range between the smallest and the largest specimens (see Supporting Information, Figs S3, S4). The most complete maxillae are the 
<typeStatus id="54CAD2F5FFC4B456FA96F8F20923F944" box="[1300,1392,1681,1705]" pageId="10" pageNumber="11" type="holotype">holotype</typeStatus>
(ONM CBI-1-645) and the specimens ONM CBI-1-649 (
<figureCitation id="134A70D2FFC4B456FAD0F8D309F2F925" box="[1362,1441,1712,1736]" captionStart="Figure 9" captionStartId="14.[129,194,1836,1860]" captionTargetBox="[162,1439,151,1799]" captionTargetId="figure-8@14.[156,1446,144,1808]" captionTargetPageId="14" captionText="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="10" pageNumber="11">Figs 9E</figureCitation>
, F, 11J–Q) and ONM CBI-1-654 (
<figureCitation id="134A70D2FFC4B456FBFCF8AC08BAF90A" box="[1150,1257,1743,1767]" captionStart="Figure 11" captionStartId="16.[129,194,1840,1864]" captionTargetBox="[182,1423,146,1809]" captionTargetId="figure-8@16.[177,1425,144,1811]" captionTargetPageId="16" captionText="Figure 11. Maxillae of Terastiodontosaurus marcelosanchezi, small-sized individuals. μCT 3D reconstructions of: A–I, right maxilla ONM CBI-1-654 in labial (A), medial (B), anteromedial (C), ventral (D), ventromedial (E), posteromedial (F), posteroventral (G), dorsal (H), and dorsomedial (I) views; J–Q, right maxilla ONM CBI-1-649 in labial (J), medial (K), posteromedial (L), ventral (M), anteroventral (N), ventromedial (O), dorsal (P), and anterior (Q) views; and R–U, posterior portion of left maxilla ONM CBI-1-653 in labial (R), medial (S), ventral (T), and posteromedial (U) views." figureDoi="http://doi.org/10.5281/zenodo.14514577" httpUri="https://zenodo.org/record/14514577/files/figure.png" pageId="10" pageNumber="11">Fig. 11A–I</figureCitation>
), which are both significantly smaller. In fact, the 
<typeStatus id="54CAD2F5FFC4B456FBDDF88C08E8F8EA" box="[1119,1211,1775,1799]" pageId="10" pageNumber="11" type="holotype">holotype</typeStatus>
ONM CBI-1-645 (
<figureCitation id="134A70D2FFC4B456FA03F88C09EFF8EA" box="[1409,1468,1775,1799]" captionStart="Figure 2" captionStartId="6.[129,194,1839,1863]" captionTargetBox="[155,1448,146,1809]" captionTargetId="figure-75@6.[153,1450,144,1811]" captionTargetPageId="6" captionText="Figure 2. Holotype right maxilla (ONM CBI-1-645) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), dorsal (C), and ventral (D) views." figureDoi="http://doi.org/10.5281/zenodo.14514555" httpUri="https://zenodo.org/record/14514555/files/figure.png" pageId="10" pageNumber="11">Figs 2</figureCitation>
, 3) represents the largest known individual of 
<taxonomicName id="4C7117D4FFC4B456FB7CF96D0F84F8A8" authorityName="Georgalis &amp; Smith" authorityYear="2024" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFC4B456FB7CF96D0F84F8A8" italics="true" pageId="10" pageNumber="11">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
, whereas ONM CBI-1-649 represents one of the smallest among our sample. Nevertheless, the morphology of the maxillae overall is very similar. Here, we focus on comparisons.
</paragraph>
<caption id="DF0E3CDFFFC5B457FFF3F81E0930F95C" ID-DOI="http://doi.org/10.5281/zenodo.14514567" ID-Zenodo-Dep="14514567" httpUri="https://zenodo.org/record/14514567/files/figure.png" pageId="11" pageNumber="12" startId="11.[113,178,1661,1685]" targetBox="[116,1455,146,1632]" targetPageId="11" targetType="figure">
<paragraph id="8BCE6C57FFC5B457FFF3F81E0930F95C" blockId="11.[113,1443,1661,1714]" pageId="11" pageNumber="12">
<emphasis id="B905B045FFC5B457FFF3F81E0C9AF978" bold="true" box="[113,201,1661,1685]" pageId="11" pageNumber="12">Figure 6.</emphasis>
Premaxillae of 
<taxonomicName id="4C7117D4FFC5B457FED8F81E0E95F978" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[346,710,1661,1685]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFC5B457FED8F81E0E95F978" bold="true" box="[346,710,1661,1685]" italics="true" pageId="11" pageNumber="12">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. Photographs of the larger specimens: A–C, ONM CBI-1-711 in anterior (A), right lateral (B), and posterior (C) views; and D–G, ONM CBI-1-658 in anterior (D), posterior (E), dorsal (F), and ventral (G) views.
</paragraph>
</caption>
<paragraph id="8BCE6C57FFC5B450FF0FF8820E0EF939" blockId="11.[113,763,1761,1973]" lastBlockId="12.[129,779,1536,1967]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
The anterodorsally trending premaxillary process is best preserved or complete in ONM CBI-1-654 (
<figureCitation id="134A70D2FFC5B457FDB2F9620EF3F8F4" box="[560,672,1793,1817]" captionStart="Figure 11" captionStartId="16.[129,194,1840,1864]" captionTargetBox="[182,1423,146,1809]" captionTargetId="figure-8@16.[177,1425,144,1811]" captionTargetPageId="16" captionText="Figure 11. Maxillae of Terastiodontosaurus marcelosanchezi, small-sized individuals. μCT 3D reconstructions of: A–I, right maxilla ONM CBI-1-654 in labial (A), medial (B), anteromedial (C), ventral (D), ventromedial (E), posteromedial (F), posteroventral (G), dorsal (H), and dorsomedial (I) views; J–Q, right maxilla ONM CBI-1-649 in labial (J), medial (K), posteromedial (L), ventral (M), anteroventral (N), ventromedial (O), dorsal (P), and anterior (Q) views; and R–U, posterior portion of left maxilla ONM CBI-1-653 in labial (R), medial (S), ventral (T), and posteromedial (U) views." figureDoi="http://doi.org/10.5281/zenodo.14514577" httpUri="https://zenodo.org/record/14514577/files/figure.png" pageId="11" pageNumber="12">Fig. 11A–I</figureCitation>
), ONM CBI-1-649 (
<figureCitation id="134A70D2FFC5B457FF72F9430D6FF8D5" box="[240,316,1824,1848]" captionStart="Figure 9" captionStartId="14.[129,194,1836,1860]" captionTargetBox="[162,1439,151,1799]" captionTargetId="figure-8@14.[156,1446,144,1808]" captionTargetPageId="14" captionText="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="11" pageNumber="12">Figs 9E</figureCitation>
, F, 11J–Q), the 
<typeStatus id="54CAD2F5FFC5B457FE61F9430E6CF8D5" box="[483,575,1824,1848]" pageId="11" pageNumber="12" type="holotype">holotype</typeStatus>
ONM CBI-1-645 (
<figureCitation id="134A70D2FFC5B457FFFEF95C0CE9F8B5" box="[124,186,1855,1880]" captionStart="Figure 2" captionStartId="6.[129,194,1839,1863]" captionTargetBox="[155,1448,146,1809]" captionTargetId="figure-75@6.[153,1450,144,1811]" captionTargetPageId="6" captionText="Figure 2. Holotype right maxilla (ONM CBI-1-645) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), dorsal (C), and ventral (D) views." figureDoi="http://doi.org/10.5281/zenodo.14514555" httpUri="https://zenodo.org/record/14514555/files/figure.png" pageId="11" pageNumber="12">Figs 2</figureCitation>
, 3), and the fragmentary specimens ONM CBI-1-1012 (
<figureCitation id="134A70D2FFC5B457FFFEF93C0CA0F89A" box="[124,243,1887,1911]" captionStart="Figure 12" captionStartId="17.[113,178,1844,1868]" captionTargetBox="[165,1408,151,1814]" captionTargetId="figure-7@17.[163,1410,149,1816]" captionTargetPageId="17" captionText="Figure 12. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, posterior fragment of right maxilla ONM CBI-1-667 in labial (A), medial (B), and ventral (C) views; D–F, posterior fragment of right maxilla ONM CBI-1-1017 in labial (D), ventromedial (E), and ventral (E) views; G–I, anterior fragment of left maxilla ONM CBI-1-1012 in labial (G), medial (H), and ventral (I) views; J–L, anterior fragment of left maxilla ONM CBI-1-1016 in labial (J), medial (K), and ventral (L) views; M–O, anterior fragment of left maxilla ONM CBI-1-1018 in labial (M), medial (N), and ventral (O) views; and P–R, anterior fragment of left maxilla ONM CBI-1-1022 in labial (P), medial (Q), and ventral (R) views." figureDoi="http://doi.org/10.5281/zenodo.14514579" httpUri="https://zenodo.org/record/14514579/files/figure.png" pageId="11" pageNumber="12">Fig. 12G–I</figureCitation>
), ONM CBI-1-1016 (
<figureCitation id="134A70D2FFC5B457FE6CF93C0E0CF89A" box="[494,607,1887,1911]" captionStart="Figure 12" captionStartId="17.[113,178,1844,1868]" captionTargetBox="[165,1408,151,1814]" captionTargetId="figure-7@17.[163,1410,149,1816]" captionTargetPageId="17" captionText="Figure 12. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, posterior fragment of right maxilla ONM CBI-1-667 in labial (A), medial (B), and ventral (C) views; D–F, posterior fragment of right maxilla ONM CBI-1-1017 in labial (D), ventromedial (E), and ventral (E) views; G–I, anterior fragment of left maxilla ONM CBI-1-1012 in labial (G), medial (H), and ventral (I) views; J–L, anterior fragment of left maxilla ONM CBI-1-1016 in labial (J), medial (K), and ventral (L) views; M–O, anterior fragment of left maxilla ONM CBI-1-1018 in labial (M), medial (N), and ventral (O) views; and P–R, anterior fragment of left maxilla ONM CBI-1-1022 in labial (P), medial (Q), and ventral (R) views." figureDoi="http://doi.org/10.5281/zenodo.14514579" httpUri="https://zenodo.org/record/14514579/files/figure.png" pageId="11" pageNumber="12">Fig. 12J–L</figureCitation>
), ONM CBI-1-1018 (
<figureCitation id="134A70D2FFC5B457FF4DF91D0D02F87B" box="[207,337,1918,1942]" captionStart="Figure 12" captionStartId="17.[113,178,1844,1868]" captionTargetBox="[165,1408,151,1814]" captionTargetId="figure-7@17.[163,1410,149,1816]" captionTargetPageId="17" captionText="Figure 12. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, posterior fragment of right maxilla ONM CBI-1-667 in labial (A), medial (B), and ventral (C) views; D–F, posterior fragment of right maxilla ONM CBI-1-1017 in labial (D), ventromedial (E), and ventral (E) views; G–I, anterior fragment of left maxilla ONM CBI-1-1012 in labial (G), medial (H), and ventral (I) views; J–L, anterior fragment of left maxilla ONM CBI-1-1016 in labial (J), medial (K), and ventral (L) views; M–O, anterior fragment of left maxilla ONM CBI-1-1018 in labial (M), medial (N), and ventral (O) views; and P–R, anterior fragment of left maxilla ONM CBI-1-1022 in labial (P), medial (Q), and ventral (R) views." figureDoi="http://doi.org/10.5281/zenodo.14514579" httpUri="https://zenodo.org/record/14514579/files/figure.png" pageId="11" pageNumber="12">Fig. 12M–O</figureCitation>
), and ONM CBI-1-1022 (
<figureCitation id="134A70D2FFC5B457FDF6F91D0EB9F87B" box="[628,746,1918,1942]" captionStart="Figure 12" captionStartId="17.[113,178,1844,1868]" captionTargetBox="[165,1408,151,1814]" captionTargetId="figure-7@17.[163,1410,149,1816]" captionTargetPageId="17" captionText="Figure 12. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, posterior fragment of right maxilla ONM CBI-1-667 in labial (A), medial (B), and ventral (C) views; D–F, posterior fragment of right maxilla ONM CBI-1-1017 in labial (D), ventromedial (E), and ventral (E) views; G–I, anterior fragment of left maxilla ONM CBI-1-1012 in labial (G), medial (H), and ventral (I) views; J–L, anterior fragment of left maxilla ONM CBI-1-1016 in labial (J), medial (K), and ventral (L) views; M–O, anterior fragment of left maxilla ONM CBI-1-1018 in labial (M), medial (N), and ventral (O) views; and P–R, anterior fragment of left maxilla ONM CBI-1-1022 in labial (P), medial (Q), and ventral (R) views." figureDoi="http://doi.org/10.5281/zenodo.14514579" httpUri="https://zenodo.org/record/14514579/files/figure.png" pageId="11" pageNumber="12">Fig. 12P–R</figureCitation>
