587 lines
103 KiB
XML
587 lines
103 KiB
XML
<document ID-DOI="10.1038/s42003-022-04119-9" ID-GBIF-Dataset="b49b6ede-730b-460a-a2c1-f1b7f221399d" ID-Zenodo-Dep="7390392" approvedBy="castro_plazi" checkinTime="1669987396771" checkinUser="jonas" docAuthor="Lee, Sungjin, Lee, Yuong-Nam, Currie, Philip J., Sissons, Robin, Park, Jin-Young, Kim