). The outwardly flaring posterior process of the maxilla is most complete in ONM CBI-1-654 (
<figureCitation id="134A70D2FFC5B457FBFAF88208B5F917" box="[1144,1254,1761,1786]" captionStart="Figure 11" captionStartId="16.[129,194,1840,1864]" captionTargetBox="[182,1423,146,1809]" captionTargetId="figure-8@16.[177,1425,144,1811]" captionTargetPageId="16" captionText="Figure 11. Maxillae of Terastiodontosaurus marcelosanchezi, small-sized individuals. μCT 3D reconstructions of: A–I, right maxilla ONM CBI-1-654 in labial (A), medial (B), anteromedial (C), ventral (D), ventromedial (E), posteromedial (F), posteroventral (G), dorsal (H), and dorsomedial (I) views; J–Q, right maxilla ONM CBI-1-649 in labial (J), medial (K), posteromedial (L), ventral (M), anteroventral (N), ventromedial (O), dorsal (P), and anterior (Q) views; and R–U, posterior portion of left maxilla ONM CBI-1-653 in labial (R), medial (S), ventral (T), and posteromedial (U) views." figureDoi="http://doi.org/10.5281/zenodo.14514577" httpUri="https://zenodo.org/record/14514577/files/figure.png" pageId="11" pageNumber="12">Fig. 11A–I</figureCitation>
), followed in completeness by the 
<typeStatus id="54CAD2F5FFC5B457FC5AF9620867F8F4" box="[984,1076,1793,1817]" pageId="11" pageNumber="12" type="holotype">holotype</typeStatus>
ONM CBI-1-645 (
<figureCitation id="134A70D2FFC5B457FA81F9620912F8F4" box="[1283,1345,1793,1817]" captionStart="Figure 2" captionStartId="6.[129,194,1839,1863]" captionTargetBox="[155,1448,146,1809]" captionTargetId="figure-75@6.[153,1450,144,1811]" captionTargetPageId="6" captionText="Figure 2. Holotype right maxilla (ONM CBI-1-645) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), dorsal (C), and ventral (D) views." figureDoi="http://doi.org/10.5281/zenodo.14514555" httpUri="https://zenodo.org/record/14514555/files/figure.png" pageId="11" pageNumber="12">Figs 2</figureCitation>
, 3), ONM CBI-1-649 (
<figureCitation id="134A70D2FFC5B457FC2BF9430FA7F8D5" box="[937,1012,1824,1848]" captionStart="Figure 9" captionStartId="14.[129,194,1836,1860]" captionTargetBox="[162,1439,151,1799]" captionTargetId="figure-8@14.[156,1446,144,1808]" captionTargetPageId="14" captionText="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="11" pageNumber="12">Figs 9E</figureCitation>
, F, 11J–Q), ONM CBI-1-651 (
<figureCitation id="134A70D2FFC5B457FAB5F94309F1F8D5" box="[1335,1442,1824,1848]" captionStart="Figure 10" captionStartId="15.[113,178,1840,1864]" captionTargetBox="[150,1421,148,1810]" captionTargetId="figure-7@15.[148,1423,145,1812]" captionTargetPageId="15" captionText="Figure 10. Maxillae of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of: A–D, left maxilla ONM CBI-1-648 in labial (A), medial (B), dorsal (C), and ventral (D) views; E–J, right maxilla ONM CBI-1-651 in labial (E), ventral (F), dorsal (G), medial (H), ventromedial (I), and posteroventral (J) views; and K–M, fragment of right maxilla ONM CBI-1-650 in labial (K), medial (L), and ventral (M) views. Note that specimen ONM CBI-1-648 was slightly damaged during micro-computed tomography scanning, hence the difference from the photographs in Figure 9." figureDoi="http://doi.org/10.5281/zenodo.14514575" httpUri="https://zenodo.org/record/14514575/files/figure.png" pageId="11" pageNumber="12">Fig. 10E–J</figureCitation>
), ONM CBI-1-653 (
<figureCitation id="134A70D2FFC5B457FC71F95C083FF8B5" box="[1011,1132,1855,1880]" captionStart="Figure 11" captionStartId="16.[129,194,1840,1864]" captionTargetBox="[182,1423,146,1809]" captionTargetId="figure-8@16.[177,1425,144,1811]" captionTargetPageId="16" captionText="Figure 11. Maxillae of Terastiodontosaurus marcelosanchezi, small-sized individuals. μCT 3D reconstructions of: A–I, right maxilla ONM CBI-1-654 in labial (A), medial (B), anteromedial (C), ventral (D), ventromedial (E), posteromedial (F), posteroventral (G), dorsal (H), and dorsomedial (I) views; J–Q, right maxilla ONM CBI-1-649 in labial (J), medial (K), posteromedial (L), ventral (M), anteroventral (N), ventromedial (O), dorsal (P), and anterior (Q) views; and R–U, posterior portion of left maxilla ONM CBI-1-653 in labial (R), medial (S), ventral (T), and posteromedial (U) views." figureDoi="http://doi.org/10.5281/zenodo.14514577" httpUri="https://zenodo.org/record/14514577/files/figure.png" pageId="11" pageNumber="12">Fig. 11R–U</figureCitation>
), ONM CBI-1-667 (
<figureCitation id="134A70D2FFC5B457FACFF95C0F6FF89A" captionStart="Figure 12" captionStartId="17.[113,178,1844,1868]" captionTargetBox="[165,1408,151,1814]" captionTargetId="figure-7@17.[163,1410,149,1816]" captionTargetPageId="17" captionText="Figure 12. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, posterior fragment of right maxilla ONM CBI-1-667 in labial (A), medial (B), and ventral (C) views; D–F, posterior fragment of right maxilla ONM CBI-1-1017 in labial (D), ventromedial (E), and ventral (E) views; G–I, anterior fragment of left maxilla ONM CBI-1-1012 in labial (G), medial (H), and ventral (I) views; J–L, anterior fragment of left maxilla ONM CBI-1-1016 in labial (J), medial (K), and ventral (L) views; M–O, anterior fragment of left maxilla ONM CBI-1-1018 in labial (M), medial (N), and ventral (O) views; and P–R, anterior fragment of left maxilla ONM CBI-1-1022 in labial (P), medial (Q), and ventral (R) views." figureDoi="http://doi.org/10.5281/zenodo.14514579" httpUri="https://zenodo.org/record/14514579/files/figure.png" pageId="11" pageNumber="12">Fig. 12A– C</figureCitation>
), and ONM CBI-1-1017 (
<figureCitation id="134A70D2FFC5B457FBCCF93C0891F89A" box="[1102,1218,1887,1911]" captionStart="Figure 12" captionStartId="17.[113,178,1844,1868]" captionTargetBox="[165,1408,151,1814]" captionTargetId="figure-7@17.[163,1410,149,1816]" captionTargetPageId="17" captionText="Figure 12. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, posterior fragment of right maxilla ONM CBI-1-667 in labial (A), medial (B), and ventral (C) views; D–F, posterior fragment of right maxilla ONM CBI-1-1017 in labial (D), ventromedial (E), and ventral (E) views; G–I, anterior fragment of left maxilla ONM CBI-1-1012 in labial (G), medial (H), and ventral (I) views; J–L, anterior fragment of left maxilla ONM CBI-1-1016 in labial (J), medial (K), and ventral (L) views; M–O, anterior fragment of left maxilla ONM CBI-1-1018 in labial (M), medial (N), and ventral (O) views; and P–R, anterior fragment of left maxilla ONM CBI-1-1022 in labial (P), medial (Q), and ventral (R) views." figureDoi="http://doi.org/10.5281/zenodo.14514579" httpUri="https://zenodo.org/record/14514579/files/figure.png" pageId="11" pageNumber="12">Fig. 12D–F</figureCitation>
). A medial process that contacts the horizontal wing of the vomer is clearest in the 
<typeStatus id="54CAD2F5FFC5B457FAFEF91D0F0BF85B" pageId="11" pageNumber="12" type="holotype">holotype</typeStatus>
(ONM CBI-1-645) and especially in ONM CBI-1-648 (
<figureCitation id="134A70D2FFC2B450FF09F8630C85F9F5" box="[139,214,1536,1560]" captionStart="Figure 9" captionStartId="14.[129,194,1836,1860]" captionTargetBox="[162,1439,151,1799]" captionTargetId="figure-8@14.[156,1446,144,1808]" captionTargetPageId="14" captionText="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="12" pageNumber="13">Figs 9B</figureCitation>
, C, 10B, C) and ONM CBI-1-649 (
<figureCitation id="134A70D2FFC2B450FDBDF8630ED9F9F5" box="[575,650,1536,1560]" captionStart="Figure 9" captionStartId="14.[129,194,1836,1860]" captionTargetBox="[162,1439,151,1799]" captionTargetId="figure-8@14.[156,1446,144,1808]" captionTargetPageId="14" captionText="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="12" pageNumber="13">Figs 9E</figureCitation>
, F, 11K–Q). The porosity and ridge-like form of the labial surface, observed in the 
<typeStatus id="54CAD2F5FFC2B450FF40F85D0D4DF9BB" box="[194,286,1598,1622]" pageId="12" pageNumber="13" type="holotype">holotype</typeStatus>
ONM CBI-1-645 (
<figureCitation id="134A70D2FFC2B450FE64F85D0E60F9BA" box="[486,563,1598,1623]" captionStart="Figure 2" captionStartId="6.[129,194,1839,1863]" captionTargetBox="[155,1448,146,1809]" captionTargetId="figure-75@6.[153,1450,144,1811]" captionTargetPageId="6" captionText="Figure 2. Holotype right maxilla (ONM CBI-1-645) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), dorsal (C), and ventral (D) views." figureDoi="http://doi.org/10.5281/zenodo.14514555" httpUri="https://zenodo.org/record/14514555/files/figure.png" pageId="12" pageNumber="13">Figs 2A</figureCitation>
, 3A, F), is otherwise evident only in the second largest specimen (ONM CBI-1-651; 
<figureCitation id="134A70D2FFC2B450FF03F81E0C86F978" box="[129,213,1661,1685]" captionStart="Figure 10" captionStartId="15.[113,178,1840,1864]" captionTargetBox="[150,1421,148,1810]" captionTargetId="figure-7@15.[148,1423,145,1812]" captionTargetPageId="15" captionText="Figure 10. Maxillae of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of: A–D, left maxilla ONM CBI-1-648 in labial (A), medial (B), dorsal (C), and ventral (D) views; E–J, right maxilla ONM CBI-1-651 in labial (E), ventral (F), dorsal (G), medial (H), ventromedial (I), and posteroventral (J) views; and K–M, fragment of right maxilla ONM CBI-1-650 in labial (K), medial (L), and ventral (M) views. Note that specimen ONM CBI-1-648 was slightly damaged during micro-computed tomography scanning, hence the difference from the photographs in Figure 9." figureDoi="http://doi.org/10.5281/zenodo.14514575" httpUri="https://zenodo.org/record/14514575/files/figure.png" pageId="12" pageNumber="13">Fig. 10E</figureCitation>
) and is absent in all other smaller specimens, in which this surface is almost completely smooth. This suggests that this feature is subject to size/ontogenetic variation.
</paragraph>
<caption id="DF0E3CDFFFC2B450FF03FBE30D0FFA3D" ID-DOI="http://doi.org/10.5281/zenodo.14514569" ID-Zenodo-Dep="14514569" httpUri="https://zenodo.org/record/14514569/files/figure.png" pageId="12" pageNumber="13" startId="12.[129,194,1408,1432]" targetBox="[131,1471,146,1378]" targetPageId="12" targetType="figure">
<paragraph id="8BCE6C57FFC2B450FF03FBE30D0FFA3D" blockId="12.[129,1473,1407,1488]" pageId="12" pageNumber="13">
<emphasis id="B905B045FFC2B450FF03FBE30C8BFA75" bold="true" box="[129,216,1408,1432]" pageId="12" pageNumber="13">Figure 7.</emphasis>
Premaxillae of 
<taxonomicName id="4C7117D4FFC2B450FEEBFB1C0E86FA7A" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[361,725,1407,1431]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFC2B450FEEBFB1C0E86FA7A" bold="true" box="[361,725,1407,1431]" italics="true" pageId="12" pageNumber="13">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. Photographs of the smaller specimens: A–D, ONM CBI-1-672 in anterior (A), right lateral (B), posterior (C), and dorsal (D) views; and E–H, ONM CBI-1-1021 in anterior (E), right anterolateral (F), posterior (G), right dorsolateral (H) views.
</paragraph>
</caption>
<paragraph id="8BCE6C57FFC2B450FF1EF8B8097BF978" blockId="12.[129,779,1536,1967]" lastBlockId="12.[825,1475,1536,1967]" pageId="12" pageNumber="13">
There are three labial foramina, placed almost in a row, in the 
<typeStatus id="54CAD2F5FFC2B450FF03F8990C8EF8FF" box="[129,221,1786,1810]" pageId="12" pageNumber="13" type="holotype">holotype</typeStatus>
ONM CBI-1-645 (
<figureCitation id="134A70D2FFC2B450FE2BF8990DA4F8FE" box="[425,503,1786,1811]" captionStart="Figure 2" captionStartId="6.[129,194,1839,1863]" captionTargetBox="[155,1448,146,1809]" captionTargetId="figure-75@6.[153,1450,144,1811]" captionTargetPageId="6" captionText="Figure 2. Holotype right maxilla (ONM CBI-1-645) of Terastiodontosaurus marcelosanchezi. Photographs of the specimen in labial (A), medial (B), dorsal (C), and ventral (D) views." figureDoi="http://doi.org/10.5281/zenodo.14514555" httpUri="https://zenodo.org/record/14514555/files/figure.png" pageId="12" pageNumber="13">Figs 2A</figureCitation>
, 3A) and in ONM CBI-1- 649 (
<figureCitation id="134A70D2FFC2B450FF38F9790D58F8DF" box="[186,267,1818,1842]" captionStart="Figure 9" captionStartId="14.[129,194,1836,1860]" captionTargetBox="[162,1439,151,1799]" captionTargetId="figure-8@14.[156,1446,144,1808]" captionTargetPageId="14" captionText="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="12" pageNumber="13">Figs 9G</figureCitation>
, 11J); in both these specimens, the two anterior foramina are more closely spaced. ONM CBI-1-648 also has three labial foramina (
<figureCitation id="134A70D2FFC2B450FEE5F93B0DE2F89C" box="[359,433,1880,1905]" captionStart="Figure 9" captionStartId="14.[129,194,1836,1860]" captionTargetBox="[162,1439,151,1799]" captionTargetId="figure-8@14.[156,1446,144,1808]" captionTargetPageId="14" captionText="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="12" pageNumber="13">Fig. 9A</figureCitation>
), but the posterior two foramina are more closely spaced. In ONM CBI-1-649, there is also a further foramen situated dorsal to the row of the three foramina, situated approximately above the first foramen. Even more, in ONM CBI-1-654, there are two small foramina above the two of three foramina (
<figureCitation id="134A70D2FFC2B450FB83F85D0809F9BA" box="[1025,1114,1598,1623]" captionStart="Figure 11" captionStartId="16.[129,194,1840,1864]" captionTargetBox="[182,1423,146,1809]" captionTargetId="figure-8@16.[177,1425,144,1811]" captionTargetPageId="16" captionText="Figure 11. Maxillae of Terastiodontosaurus marcelosanchezi, small-sized individuals. μCT 3D reconstructions of: A–I, right maxilla ONM CBI-1-654 in labial (A), medial (B), anteromedial (C), ventral (D), ventromedial (E), posteromedial (F), posteroventral (G), dorsal (H), and dorsomedial (I) views; J–Q, right maxilla ONM CBI-1-649 in labial (J), medial (K), posteromedial (L), ventral (M), anteroventral (N), ventromedial (O), dorsal (P), and anterior (Q) views; and R–U, posterior portion of left maxilla ONM CBI-1-653 in labial (R), medial (S), ventral (T), and posteromedial (U) views." figureDoi="http://doi.org/10.5281/zenodo.14514577" httpUri="https://zenodo.org/record/14514577/files/figure.png" pageId="12" pageNumber="13">Fig. 11A</figureCitation>
). The number of foramina cannot be assessed fully in the remaining incomplete maxillae. The superior alveolar foramen is usually relatively large.
</paragraph>
<paragraph id="8BCE6C57FFC2B44EFCD7F8FF0EABFD93" blockId="12.[825,1475,1536,1967]" lastBlockId="18.[128,779,144,1077]" lastPageId="18" lastPageNumber="19" pageId="12" pageNumber="13">
Similar to the 
<typeStatus id="54CAD2F5FFC2B450FC6AF8FF0817F959" box="[1000,1092,1692,1716]" pageId="12" pageNumber="13" type="holotype">holotype</typeStatus>
(ONM CBI-1-645), maxillae almost always bear three teeth, of which the second is by far the largest and, in fact, covers much of the ventral portion of the bone; this tooth is followed in size by the anteriormost tooth, while the posteriormost tooth is tiny. However, there are two notable exceptions, denoting some degree of variability: in ONM CBI-1-651, there is a fourth tiny tooth located posteromedially to the third tooth, not in line with it (
<figureCitation id="134A70D2FFC2B450FB3CF91B0941F87D" box="[1214,1298,1912,1936]" captionStart="Figure 10" captionStartId="15.[113,178,1840,1864]" captionTargetBox="[150,1421,148,1810]" captionTargetId="figure-7@15.[148,1423,145,1812]" captionTargetPageId="15" captionText="Figure 10. Maxillae of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of: A–D, left maxilla ONM CBI-1-648 in labial (A), medial (B), dorsal (C), and ventral (D) views; E–J, right maxilla ONM CBI-1-651 in labial (E), ventral (F), dorsal (G), medial (H), ventromedial (I), and posteroventral (J) views; and K–M, fragment of right maxilla ONM CBI-1-650 in labial (K), medial (L), and ventral (M) views. Note that specimen ONM CBI-1-648 was slightly damaged during micro-computed tomography scanning, hence the difference from the photographs in Figure 9." figureDoi="http://doi.org/10.5281/zenodo.14514575" httpUri="https://zenodo.org/record/14514575/files/figure.png" pageId="12" pageNumber="13">Fig. 10F</figureCitation>
), while in ONM CBI-1-649 the tiniest tooth is absent, meaning that specimen bears only two teeth (
<figureCitation id="134A70D2FFDCB44EFEE5FEF30DEBFF45" box="[359,440,144,168]" captionStart="Figure 9" captionStartId="14.[129,194,1836,1860]" captionTargetBox="[162,1439,151,1799]" captionTargetId="figure-8@14.[156,1446,144,1808]" captionTargetPageId="14" captionText="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="18" pageNumber="19">Figs 9G</figureCitation>
, 11M–O). A further, interesting variation occurs also in the right maxilla fragment ONM CBI-1-650, where the anterior (smaller) preserved tooth is different from the same tooth in other specimens, being extremely narrow (
<figureCitation id="134A70D2FFDCB44EFF09FF6E0C8DFEC8" box="[139,222,269,293]" captionStart="Figure 9" captionStartId="14.[129,194,1836,1860]" captionTargetBox="[162,1439,151,1799]" captionTargetId="figure-8@14.[156,1446,144,1808]" captionTargetPageId="14" captionText="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="18" pageNumber="19">Figs 9D</figureCitation>
, 10K–M). Tentatively, we attribute this variation as being intraspecific (or even ontogenetic). This could indeed be the case, taking into consideration that especially if the more anterior teeth are being ‘de-emphasized’ in favour of the huge tooth (which serves as the prey ‘cracker’), and thus the anterior teeth perhaps become more of a remnant/vestige, then one would expect a greater variation. Moreover, again in the same specimen, there seems to be a considerable gap between the two preserved teeth (
<figureCitation id="134A70D2FFDCB44EFF4AFC6B0D49FDCD" box="[200,282,520,544]" captionStart="Figure 9" captionStartId="14.[129,194,1836,1860]" captionTargetBox="[162,1439,151,1799]" captionTargetId="figure-8@14.[156,1446,144,1808]" captionTargetPageId="14" captionText="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="18" pageNumber="19">Figs 9D</figureCitation>
, 10K–M); this is unusual, because in most remaining maxillae and dentaries, the teeth are closely appressed, with no interdental gaps between them (but in ONM CBI-1-653 the third tooth is somewhat more widely separated; 
<figureCitation id="134A70D2FFDCB44EFD12FC050EB4FD93" box="[656,743,614,638]" captionStart="Figure 11" captionStartId="16.[129,194,1840,1864]" captionTargetBox="[182,1423,146,1809]" captionTargetId="figure-8@16.[177,1425,144,1811]" captionTargetPageId="16" captionText="Figure 11. Maxillae of Terastiodontosaurus marcelosanchezi, small-sized individuals. μCT 3D reconstructions of: A–I, right maxilla ONM CBI-1-654 in labial (A), medial (B), anteromedial (C), ventral (D), ventromedial (E), posteromedial (F), posteroventral (G), dorsal (H), and dorsomedial (I) views; J–Q, right maxilla ONM CBI-1-649 in labial (J), medial (K), posteromedial (L), ventral (M), anteroventral (N), ventromedial (O), dorsal (P), and anterior (Q) views; and R–U, posterior portion of left maxilla ONM CBI-1-653 in labial (R), medial (S), ventral (T), and posteromedial (U) views." figureDoi="http://doi.org/10.5281/zenodo.14514577" httpUri="https://zenodo.org/record/14514577/files/figure.png" pageId="18" pageNumber="19">Fig. 11T</figureCitation>
).
</paragraph>
<caption id="DF0E3CDFFFC3B451FFF3F94C0E6BF876" ID-DOI="http://doi.org/10.5281/zenodo.14514571" ID-Zenodo-Dep="14514571" httpUri="https://zenodo.org/record/14514571/files/figure.png" pageId="13" pageNumber="14" startId="13.[113,178,1839,1863]" targetBox="[165,1408,146,1809]" targetPageId="13" targetType="figure">
<paragraph id="8BCE6C57FFC3B451FFF3F94C0E6BF876" blockId="13.[113,1431,1838,1947]" pageId="13" pageNumber="14">
<emphasis id="B905B045FFC3B451FFF3F94C0C9AF8AA" bold="true" box="[113,201,1839,1863]" pageId="13" pageNumber="14">Figure 8.</emphasis>
Premaxillae of 
<taxonomicName id="4C7117D4FFC3B451FED8F94D0E95F8AB" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[346,710,1838,1862]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFC3B451FED8F94D0E95F8AB" bold="true" box="[346,710,1838,1862]" italics="true" pageId="13" pageNumber="14">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. μCT 3D reconstructions of: A–F, ONM CBI-1-711 in anterior (A), right lateral (B), left ventrolateral (C), posterior (D), ventral (E), and dorsal (F) views; G–L, ONM CBI-1-658 in anterior (G), right lateral (H), left lateral (I), posterior (J), dorsal (K), and ventral (L) views; and M–R, ONM CBI-1-672 in anterior (M), right lateral (N), left lateral (O), posterior (P), ventral (Q), and dorsal (R) views.
</paragraph>
</caption>
<caption id="DF0E3CDFFFC0B452FF03F94F0ECFF891" ID-DOI="http://doi.org/10.5281/zenodo.14514573" ID-Zenodo-Dep="14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="14" pageNumber="15" startId="14.[129,194,1836,1860]" targetBox="[162,1439,151,1799]" targetPageId="14" targetType="figure">
<paragraph id="8BCE6C57FFC0B452FF03F94F0ECFF891" blockId="14.[129,1456,1836,1916]" pageId="14" pageNumber="15">
<emphasis id="B905B045FFC0B452FF03F94F0C8BF8A9" bold="true" box="[129,216,1836,1860]" pageId="14" pageNumber="15">Figure 9.</emphasis>
Maxillae of 
<taxonomicName id="4C7117D4FFC0B452FEC9F94F0EE4F8A9" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[331,695,1836,1860]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFC0B452FEC9F94F0EE4F8A9" bold="true" box="[331,695,1836,1860]" italics="true" pageId="14" pageNumber="15">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views.
</paragraph>
</caption>
<caption id="DF0E3CDFFFC1B453FFF3F9530DFCF855" ID-DOI="http://doi.org/10.5281/zenodo.14514575" ID-Zenodo-Dep="14514575" httpUri="https://zenodo.org/record/14514575/files/figure.png" pageId="15" pageNumber="16" startId="15.[113,178,1840,1864]" targetBox="[150,1421,148,1810]" targetPageId="15" targetType="figure">
<paragraph id="8BCE6C57FFC1B453FFF3F9530DFCF855" blockId="15.[113,1425,1840,1977]" pageId="15" pageNumber="16">
<emphasis id="B905B045FFC1B453FFF3F9530C85F8A5" bold="true" box="[113,214,1840,1864]" pageId="15" pageNumber="16">Figure 10.</emphasis>
Maxillae of 
<taxonomicName id="4C7117D4FFC1B453FECBF9530EE6F8A5" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[329,693,1840,1864]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFC1B453FECBF9530EE6F8A5" bold="true" box="[329,693,1840,1864]" italics="true" pageId="15" pageNumber="16">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. μCT 3D reconstructions of: A–D, left maxilla ONM CBI-1-648 in labial (A), medial (B), dorsal (C), and ventral (D) views; E–J, right maxilla ONM CBI-1-651 in labial (E), ventral (F), dorsal (G), medial (H), ventromedial (I), and posteroventral (J) views; and K–M, fragment of right maxilla ONM CBI-1-650 in labial (K), medial (L), and ventral (M) views. Note that specimen ONM CBI-1-648 was slightly damaged during micro-computed tomography scanning, hence the difference from the photographs in Figure 9.
</paragraph>
</caption>
<caption id="DF0E3CDFFFDEB44CFF03F9530E43F855" ID-DOI="http://doi.org/10.5281/zenodo.14514577" ID-Zenodo-Dep="14514577" httpUri="https://zenodo.org/record/14514577/files/figure.png" pageId="16" pageNumber="17" startId="16.[129,194,1840,1864]" targetBox="[182,1423,146,1809]" targetPageId="16" targetType="figure">
<paragraph id="8BCE6C57FFDEB44CFF03F9530E43F855" blockId="16.[129,1448,1840,1976]" pageId="16" pageNumber="17">
<emphasis id="B905B045FFDEB44CFF03F9530CB6F8A5" bold="true" box="[129,229,1840,1864]" pageId="16" pageNumber="17">Figure 11.</emphasis>
Maxillae of 
<taxonomicName id="4C7117D4FFDEB44CFEDAF9530E97F8A5" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[344,708,1840,1864]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFDEB44CFEDAF9530E97F8A5" bold="true" box="[344,708,1840,1864]" italics="true" pageId="16" pageNumber="17">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
, small-sized individuals. μCT 3D reconstructions of: A–I, right maxilla ONM CBI-1-654 in labial (A), medial (B), anteromedial (C), ventral (D), ventromedial (E), posteromedial (F), posteroventral (G), dorsal (H), and dorsomedial (I) views; J–Q, right maxilla ONM CBI-1-649 in labial (J), medial (K), posteromedial (L), ventral (M), anteroventral (N), ventromedial (O), dorsal (P), and anterior (Q) views; and R–U, posterior portion of left maxilla ONM CBI-1-653 in labial (R), medial (S), ventral (T), and posteromedial (U) views.
</paragraph>
</caption>
<caption id="DF0E3CDFFFDFB44DFFF3F9570D20F835" ID-DOI="http://doi.org/10.5281/zenodo.14514579" ID-Zenodo-Dep="14514579" httpUri="https://zenodo.org/record/14514579/files/figure.png" pageId="17" pageNumber="18" startId="17.[113,178,1844,1868]" targetBox="[165,1408,151,1814]" targetPageId="17" targetType="figure">
<paragraph id="8BCE6C57FFDFB44DFFF3F9570D20F835" blockId="17.[113,1443,1844,2008]" pageId="17" pageNumber="18">
<emphasis id="B905B045FFDFB44DFFF3F9570C85F8A1" bold="true" box="[113,214,1844,1868]" pageId="17" pageNumber="18">Figure 12.</emphasis>
Maxillae of 
<taxonomicName id="4C7117D4FFDFB44DFECBF9570EE6F8A1" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[329,693,1844,1868]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFDFB44DFECBF9570EE6F8A1" bold="true" box="[329,693,1844,1868]" italics="true" pageId="17" pageNumber="18">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. Photographs of: A–C, posterior fragment of right maxilla ONM CBI-1-667 in labial (A), medial (B), and ventral (C) views; D–F, posterior fragment of right maxilla ONM CBI-1-1017 in labial (D), ventromedial (E), and ventral (E) views; G–I, anterior fragment of left maxilla ONM CBI-1-1012 in labial (G), medial (H), and ventral (I) views; J–L, anterior fragment of left maxilla ONM CBI-1-1016 in labial (J), medial (K), and ventral (L) views; M–O, anterior fragment of left maxilla ONM CBI-1-1018 in labial (M), medial (N), and ventral (O) views; and P–R, anterior fragment of left maxilla ONM CBI-1-1022 in labial (P), medial (Q), and ventral (R) views.
</paragraph>
</caption>
<paragraph id="8BCE6C57FFDCB44EFF1EFCE60F57FD16" blockId="18.[128,779,144,1077]" pageId="18" pageNumber="19">
Notably also, in some cases, there is a distinct ‘hill’ forming on the smallest tooth or teeth. This is the case with ONM CBI-1-651 (where the ‘hill’ is prominent in both the small third and fourth teeth; 
<figureCitation id="134A70D2FFDCB44EFE85FC800D0FFD16" box="[263,348,739,763]" captionStart="Figure 10" captionStartId="15.[113,178,1840,1864]" captionTargetBox="[150,1421,148,1810]" captionTargetId="figure-7@15.[148,1423,145,1812]" captionTargetPageId="15" captionText="Figure 10. Maxillae of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of: A–D, left maxilla ONM CBI-1-648 in labial (A), medial (B), dorsal (C), and ventral (D) views; E–J, right maxilla ONM CBI-1-651 in labial (E), ventral (F), dorsal (G), medial (H), ventromedial (I), and posteroventral (J) views; and K–M, fragment of right maxilla ONM CBI-1-650 in labial (K), medial (L), and ventral (M) views. Note that specimen ONM CBI-1-648 was slightly damaged during micro-computed tomography scanning, hence the difference from the photographs in Figure 9." figureDoi="http://doi.org/10.5281/zenodo.14514575" httpUri="https://zenodo.org/record/14514575/files/figure.png" pageId="18" pageNumber="19">Fig. 10E</figureCitation>
, I, J) and ONM CBI-1-653 (
<figureCitation id="134A70D2FFDCB44EFD03FC800E96FD16" box="[641,709,739,763]" captionStart="Figure 11" captionStartId="16.[129,194,1840,1864]" captionTargetBox="[182,1423,146,1809]" captionTargetId="figure-8@16.[177,1425,144,1811]" captionTargetPageId="16" captionText="Figure 11. Maxillae of Terastiodontosaurus marcelosanchezi, small-sized individuals. μCT 3D reconstructions of: A–I, right maxilla ONM CBI-1-654 in labial (A), medial (B), anteromedial (C), ventral (D), ventromedial (E), posteromedial (F), posteroventral (G), dorsal (H), and dorsomedial (I) views; J–Q, right maxilla ONM CBI-1-649 in labial (J), medial (K), posteromedial (L), ventral (M), anteroventral (N), ventromedial (O), dorsal (P), and anterior (Q) views; and R–U, posterior portion of left maxilla ONM CBI-1-653 in labial (R), medial (S), ventral (T), and posteromedial (U) views." figureDoi="http://doi.org/10.5281/zenodo.14514577" httpUri="https://zenodo.org/record/14514577/files/figure.png" pageId="18" pageNumber="19">Fig. 11</figureCitation>
S–U).
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFF1EFD600D6EFC3A" blockId="18.[128,779,144,1077]" pageId="18" pageNumber="19">
Several nutritive foramina are always present at the base of the teeth in all specimens, in the remnants of the subdental gutter; however, their number and size are variable. Usually, these foramina are mostly at the medial side of the maxilla; however, there is also some variation: in ONM CBI-1-650, they are equally present in both labial and medial aspects of the bone (
<figureCitation id="134A70D2FFDCB44EFF09FDDC0C89FC3A" box="[139,218,959,983]" captionStart="Figure 9" captionStartId="14.[129,194,1836,1860]" captionTargetBox="[162,1439,151,1799]" captionTargetId="figure-8@14.[156,1446,144,1808]" captionTargetPageId="14" captionText="Figure 9. Maxillae of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left maxilla ONM CBI-1-648 in labial (A), medial (B), and dorsal (C) views; D, fragment of right maxilla ONM CBI-1-650 in medial view; and E–G, right maxilla ONM CBI-1-649 of a small-sized individual in dorsal (E), medial (F), and labial (G) views." figureDoi="http://doi.org/10.5281/zenodo.14514573" httpUri="https://zenodo.org/record/14514573/files/figure.png" pageId="18" pageNumber="19">Figs 9D</figureCitation>
, 10K, L).
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFF1EFDBD0F59FBD8" blockId="18.[128,779,144,1077]" pageId="18" pageNumber="19">
In order to facilitate quantitative investigation, we introduce the ratio of largest tooth length on the maxilla to total tooth row length of the maxilla. This was complete in only 
<specimenCount id="9D77A7DEFFDCB44EFDE5FA7E0F55FBD8" box="[615,774,1053,1077]" count="4" pageId="18" pageNumber="19" type="generic">four specimens</specimenCount>
:
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFF0DFA380E04FB7F" blockId="18.[143,778,1115,1359]" pageId="18" pageNumber="19">
<typeStatus id="54CAD2F5FFDCB44EFF0DFA380CBEFB9E" box="[143,237,1115,1139]" pageId="18" pageNumber="19" type="holotype">Holotype</typeStatus>
, ONM CBI-1-645: largest tooth length, 
<quantity id="4C89C1B2FFDCB44EFD2DFA380F52FB9E" box="[687,769,1115,1139]" metricMagnitude="-3" metricUnit="m" metricValue="4.9" pageId="18" pageNumber="19" unit="mm" value="4.9">4.9 mm</quantity>
/ total tooth row length, 
<quantity id="4C89C1B2FFDCB44EFE12FA190DB2FB7F" box="[400,481,1146,1170]" metricMagnitude="-3" metricUnit="m" metricValue="9.5" pageId="18" pageNumber="19" unit="mm" value="9.5">9.5 mm</quantity>
; ratio, 0.52.
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFF0DFAF90DB5FB3C" blockId="18.[143,778,1115,1359]" pageId="18" pageNumber="19">
ONM CBI-1-649: largest tooth length, 
<quantity id="4C89C1B2FFDCB44EFDBFFAF90EDEFB5F" box="[573,653,1178,1202]" metricMagnitude="-3" metricUnit="m" metricValue="2.6" pageId="18" pageNumber="19" unit="mm" value="2.6">2.6 mm</quantity>
/total tooth row length, 
<quantity id="4C89C1B2FFDCB44EFE9DFADA0D23FB3C" box="[287,368,1209,1233]" metricMagnitude="-3" metricUnit="m" metricValue="3.7" pageId="18" pageNumber="19" unit="mm" value="3.7">3.7 mm</quantity>
; ratio, 0.70.
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFF0DFABA0DB5FAFD" blockId="18.[143,778,1115,1359]" pageId="18" pageNumber="19">
ONM CBI-1-651: largest tooth length, 
<quantity id="4C89C1B2FFDCB44EFDBFFABA0EDEFB1C" box="[573,653,1241,1265]" metricMagnitude="-3" metricUnit="m" metricValue="3.7" pageId="18" pageNumber="19" unit="mm" value="3.7">3.7 mm</quantity>
/total tooth row length, 
<quantity id="4C89C1B2FFDCB44EFE9DFA9B0D23FAE2" box="[287,368,1272,1296]" metricMagnitude="-3" metricUnit="m" metricValue="7.3" pageId="18" pageNumber="19" unit="mm" value="7.3">7.3 mm</quantity>
; ratio, 0.51.
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFF0DFB740DB5FAA3" blockId="18.[143,778,1115,1359]" pageId="18" pageNumber="19">
ONM CBI-1-654: largest tooth length, 
<quantity id="4C89C1B2FFDCB44EFDBFFB740EDEFAC2" box="[573,653,1303,1327]" metricMagnitude="-3" metricUnit="m" metricValue="2.3" pageId="18" pageNumber="19" unit="mm" value="2.3">2.3 mm</quantity>
/total tooth row length, 
<quantity id="4C89C1B2FFDCB44EFE9DFB540D23FAA3" box="[287,368,1335,1359]" metricMagnitude="-3" metricUnit="m" metricValue="4.1" pageId="18" pageNumber="19" unit="mm" value="4.1">4.1 mm</quantity>
; ratio, 0.56.
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFF1EFB160E69FA40" blockId="18.[129,777,1397,1453]" pageId="18" pageNumber="19">Besides, some information on this ratio can also be gleaned tentatively from some incomplete maxillae:</paragraph>
<paragraph id="8BCE6C57FFDCB44EFF0DFBB00CBAF9C7" blockId="18.[143,778,1491,1672]" pageId="18" pageNumber="19">
ONM CBI-1-650: largest tooth length, 
<quantity id="4C89C1B2FFDCB44EFDC9FBB00EC9FA06" box="[587,666,1491,1515]" metricMagnitude="-3" metricUnit="m" metricValue="4.8" pageId="18" pageNumber="19" unit="mm" value="4.8">4.8 mm</quantity>
/preserved tooth row length (incomplete), 
<quantity id="4C89C1B2FFDCB44EFD87FB900E05F9E7" box="[517,598,1523,1547]" metricMagnitude="-3" metricUnit="m" metricValue="7.8" pageId="18" pageNumber="19" unit="mm" value="7.8">7.8 mm</quantity>
; estimated ratio, &lt;0.62.
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFF0DF8520CBAF965" blockId="18.[143,778,1491,1672]" pageId="18" pageNumber="19">
ONM CBI-1-648: largest tooth length, 
<quantity id="4C89C1B2FFDCB44EFDC9F8520EC9F9A4" box="[587,666,1585,1609]" metricMagnitude="-3" metricUnit="m" metricValue="3.2" pageId="18" pageNumber="19" unit="mm" value="3.2">3.2 mm</quantity>
/preserved tooth row length (incomplete), 
<quantity id="4C89C1B2FFDCB44EFD87F8330E05F985" box="[517,598,1616,1640]" metricMagnitude="-3" metricUnit="m" metricValue="5.6" pageId="18" pageNumber="19" unit="mm" value="5.6">5.6 mm</quantity>
; estimated ratio, &lt;0.57.
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFED7F8CB0E66F92F" blockId="18.[127,779,1703,1986]" box="[341,565,1703,1730]" pageId="18" pageNumber="19">
<emphasis id="B905B045FFDCB44EFED7F8CB0E66F92F" box="[341,565,1703,1730]" italics="true" pageId="18" pageNumber="19">
Dentaries (
<figureCitation id="134A70D2FFDCB44EFE3CF8CB0E78F92C" box="[446,555,1703,1730]" captionStart-0="Figure 13" captionStart-1="Figure 14" captionStart-2="Figure 15" captionStart-3="Figure 16" captionStartId-0="19.[113,178,1288,1312]" captionStartId-1="20.[129,194,1626,1650]" captionStartId-2="21.[113,178,1840,1864]" captionStartId-3="22.[129,194,1840,1864]" captionTargetBox-0="[116,1456,146,1258]" captionTargetBox-1="[136,1459,156,1585]" captionTargetBox-2="[162,1409,147,1810]" captionTargetBox-3="[180,1423,146,1810]" captionTargetId-0="figure-358@19.[114,1458,144,1260]" captionTargetId-1="figure-160@20.[129,1473,144,1599]" captionTargetId-2="figure-6@21.[160,1411,145,1812]" captionTargetId-3="figure-7@22.[178,1425,144,1812]" captionTargetPageId-0="19" captionTargetPageId-1="20" captionTargetPageId-2="21" captionTargetPageId-3="22" captionText-0="Figure 13. Dentary of Terastiodontosaurus marcelosanchezi. Photographs of left dentary ONM CBI-1-647 in labial (A) and medial (B) views." captionText-1="Figure 14. Dentaries of Terastiodontosaurus marcelosanchezi. Photographs of left dentary ONM CBI-1-657 in medial view (A); left dentary ONM CBI-1-655 in labial view (B); and right dentary ONM CBI-1-656 in medial view (C)." captionText-2="Figure 15. Dentaries of Terastiodontosaurus marcelosanchezi. μCT 3D reconstructions of: A–C, left dentary ONM CBI-1-657 in labial (A), medial (B), and dorsal (C) views; D–F, left dentary ONM CBI-1-659 in labial (D), medial (E), and dorsal (F) views; G–I, right dentary ONM CBI-1-660 in labial (G), medial (H), and dorsal (I) views; J–L, left dentary ONM CBI-1-661 in labial (J), medial (K), and dorsal (L) views; M–O, left dentary ONM CBI-1-668 in labial (M), medial (N), and dorsal (O) views; and P–S, left dentary ONM CBI-1-670 in labial (P), medial (Q), dorsal (R), and posterior (S) views." captionText-3="Figure 16. Dentaries of Terastiodontosaurus marcelosanchezi. Photographs of: A–C, left dentary ONM CBI-1-662 in labial (A), medial (B), and dorsal (C) views; D, E, right dentary ONM CBI-1-1013 in labial (D) and dorsal (E) views; F–H, right dentary ONM CBI-1-666 in labial (F), dorsolabial (G), and medial (H) views; I–K, anterior fragment of right dentary ONM CBI-1-1020 in labial (I), dorsal (J), and medial (K) views; L–N, anterior fragment of left dentary ONM CBI-1-1015 in labial (L), dorsal (M), and medial (N) views; and O–Q, anterior fragment of left dentary ONM CBI-1-1014 in dorsal (O), labial (P), and medial (Q) views." figureDoi-0="http://doi.org/10.5281/zenodo.14514581" figureDoi-1="http://doi.org/10.5281/zenodo.14514583" figureDoi-2="http://doi.org/10.5281/zenodo.14514585" figureDoi-3="http://doi.org/10.5281/zenodo.14514587" httpUri-0="https://zenodo.org/record/14514581/files/figure.png" httpUri-1="https://zenodo.org/record/14514583/files/figure.png" httpUri-2="https://zenodo.org/record/14514585/files/figure.png" httpUri-3="https://zenodo.org/record/14514587/files/figure.png" pageId="18" pageNumber="19">Figs 13–16</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFF03F8AD0F97FEC8" blockId="18.[127,779,1703,1986]" lastBlockId="18.[824,1477,144,1703]" pageId="18" pageNumber="19">
Apart from the 
<typeStatus id="54CAD2F5FFDCB44EFE98F8AD0D20F90B" box="[282,371,1742,1766]" pageId="18" pageNumber="19" type="paratype">paratype</typeStatus>
dentary ONM CBI-1-646, all remaining dentaries are rather incomplete. The available sample denotes a range of sizes, but substantially less disparate than the maxillary sample (cf. Supporting Information, Figs S3–S6). The 
<typeStatus id="54CAD2F5FFDCB44EFD32F94F0F5AF8A9" box="[688,777,1836,1860]" pageId="18" pageNumber="19" type="paratype">paratype</typeStatus>
ONM CBI-1-646 represents one of the largest individuals, with the fragmentary dentary ONM CBI-1-659 pertaining to a more or less similar size. All dentaries closely approach in overall morphology the 
<typeStatus id="54CAD2F5FFDCB44EFEB7F9C90DDDF82F" box="[309,398,1962,1986]" pageId="18" pageNumber="19" type="paratype">paratype</typeStatus>
ONM CBI-1-646 described above. Some specimens are nevertheless highly incomplete, sometimes preserving only the anterior portion of the dentary (e.g. ONM CBI-1-1014, ONM CBI-1-1015, and ONM CBI-1-1020), while ONM CBI-1-664 is only a fragment of the coronoid process of a left dentary.
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFCD7FF4E0990FBB9" blockId="18.[824,1477,144,1703]" pageId="18" pageNumber="19">
The tooth row is complete only in the 
<typeStatus id="54CAD2F5FFDCB44EFB55FF4E0963FEA8" box="[1239,1328,301,325]" pageId="18" pageNumber="19" type="paratype">paratype</typeStatus>
ONM CBI-1- 646, where it comprises eight acrodont teeth, the largest being the fourth one (counting from posteriorly). Otherwise, the tooth row is almost complete in ONM CBI-1-657 (
<figureCitation id="134A70D2FFDCB44EFA80FFE80908FE4E" box="[1282,1371,395,419]" captionStart="Figure 14" captionStartId="20.[129,194,1626,1650]" captionTargetBox="[136,1459,156,1585]" captionTargetId="figure-160@20.[129,1473,144,1599]" captionTargetPageId="20" captionText="Figure 14. Dentaries of Terastiodontosaurus marcelosanchezi. Photographs of left dentary ONM CBI-1-657 in medial view (A); left dentary ONM CBI-1-655 in labial view (B); and right dentary ONM CBI-1-656 in medial view (C)." figureDoi="http://doi.org/10.5281/zenodo.14514583" httpUri="https://zenodo.org/record/14514583/files/figure.png" pageId="18" pageNumber="19">Figs 14A</figureCitation>
, 15A–C), which preserves all but the seventh tooth (counting from posteriorly). In that specimen also, the fourth tooth is the largest one and the posteriormost tooth is the tiniest one (both counting from posteriorly). One important difference in ONM CBI-1-657 is that there appears to be a dental gap between the anteriormost tooth and the succeeding tooth position (although we cannot be certain that this is not an artefact). Otherwise, in all specimens, all teeth are almost adjoined, with almost no interdental gaps between them (at maximum only tiny empty spaces between some, but not all teeth, do exist). As in the 
<typeStatus id="54CAD2F5FFDCB44EFB28FCA70950FD31" box="[1194,1283,708,732]" pageId="18" pageNumber="19" type="paratype">paratype</typeStatus>
ONM CBI-1-646, also in ONM CBI-1-647, ONM CBI-1-655, ONM CBI-1-657, ONM CBI-1-662, ONM CBI-1-666, ONM CBI-1-1014, ONM CBI-1-1015, and ONM CBI-1-1020, the anteriormost tooth is more bulbous and dorsoventrally high and projects beyond the anterior surface of the symphysis. This is also the case for the second anteriormost tooth in the 
<typeStatus id="54CAD2F5FFDCB44EFB21FDE308AFFC75" box="[1187,1276,896,920]" pageId="18" pageNumber="19" type="paratype">paratype</typeStatus>
ONM CBI-1-646, as in ONM CBI-1-647, ONM CBI-1-655, ONM CBI-1-662, ONM CBI-1-666, ONM CBI-1-1014, ONM CBI-1-1015, and ONM CBI-1-1020, which is also relatively bulbous and dorsoventrally tall. In dentary ONM CBI-1-670, the teeth are not that flattened but apparently represent the anterior teeth [second, third, or fourth (counting from anteriorly)], near the symphysis.
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFCD7FA3F08ABFB5F" blockId="18.[824,1477,144,1703]" pageId="18" pageNumber="19">As in the maxillae, there is a distinct ‘hill’ on the tiny teeth (e.g. ONM CBI-1-646 and ONM CBI-1-657). Great enamel thickness is also found on the dentary teeth.</paragraph>
<paragraph id="8BCE6C57FFDCB44EFCD7FADA0FC2FAA2" blockId="18.[824,1477,144,1703]" pageId="18" pageNumber="19">Nutritive foramina are open at the base of various teeth above the subdental shelf in all dentaries; their number is not consistent, and it can vary between tooth positions and individuals at the same position. In ONM CBI-1-657, these are poorly developed.</paragraph>
<paragraph id="8BCE6C57FFDCB44EFCD7FB350944F9C7" blockId="18.[824,1477,144,1703]" pageId="18" pageNumber="19">
Meckel’s groove is open in all specimens where this can be studied, most notably ONM CBI-1-660. Owing to their incompleteness, besides the 
<typeStatus id="54CAD2F5FFDCB44EFB94FBF6083CFA40" box="[1046,1135,1429,1453]" pageId="18" pageNumber="19" type="paratype">paratype</typeStatus>
dentary ONM CBI-1-646, which bears four, the exact number of labial foramina cannot be assessed in the remaining dentaries. Interestingly, however, ONM CBI-1-659 is pierced by several (at least nine) tiny foramina in its labial surface, most of which are closely spaced.
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFCD7F85108B6F94A" blockId="18.[824,1477,144,1703]" pageId="18" pageNumber="19">
As in the case of the maxillae above, in order to facilitate further quantitative investigation, we introduce the ratio of largest dentary tooth length to the total tooth row length of the maxilla. This was complete in only 
<specimenCount id="9D77A7DEFFDCB44EFBC7F8F308B2F94A" box="[1093,1249,1679,1703]" count="2" pageId="18" pageNumber="19" type="generic">two specimens</specimenCount>
:
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFCCAF8AD08BAF8E8" blockId="18.[840,1474,1742,1860]" pageId="18" pageNumber="19">
<typeStatus id="54CAD2F5FFDCB44EFCCAF8AD0FF1F90B" box="[840,930,1742,1766]" pageId="18" pageNumber="19" type="paratype">Paratype</typeStatus>
ONM CBI-1-646: largest tooth length, 
<quantity id="4C89C1B2FFDCB44EFABBF8AD09DAF90B" box="[1337,1417,1742,1766]" metricMagnitude="-3" metricUnit="m" metricValue="2.8" pageId="18" pageNumber="19" unit="mm" value="2.8">2.8 mm</quantity>
/total tooth row length, 
<quantity id="4C89C1B2FFDCB44EFB94F88E0820F8E8" box="[1046,1139,1773,1797]" metricMagnitude="-2" metricUnit="m" metricValue="1.19" pageId="18" pageNumber="19" unit="mm" value="11.9">11.9 mm</quantity>
; ratio, 0.24.
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFCCAF96E08CCF8A9" blockId="18.[840,1474,1742,1860]" pageId="18" pageNumber="19">
ONM CBI-1-657: largest tooth length, 
<quantity id="4C89C1B2FFDCB44EFB74F96E0915F8C8" box="[1270,1350,1805,1829]" metricMagnitude="-3" metricUnit="m" metricValue="1.6" pageId="18" pageNumber="19" unit="mm" value="1.6">1.6 mm</quantity>
/total tooth row length, 
<quantity id="4C89C1B2FFDCB44EFC5AF94F087AF8AE" box="[984,1065,1836,1860]" metricMagnitude="-3" metricUnit="m" metricValue="6.4" pageId="18" pageNumber="19" unit="mm" value="6.4">6.4 mm</quantity>
; ratio, 0.25.
</paragraph>
<paragraph id="8BCE6C57FFDCB44EFCD7F9080802F82C" blockId="18.[826,1474,1899,1985]" pageId="18" pageNumber="19">Besides these specimens for which ratios could be calculated, the following dentaries provide data on the length of the largest tooth in further specimens:</paragraph>
<caption id="DF0E3CDFFFDDB44FFFF3FB6B09E1FACD" ID-DOI="http://doi.org/10.5281/zenodo.14514581" ID-Zenodo-Dep="14514581" box="[113,1458,1287,1312]" httpUri="https://zenodo.org/record/14514581/files/figure.png" pageId="19" pageNumber="20" startId="19.[113,178,1288,1312]" targetBox="[116,1456,146,1258]" targetPageId="19" targetType="figure">
<paragraph id="8BCE6C57FFDDB44FFFF3FB6B09E1FACD" blockId="19.[113,1458,1287,1312]" box="[113,1458,1287,1312]" pageId="19" pageNumber="20">
<emphasis id="B905B045FFDDB44FFFF3FB6B0C86FACD" bold="true" box="[113,213,1288,1312]" pageId="19" pageNumber="20">Figure 13.</emphasis>
Dentary of 
<taxonomicName id="4C7117D4FFDDB44FFEC7FB640EE2FAF2" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[325,689,1287,1311]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="19" pageNumber="20" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFDDB44FFEC7FB640EE2FAF2" bold="true" box="[325,689,1287,1311]" italics="true" pageId="19" pageNumber="20">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. Photographs of left dentary ONM CBI-1-647 in labial (A) and medial (B) views.
</paragraph>
</caption>
<paragraph id="8BCE6C57FFDDB44FFF02FB330E37FA8A" blockId="19.[128,612,1360,1478]" box="[128,612,1360,1384]" pageId="19" pageNumber="20">
ONM CBI-1-647: largest tooth length, 
<quantity id="4C89C1B2FFDDB44FFD92FB330E32FA8A" box="[528,609,1360,1384]" metricMagnitude="-3" metricUnit="m" metricValue="2.9" pageId="19" pageNumber="20" unit="mm" value="2.9">2.9 mm</quantity>
.
</paragraph>
<paragraph id="8BCE6C57FFDDB44FFF02FB0C0E37FA6A" blockId="19.[128,612,1360,1478]" box="[128,612,1391,1415]" pageId="19" pageNumber="20">
ONM CBI-1-659: largest tooth length, 
<quantity id="4C89C1B2FFDDB44FFD92FB0C0E32FA6A" box="[528,609,1391,1415]" metricMagnitude="-3" metricUnit="m" metricValue="2.3" pageId="19" pageNumber="20" unit="mm" value="2.3">2.3 mm</quantity>
.
</paragraph>
<paragraph id="8BCE6C57FFDDB44FFF02FBEC0E37FA4B" blockId="19.[128,612,1360,1478]" box="[128,612,1422,1447]" pageId="19" pageNumber="20">
ONM CBI-1-655: largest tooth length, 
<quantity id="4C89C1B2FFDDB44FFD92FBEC0E32FA4B" box="[528,609,1423,1447]" metricMagnitude="-3" metricUnit="m" metricValue="1.9" pageId="19" pageNumber="20" unit="mm" value="1.9">1.9 mm</quantity>
.
</paragraph>
<paragraph id="8BCE6C57FFDDB44FFF02FBCD0E37FA28" blockId="19.[128,612,1360,1478]" box="[128,612,1454,1478]" pageId="19" pageNumber="20">
ONM CBI-1-656: largest tooth length, 
<quantity id="4C89C1B2FFDDB44FFD92FBCD0E32FA28" box="[528,609,1454,1478]" metricMagnitude="-3" metricUnit="m" metricValue="1.5" pageId="19" pageNumber="20" unit="mm" value="1.5">1.5 mm</quantity>
.
</paragraph>
<paragraph id="8BCE6C57FFDDB44FFEDCFB860E5DF9ED" blockId="19.[113,763,1509,1980]" box="[350,526,1509,1536]" pageId="19" pageNumber="20">
<emphasis id="B905B045FFDDB44FFEDCFB860E5DF9ED" box="[350,526,1509,1536]" italics="true" pageId="19" pageNumber="20">
Vertebrae (
<figureCitation id="134A70D2FFDDB44FFE46FB850E50FA12" box="[452,515,1509,1535]" captionStart="Figure 17" captionStartId="23.[113,178,1844,1868]" captionTargetBox="[166,1406,151,1814]" captionTargetId="figure-6@23.[164,1408,149,1816]" captionTargetPageId="23" captionText="Figure 17. Vertebrae tentatively attributed to Terastiodontosaurus marcelosanchezi. A–E, presacral vertebra ONM CBI-1-833 in anterior (A), posterior (B), right lateral (C), dorsal (D), and ventral (E) views; F–J, presacral vertebra ONM CBI-1-860 in anterior (F), posterior (G), left lateral (H), dorsal (I), and ventral (J) views; K–O, presacral vertebra ONM CBI-1-820 in anterior (K), posterior (L), left lateral (M), dorsal (N), and ventral (O) views; P–R, presacral vertebra ONM CBI-1-682 in anterior (P), dorsal (Q), and ventral (R) views; S–U, presacral vertebra ONM CBI-1-687 in dorsal (S), ventral (T), and right lateral (U) views; V, W, anterior caudal vertebra ONM CBI-1-689 in left lateral (V) and ventral (W) views; X, Y, posterior caudal vertebra ONM CBI-1-686 in ventral (X) and left lateral (Y) views." figureDoi="http://doi.org/10.5281/zenodo.14514589" httpUri="https://zenodo.org/record/14514589/files/figure.png" pageId="19" pageNumber="20">Fig.17</figureCitation>
)
</emphasis>
</paragraph>
<paragraph id="8BCE6C57FFDDB44FFFF3F86F0C91F96F" blockId="19.[113,763,1509,1980]" pageId="19" pageNumber="20">Vertebrae are referred tentatively to the same taxon on the basis of co-occurrence and the fact that large numbers of jaws have yielded only a single species of amphisbaenian thus far from the locality.</paragraph>
<paragraph id="8BCE6C57FFDDB44FFF0FF8EA086BF912" blockId="19.[113,763,1509,1980]" lastBlockId="19.[810,1460,1360,1979]" pageId="19" pageNumber="20">
Presacral vertebrae range in size between ~1 and 
<quantity id="4C89C1B2FFDDB44FFD3CF8EA0EA9F94C" box="[702,762,1673,1697]" metricMagnitude="-3" metricUnit="m" metricValue="5.0" pageId="19" pageNumber="20" unit="mm" value="5.0">5 mm</quantity>
(
<figureCitation id="134A70D2FFDDB44FFFFEF8CB0CAAF92C" box="[124,249,1704,1729]" captionStart="Figure 17" captionStartId="23.[113,178,1844,1868]" captionTargetBox="[166,1406,151,1814]" captionTargetId="figure-6@23.[164,1408,149,1816]" captionTargetPageId="23" captionText="Figure 17. Vertebrae tentatively attributed to Terastiodontosaurus marcelosanchezi. A–E, presacral vertebra ONM CBI-1-833 in anterior (A), posterior (B), right lateral (C), dorsal (D), and ventral (E) views; F–J, presacral vertebra ONM CBI-1-860 in anterior (F), posterior (G), left lateral (H), dorsal (I), and ventral (J) views; K–O, presacral vertebra ONM CBI-1-820 in anterior (K), posterior (L), left lateral (M), dorsal (N), and ventral (O) views; P–R, presacral vertebra ONM CBI-1-682 in anterior (P), dorsal (Q), and ventral (R) views; S–U, presacral vertebra ONM CBI-1-687 in dorsal (S), ventral (T), and right lateral (U) views; V, W, anterior caudal vertebra ONM CBI-1-689 in left lateral (V) and ventral (W) views; X, Y, posterior caudal vertebra ONM CBI-1-686 in ventral (X) and left lateral (Y) views." figureDoi="http://doi.org/10.5281/zenodo.14514589" httpUri="https://zenodo.org/record/14514589/files/figure.png" pageId="19" pageNumber="20">Fig. 17A–U</figureCitation>
). They are procoelous and dorsoventrally compressed. In anterior view, the prezygapophyses are strongly inclined, much exceeding in height the anterodorsal edge of the neural canal. There is no zygosphene. The cotyle is elliptical and strongly depressed. In posterior view, the condyle is also elliptical and strongly depressed. The neural arch is depressed. The lateral walls of the neural arch form moderately robust centropostzygapophyseal laminae (
<emphasis id="B905B045FFDDB44FFD47F9E60EA9F871" box="[709,762,1925,1948]" italics="true" pageId="19" pageNumber="20">sensu</emphasis>
<bibRefCitation id="EFE011A6FFDDB44FFFF3F9C00D3BF856" author="Georgalis GL" box="[113,360,1955,1979]" pageId="19" pageNumber="20" pagination="155 - 9" refId="ref24868" refString="Georgalis GL * Villa A * Delfino M. The last amphisbaenian (ºquamata) from continental Eastern Europe. Annales de Paleontologie 2018 b; 104: 155 - 9. https: // doi. org / 10.1016 / j. annpal. 2018.03.002" type="journal article" year="2018">
Georgalis 
<emphasis id="B905B045FFDDB44FFF61F9C70D49F856" box="[227,282,1955,1979]" italics="true" pageId="19" pageNumber="20">et al.</emphasis>
2018b
</bibRefCitation>
). There is no zygantrum. In dorsal view, the prezygapophyses extend anterolaterally. The prezygapophyseal articular facets are large and broad; in some specimens, there are prominent prezygapophyseal accessory processes. There is no neural spine. The interzygapophyseal constriction is deep. There is practically no posterior median notch of the neural arch. In ventral view, the centrum is flattened, with only slightly concave lateral margins. Two usually large, occasionally asymmetrical subcentral foramina are present, one at each lateral side of the ventral surface of the centrum. The synapophyses are robust and more or less rounded. In lateral view, the neural arch rises distinctly, with a gentle curve towards its posterior end. Each prezygapophysis is connected to the related postzygapophysis by a relatively low interzygapophyseal ridge.
</paragraph>
<paragraph id="8BCE6C57FFDDB448FCC7F9650C9EF91B" blockId="19.[810,1460,1360,1979]" lastBlockId="20.[129,778,1726,1782]" lastPageId="20" lastPageNumber="21" pageId="19" pageNumber="20">
Caudal vertebrae have haemapophyses fused to the centrum. A short anterior caudal vertebra has forked lymphapophyses (
<figureCitation id="134A70D2FFDDB44FFCB7F9260FDFF8B0" box="[821,908,1861,1885]" captionStart="Figure 17" captionStartId="23.[113,178,1844,1868]" captionTargetBox="[166,1406,151,1814]" captionTargetId="figure-6@23.[164,1408,149,1816]" captionTargetPageId="23" captionText="Figure 17. Vertebrae tentatively attributed to Terastiodontosaurus marcelosanchezi. A–E, presacral vertebra ONM CBI-1-833 in anterior (A), posterior (B), right lateral (C), dorsal (D), and ventral (E) views; F–J, presacral vertebra ONM CBI-1-860 in anterior (F), posterior (G), left lateral (H), dorsal (I), and ventral (J) views; K–O, presacral vertebra ONM CBI-1-820 in anterior (K), posterior (L), left lateral (M), dorsal (N), and ventral (O) views; P–R, presacral vertebra ONM CBI-1-682 in anterior (P), dorsal (Q), and ventral (R) views; S–U, presacral vertebra ONM CBI-1-687 in dorsal (S), ventral (T), and right lateral (U) views; V, W, anterior caudal vertebra ONM CBI-1-689 in left lateral (V) and ventral (W) views; X, Y, posterior caudal vertebra ONM CBI-1-686 in ventral (X) and left lateral (Y) views." figureDoi="http://doi.org/10.5281/zenodo.14514589" httpUri="https://zenodo.org/record/14514589/files/figure.png" pageId="19" pageNumber="20">Fig. 17V</figureCitation>
, W), whereas more elongate posterior caudal vertebrae have unitary pleurapophyses (
<figureCitation id="134A70D2FFDDB44FFB14F90708BEF890" box="[1174,1261,1892,1917]" captionStart="Figure 17" captionStartId="23.[113,178,1844,1868]" captionTargetBox="[166,1406,151,1814]" captionTargetId="figure-6@23.[164,1408,149,1816]" captionTargetPageId="23" captionText="Figure 17. Vertebrae tentatively attributed to Terastiodontosaurus marcelosanchezi. A–E, presacral vertebra ONM CBI-1-833 in anterior (A), posterior (B), right lateral (C), dorsal (D), and ventral (E) views; F–J, presacral vertebra ONM CBI-1-860 in anterior (F), posterior (G), left lateral (H), dorsal (I), and ventral (J) views; K–O, presacral vertebra ONM CBI-1-820 in anterior (K), posterior (L), left lateral (M), dorsal (N), and ventral (O) views; P–R, presacral vertebra ONM CBI-1-682 in anterior (P), dorsal (Q), and ventral (R) views; S–U, presacral vertebra ONM CBI-1-687 in dorsal (S), ventral (T), and right lateral (U) views; V, W, anterior caudal vertebra ONM CBI-1-689 in left lateral (V) and ventral (W) views; X, Y, posterior caudal vertebra ONM CBI-1-686 in ventral (X) and left lateral (Y) views." figureDoi="http://doi.org/10.5281/zenodo.14514589" httpUri="https://zenodo.org/record/14514589/files/figure.png" pageId="19" pageNumber="20">Fig. 17X</figureCitation>
, Y). If the number of caudal to presacral vertebrae can be determined, the methodology of Smith (2013) might be used to estimate the proportion of caudal vertebrae and thus to constrain the relative length of the tail.
</paragraph>
<caption id="DF0E3CDFFFDAB448FF03F8390FB1F962" ID-DOI="http://doi.org/10.5281/zenodo.14514583" ID-Zenodo-Dep="14514583" httpUri="https://zenodo.org/record/14514583/files/figure.png" pageId="20" pageNumber="21" startId="20.[129,194,1626,1650]" targetBox="[136,1459,156,1585]" targetPageId="20" targetType="figure">
<paragraph id="8BCE6C57FFDAB448FF03F8390FB1F962" blockId="20.[129,1458,1626,1679]" pageId="20" pageNumber="21">
<emphasis id="B905B045FFDAB448FF03F8390CB6F99F" bold="true" box="[129,229,1626,1650]" pageId="20" pageNumber="21">Figure 14.</emphasis>
Dentaries of 
<taxonomicName id="4C7117D4FFDAB448FEE0F8390E9DF99F" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[354,718,1626,1650]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFDAB448FEE0F8390E9DF99F" bold="true" box="[354,718,1626,1650]" italics="true" pageId="20" pageNumber="21">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
. Photographs of left dentary ONM CBI-1-657 in medial view (A); left dentary ONM CBI-1-655 in labial view (B); and right dentary ONM CBI-1-656 in medial view (C).
</paragraph>
</caption>
<paragraph id="8BCE6C57FFDAB448FE88F9760E2CF8C2" blockId="20.[266,639,1813,1839]" box="[266,639,1813,1839]" pageId="20" pageNumber="21">
<emphasis id="B905B045FFDAB448FE88F9760E2CF8C2" bold="true" box="[266,639,1813,1839]" pageId="20" pageNumber="21">Results of the phylogenetic analysis</emphasis>
</paragraph>
<paragraph id="8BCE6C57FFDAB446FF03F95F0D38FB9E" blockId="20.[129,778,1852,1970]" lastBlockId="26.[128,779,144,1139]" lastPageId="26" lastPageNumber="27" pageId="20" pageNumber="21">
Our main analysis used only two topological constraints: Afrobaenia and South American 
<taxonomicName id="4C7117D4FFDAB448FE60F9380EC3F89E" box="[482,656,1883,1907]" class="Squamata" family="Amphisbaenidae" kingdom="Animalia" pageId="20" pageNumber="21" phylum="Chordata" rank="family">Amphisbaenidae</taxonomicName>
. Fully consistent with all recent phylogenetic analyses (e.g. 
<bibRefCitation id="EFE011A6FFDAB448FD36F9180CE6F85F" author="Kearney M" pageId="20" pageNumber="21" pagination="1 - 74" refId="ref26695" refString="Kearney M. ºystematics of the Amphisbaenia (Lepidosauria: ºquamata) based on morphological evidence from recent and fossil forms. Herpetological Monographs 2003; 17: 1 - 74. https: // doi. org / 10.1655 / 0733 - 1347 (2003) 017 [0001: sotalb] 2.0. co; 2" type="journal article" year="2003">Kearney 2003</bibRefCitation>
, 
<bibRefCitation id="EFE011A6FFDAB448FF43F9F90D24F85F" author="Muller J" box="[193,375,1946,1970]" pageId="20" pageNumber="21" pagination="364 - 7" refId="ref28298" refString="Muller J * Hipsley CA * Head JJ et al. Eocene lizard from Germany reveals amphisbaenian origins. Nature 2011; 473: 364 - 7. https: // doi. org / 10.1038 / nature 09919" type="journal article" year="2011">
Müller 
<emphasis id="B905B045FFDAB448FE8FF9F80D6EF85F" box="[269,317,1946,1970]" italics="true" pageId="20" pageNumber="21">et al.</emphasis>
2011
</bibRefCitation>
, 
<bibRefCitation id="EFE011A6FFDAB448FE01F9F90E1CF85F" author="Gauthier JA" box="[387,591,1946,1970]" pageId="20" pageNumber="21" pagination="3 - 308" refId="ref24548" refString="Gauthier JA * Kearney M * Maisano JA et al. Assembling the squamate tree of life: perspectives from the phenotype and the fossil record. Bulletin of the Peabody Museum of Natural History 2012; 53: 3 - 308. https: // doi. org / 10.3374 / 014.053.0101" type="journal article" year="2012">
Gauthier 
<emphasis id="B905B045FFDAB448FE67F9F80E46F85F" box="[485,533,1946,1970]" italics="true" pageId="20" pageNumber="21">et al.</emphasis>
2012
</bibRefCitation>
, 
<bibRefCitation id="EFE011A6FFDAB448FDD9F9F90F57F85F" author="Jones MEH" box="[603,772,1946,1970]" pageId="20" pageNumber="21" refId="ref26565" refString="Jones MEH * Anderson CJ * Hipsley CA et al. Integration of molecules and new fossils supports a Triassic origin for Lepidosauria (lizards * snakes * and tuatara). BMC Evolutionary Biology 2013; 13: 208. https: // doi. org / 10.1186 / 1471 - 2148 - 13 - 208" type="journal volume" year="2013">
Jones 
<emphasis id="B905B045FFDAB448FD18F9F80E99F85F" box="[666,714,1946,1970]" italics="true" pageId="20" pageNumber="21">et al.</emphasis>
2013
</bibRefCitation>
, 
<bibRefCitation id="EFE011A6FFDAB448FCBBF8A30870F93B" author="Cernansky A" box="[825,1059,1726,1755]" pageId="20" pageNumber="21" refId="ref22773" refString="Cernansky A * Auge M * Rage J-C. A complete mandible of a new amphisbaenian reptile (ºquamata * Amphisbaenia) from the late Middle Eocene (Bartonian * MP 16) of France. Journal of Vertebrate Paleontology 2015 a; 35: e 902379. https: // doi. org / 10.1080 / 0272463 4.2014. 902379" type="journal volume" year="2015">
Čerňanský 
<emphasis id="B905B045FFDAB448FC2CF8DC0F8DF93B" box="[942,990,1726,1750]" italics="true" pageId="20" pageNumber="21">et al.</emphasis>
2015a
</bibRefCitation>
, 
<bibRefCitation id="EFE011A6FFDAB448FBADF8DD08ACF93B" author="Longrich NR" box="[1071,1279,1726,1750]" pageId="20" pageNumber="21" refId="ref27285" refString="Longrich NR * Vinther J * Pyron RA et al. Biogeography of worm lizards (Amphisbaenia) driven by end-Cretaceous mass extinction. Proceedings Biological Sciences 2015; 282: 20143034. https: // doi. org / 10.1098 / rspb. 2014.3034" type="journal volume" year="2015">
Longrich 
<emphasis id="B905B045FFDAB448FB17F8DC0896F93B" box="[1173,1221,1726,1750]" italics="true" pageId="20" pageNumber="21">et al.</emphasis>
2015
</bibRefCitation>
, Zheng and Wiens 2016, Streicher and 
<bibRefCitation id="EFE011A6FFDAB448FB84F8BD08D2F91B" author="Wiens JJ" box="[1030,1153,1758,1782]" pageId="20" pageNumber="21" refId="ref30775" refString="ºtreicher JW * Wiens JJ. Phylogenomic analyses of more than 4000 nuclear loci resolve the origin of snakes among lizard families. Biology Letters 2017; 13: 20170393. https: // doi. org / 10.1098 / rsbl. 2017.0393" type="journal volume" year="2017">Wiens 2017</bibRefCitation>
, Simões 
<emphasis id="B905B045FFDAB448FB5EF8BC0958F91B" box="[1244,1291,1758,1782]" italics="true" pageId="20" pageNumber="21">et al.</emphasis>
2018, 
<bibRefCitation id="EFE011A6FFDAB448FACDF8BD0FDAF8F8" author="Burbrink FT" pageId="20" pageNumber="21" pagination="502 - 20" refId="ref22612" refString="Burbrink FT * Grazziotin GF * Pyron RA et al. Interrogating genomic-scale data for ºquamata (lizards * snakes * and amphisbaenians) shows no support for key traditional morphological relationships. Systematic Biology 2020; 69: 502 - 20. https: // doi. org / 10.1093 / grurint / ikaa 038" type="journal article" year="2020">
Burbrink 
<emphasis id="B905B045FFDAB448FA33F8BC0F01F8F8" italics="true" pageId="20" pageNumber="21">et al.</emphasis>
2020
</bibRefCitation>
, Singhal 
<emphasis id="B905B045FFDAB448FC66F89D0841F8F8" box="[996,1042,1789,1813]" italics="true" pageId="20" pageNumber="21">et al.</emphasis>
2021, 
<bibRefCitation id="EFE011A6FFDAB448FBD6F89E0943F8F8" author="Talanda M" box="[1108,1296,1789,1813]" pageId="20" pageNumber="21" pagination="99 - 104" refId="ref31227" refString="Talanda M * Fernandez V * Panciroli E et al. ºynchrotron tomography of a stem lizard elucidates early squamate anatomy. Nature 2022; 611: 99 - 104. https: // doi. org / 10.1038 / s 41586 - 022 - 05332 - 6" type="journal article" year="2022">
Tałanda 
<emphasis id="B905B045FFDAB448FB29F89D088AF8F8" box="[1195,1241,1789,1813]" italics="true" pageId="20" pageNumber="21">et al.</emphasis>
2022
</bibRefCitation>
, 
<bibRefCitation id="EFE011A6FFDAB448FA99F89E0F3EF8D8" author="Brownstein CD" pageId="20" pageNumber="21" refId="ref22565" refString="Brownstein CD * ºimoes TR * Caldwell MW et al. The affinities of the Late Triassic Cryptovaranoides and the age of crown squamates. Royal Society Open Science 2023; 10: 230968. https: // doi. org / 10.1098 / rsos. 230968" type="journal volume" year="2023">
Brownstein 
<emphasis id="B905B045FFDAB448FA17F89D0990F8F8" box="[1429,1475,1789,1813]" italics="true" pageId="20" pageNumber="21">et al.</emphasis>
2023
</bibRefCitation>
, Čerňanský and Vasilyan 2024), our analysis finds strong support for amphisbaenian monophyly (
<figureCitation id="134A70D2FFDAB448FB75F95F0910F8B9" box="[1271,1347,1852,1876]" captionStart="Figure 18" captionStartId="24.[130,195,1834,1858]" captionTargetBox="[248,1354,142,1805]" captionTargetId="graphics-70@24.[248,1021,150,1795]" captionTargetPageId="24" captionText="Figure 18.Phylogenetic analysis of Amphisbaenia, strict consensus of maximum parsimony trees, indicating the position of Terastiodontosaurus marcelosanchezi." figureDoi="http://doi.org/10.5281/zenodo.14514591" httpUri="https://zenodo.org/record/14514591/files/figure.png" pageId="20" pageNumber="21">Fig. 18</figureCitation>
). The position of 
<taxonomicName id="4C7117D4FFDAB448FC0FF93808E1F89E" authority="Muller" authorityName="Muller" box="[909,1202,1883,1907]" class="Squamata" family="Lacertidae" genus="Cryptolacerta" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="20" pageNumber="21" phylum="Chordata" rank="species" species="hassiaca">
<emphasis id="B905B045FFDAB448FC0FF9380838F89E" box="[909,1131,1883,1907]" italics="true" pageId="20" pageNumber="21">Cryptolacerta hassiaca</emphasis>
Müller
</taxonomicName>
, Hipsley, Head, Kardjilov, Hilger, Wuttke &amp; Reisz, 2011, from the early to middle Eocene of Messel, 
<collectingCountry id="F3662CC7FFDAB448FC36F9F9085CF85F" box="[948,1039,1946,1970]" name="Germany" pageId="20" pageNumber="21">Germany</collectingCountry>
, originally described as a link between lacertids and amphisbaenians by 
<bibRefCitation id="EFE011A6FFD6B444FE66F91A0EEFF87C" author="Muller J" box="[484,700,1913,1937]" pageId="24" pageNumber="25" pagination="364 - 7" refId="ref28298" refString="Muller J * Hipsley CA * Head JJ et al. Eocene lizard from Germany reveals amphisbaenian origins. Nature 2011; 473: 364 - 7. https: // doi. org / 10.1038 / nature 09919" type="journal article" year="2011">
Müller 
<emphasis id="B905B045FFD6B444FDB6F91A0E3BF87C" box="[564,616,1913,1937]" italics="true" pageId="24" pageNumber="25">et al.</emphasis>
(2011)
</bibRefCitation>
, is unresolved (but see also 
<bibRefCitation id="EFE011A6FFD6B444FEF0F9FB0E19F85D" author="Longrich NR" box="[370,586,1944,1968]" pageId="24" pageNumber="25" refId="ref27285" refString="Longrich NR * Vinther J * Pyron RA et al. Biogeography of worm lizards (Amphisbaenia) driven by end-Cretaceous mass extinction. Proceedings Biological Sciences 2015; 282: 20143034. https: // doi. org / 10.1098 / rspb. 2014.3034" type="journal volume" year="2015">
Longrich 
<emphasis id="B905B045FFD6B444FE58F9FA0E5EF85D" box="[474,525,1944,1968]" italics="true" pageId="24" pageNumber="25">et al.</emphasis>
2015
</bibRefCitation>
, 
<bibRefCitation id="EFE011A6FFD6B444FDDBF9FB0CE6F822" author="Brownstein CD" pageId="24" pageNumber="25" refId="ref22565" refString="Brownstein CD * ºimoes TR * Caldwell MW et al. The affinities of the Late Triassic Cryptovaranoides and the age of crown squamates. Royal Society Open Science 2023; 10: 230968. https: // doi. org / 10.1098 / rsos. 230968" type="journal volume" year="2023">
Brownstein 
<emphasis id="B905B045FFD6B444FD55F9FA0F59F85D" box="[727,778,1944,1968]" italics="true" pageId="24" pageNumber="25">et al.</emphasis>
2023
</bibRefCitation>
, Čerňanský and Vasilyan 2024). In general, higher-level relationships within Amphisbaenia are poorly supported, but our analysis shares the basal position of 
<taxonomicName id="4C7117D4FFD6B444FB29F9FB0979F85D" authorityName="Vanzolini" authorityYear="1951" box="[1195,1322,1944,1968]" class="Squamata" family="Rhineuridae" kingdom="Animalia" pageId="24" pageNumber="25" phylum="Chordata" rank="family">Rhineuridae</taxonomicName>
with 
<bibRefCitation id="EFE011A6FFD6B444FAEAF9FB0FE9F822" author="Gauthier JA" pageId="24" pageNumber="25" pagination="3 - 308" refId="ref24548" refString="Gauthier JA * Kearney M * Maisano JA et al. Assembling the squamate tree of life: perspectives from the phenotype and the fossil record. Bulletin of the Peabody Museum of Natural History 2012; 53: 3 - 308. https: // doi. org / 10.3374 / 014.053.0101" type="journal article" year="2012">
Gauthier 
<emphasis id="B905B045FFD6B444FCBBF9DB0F39F822" box="[825,874,1975,1999]" italics="true" pageId="24" pageNumber="25">et al.</emphasis>
(2012)
</bibRefCitation>
, which was the first purely morphological analysis to recover this topology. A surprise was the close relationship between the unnamed amphisbaenian from Adrar-Mgorn 1 of Augé and Rage (2006) to 
<taxonomicName id="4C7117D4FFD4B446FE08FEAC0D8CFF0A" baseAuthorityName="Kearney" baseAuthorityYear="2003" box="[394,479,207,231]" class="Squamata" family="Blanidae" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="family">Blanidae</taxonomicName>
, although with poor support (BS &lt;0.50); furthermore, if all higher-level relationships within Afrobaenia (e.g. 
<bibRefCitation id="EFE011A6FFD4B446FEA9FF6E0DABFEC8" author="Graboski R" box="[299,504,269,293]" pageId="26" pageNumber="27" refId="ref25266" refString="Graboski R * Grazziotin FG * Mott T et al. The phylogenetic position of Ridley's Worm Lizard reveals the complex biogeographic history of New World insular amphisbaenids. Molecular Phylogenetics and Evolution 2022; 173: 107518. https: // doi. org / 10.1016 / j. ympev. 2022.107518" type="journal volume" year="2022">
Graboski 
<emphasis id="B905B045FFD4B446FE0DFF6D0DECFEC8" box="[399,447,269,293]" italics="true" pageId="26" pageNumber="27">et al.</emphasis>
2022
</bibRefCitation>
) are enforced, its position becomes unresolved. This unnamed pleurodont form represents the second amphisbaenian from the locality of Adrar-Mgorn 
<quantity id="4C89C1B2FFD4B446FD7EFF2F0CC6FE6E" metricMagnitude="-2" metricUnit="m" metricValue="2.54" pageId="26" pageNumber="27" unit="in" value="1.0">1 in</quantity>
<collectingCountry id="F3662CC7FFD4B446FF1EFF0F0CAAFE6E" box="[156,249,364,387]" name="Morocco" pageId="26" pageNumber="27">Morocco</collectingCountry>
(the other being 
<taxonomicName id="4C7117D4FFD4B446FE36FF080EF0FE6E" box="[436,675,363,387]" genus="Todrasaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="gheerbranti">
<emphasis id="B905B045FFD4B446FE36FF080EF0FE6E" box="[436,675,363,387]" italics="true" pageId="26" pageNumber="27">Todrasaurus gheerbranti</emphasis>
</taxonomicName>
) and was originally described by Augé and Rage (2006) as bearing some resemblance to both blanids and amphisbaenids, whereas the phylogenetic analysis of 
<bibRefCitation id="EFE011A6FFD4B446FE03FFAA0E39FE0C" author="Longrich NR" box="[385,618,457,481]" pageId="26" pageNumber="27" refId="ref27285" refString="Longrich NR * Vinther J * Pyron RA et al. Biogeography of worm lizards (Amphisbaenia) driven by end-Cretaceous mass extinction. Proceedings Biological Sciences 2015; 282: 20143034. https: // doi. org / 10.1098 / rspb. 2014.3034" type="journal volume" year="2015">
Longrich 
<emphasis id="B905B045FFD4B446FE65FFA90E4AFE0C" box="[487,537,457,481]" italics="true" pageId="26" pageNumber="27">et al.</emphasis>
(2015)
</bibRefCitation>
recovered it as an Amphisbaenia 
<emphasis id="B905B045FFD4B446FEC2FF8A0D90FDEC" box="[320,451,489,513]" italics="true" pageId="26" pageNumber="27">incertae sedis</emphasis>
. The cadeid 
<taxonomicName id="4C7117D4FFD4B446FDD2FF8A0CB4FDCD" authority="Dickerson" authorityName="Dickerson" authorityYear="1916" class="Squamata" family="Cadeidae" genus="Cadea" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="palirostrata">
<emphasis id="B905B045FFD4B446FDD2FF8A0F5AFDEC" box="[592,777,489,513]" italics="true" pageId="26" pageNumber="27">Cadea palirostrata</emphasis>
Dickerson
</taxonomicName>
, 1916 was assessed with relatively strong support as the sister taxon of Afrobaenia, i.e. the group encompassing 
<taxonomicName id="4C7117D4FFD4B446FF03FC240C8EFDB2" box="[129,221,583,607]" class="Squamata" family="Cadeidae" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="family">Cadeidae</taxonomicName>
, 
<taxonomicName id="4C7117D4FFD4B446FF73FC240DD8FDB2" authorityName="Bonaparte" authorityYear="1838" box="[241,395,583,607]" class="Squamata" family="Trogonophidae" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="family">Trogonophidae</taxonomicName>
, and 
<taxonomicName id="4C7117D4FFD4B446FE53FC240E2EFDB2" box="[465,637,583,607]" class="Squamata" family="Amphisbaenidae" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="family">Amphisbaenidae</taxonomicName>
(BS = 0.80). 
<taxonomicName id="4C7117D4FFD4B446FF03FC050DBAFD93" authorityName="Georgalis &amp; Smith" authorityYear="2024" box="[129,489,614,638]" class="Squamata" family="Trogonophidae" genus="Terastiodontosaurus" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="marcelosanchezi">
<emphasis id="B905B045FFD4B446FF03FC050DBAFD93" box="[129,489,614,638]" italics="true" pageId="26" pageNumber="27">Terastiodontosaurus marcelosanchezi</emphasis>
</taxonomicName>
was inferred with moderate support (BS = 0.65, one unambiguous character state change, Bremer support 1; 
<tableCitation id="C6F359ECFFD4B446FEC0FCC60DDEFD50" box="[322,397,677,701]" captionStart="Table 1" captionStartId="25.[113,168,143,167]" captionTargetPageId="25" captionText="Table 1. Unambiguous morphological synapomorphies (under acctran or deltran) of trogonophid clades inferred based on our phylogenetic analysis." httpUri="http://table.plazi.org/id/DF0E3CDFFFD7B445FFF3FEEC0C92FF2E" pageId="26" pageNumber="27" tableUuid="DF0E3CDFFFD7B445FFF3FEEC0C92FF2E">Table 1</tableCitation>
) to be the sister taxon of 
<taxonomicName id="4C7117D4FFD4B446FD12FCC60CBCFD31" genus="Todrasaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="gheerbranti">
<emphasis id="B905B045FFD4B446FD12FCC60CBCFD31" italics="true" pageId="26" pageNumber="27">Todrasaurus gheerbranti</emphasis>
</taxonomicName>
, and the two together were inferred with strong support to be the sister taxon of 
<taxonomicName id="4C7117D4FFD4B446FE24FC800EDDFD16" authorityName="Kaup" authorityYear="1830" box="[422,654,739,763]" class="Squamata" family="Trogonophidae" genus="Trogonophis" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="wiegmanni">
<emphasis id="B905B045FFD4B446FE24FC800EDDFD16" box="[422,654,739,763]" italics="true" pageId="26" pageNumber="27">Trogonophis wiegmanni</emphasis>
</taxonomicName>
(BS = 0.95, five unambiguous character state changes, Bremer support 3), a novel result. The herein novel topology of 
<taxonomicName id="4C7117D4FFD4B446FDE0FD410E8FFCD7" authorityName="Augé &amp; Rage" authorityYear="2006" box="[610,732,802,826]" genus="Todrasaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="genus">
<emphasis id="B905B045FFD4B446FDE0FD410E8FFCD7" box="[610,732,802,826]" italics="true" pageId="26" pageNumber="27">Todrasaurus</emphasis>
</taxonomicName>
differs from that of 
<bibRefCitation id="EFE011A6FFD4B446FEB9FD220E7AFCB4" author="Longrich NR" box="[315,553,833,857]" pageId="26" pageNumber="27" refId="ref27285" refString="Longrich NR * Vinther J * Pyron RA et al. Biogeography of worm lizards (Amphisbaenia) driven by end-Cretaceous mass extinction. Proceedings Biological Sciences 2015; 282: 20143034. https: // doi. org / 10.1098 / rspb. 2014.3034" type="journal volume" year="2015">
Longrich 
<emphasis id="B905B045FFD4B446FE21FD210D85FCB4" box="[419,470,833,857]" italics="true" pageId="26" pageNumber="27">et al.</emphasis>
(2015)
</bibRefCitation>
, who had tentatively recovered this Moroccan taxon on the stem of Afrobaenia (
<bibRefCitation id="EFE011A6FFD4B446FF09FDE30D0EFC75" author="Longrich NR" box="[139,349,896,920]" pageId="26" pageNumber="27" refId="ref27285" refString="Longrich NR * Vinther J * Pyron RA et al. Biogeography of worm lizards (Amphisbaenia) driven by end-Cretaceous mass extinction. Proceedings Biological Sciences 2015; 282: 20143034. https: // doi. org / 10.1098 / rspb. 2014.3034" type="journal volume" year="2015">
Longrich 
<emphasis id="B905B045FFD4B446FF73FDE20D71FC75" box="[241,290,896,920]" italics="true" pageId="26" pageNumber="27">et al.</emphasis>
2015
</bibRefCitation>
). Moreover, 
<taxonomicName id="4C7117D4FFD4B446FE64FDE30ED7FC75" authorityName="Bonaparte" authorityYear="1838" box="[486,644,896,920]" class="Squamata" family="Trogonophidae" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="family">Trogonophidae</taxonomicName>
(comprising 
<taxonomicName id="4C7117D4FFD4B446FF03FDFC0CAAFC5A" authorityName="Kaup" authorityYear="1830" box="[129,249,927,951]" class="Squamata" family="Trogonophidae" genus="Trogonophis" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="genus">
<emphasis id="B905B045FFD4B446FF03FDFC0CAAFC5A" box="[129,249,927,951]" italics="true" pageId="26" pageNumber="27">Trogonophis</emphasis>
</taxonomicName>
and its stem plus 
<taxonomicName id="4C7117D4FFD4B446FE31FDFC0F53FC5A" authority="Nikolskyi" authorityName="Nikolskyi" box="[435,768,927,951]" class="Squamata" family="Trogonophidae" genus="Diplometopon" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="species" species="zarudnyi">
<emphasis id="B905B045FFD4B446FE31FDFC0ECFFC5A" box="[435,668,927,951]" italics="true" pageId="26" pageNumber="27">Diplometopon zarudnyi</emphasis>
Nikolskyi
</taxonomicName>
, 1907) was inferred to be monophyletic with strong support (BS = 0.96, 26 unambiguous character state changes, Bremer support 5). Enforcing all major topological constraints within Afrobaenia (
<emphasis id="B905B045FFD4B446FE83FA7E0D77FBD8" box="[257,292,1053,1077]" italics="true" pageId="26" pageNumber="27">fide</emphasis>
<bibRefCitation id="EFE011A6FFD4B446FEABFA7E0DA1FBD8" author="Graboski R" box="[297,498,1053,1077]" pageId="26" pageNumber="27" refId="ref25266" refString="Graboski R * Grazziotin FG * Mott T et al. The phylogenetic position of Ridley's Worm Lizard reveals the complex biogeographic history of New World insular amphisbaenids. Molecular Phylogenetics and Evolution 2022; 173: 107518. https: // doi. org / 10.1016 / j. ympev. 2022.107518" type="journal volume" year="2022">
Graboski 
<emphasis id="B905B045FFD4B446FE0EFA7E0DE9FBD8" box="[396,442,1053,1077]" italics="true" pageId="26" pageNumber="27">et al.</emphasis>
2022
</bibRefCitation>
) did not affect the relationships within 
<taxonomicName id="4C7117D4FFD4B446FE83FA5F0DCCFBB9" authorityName="Bonaparte" authorityYear="1838" box="[257,415,1084,1108]" class="Squamata" family="Trogonophidae" kingdom="Animalia" pageId="26" pageNumber="27" phylum="Chordata" rank="family">Trogonophidae</taxonomicName>
(including fossil taxa) or their basal position in Afrobaenia.
</paragraph>
</subSubSection>
</treatment>
</document>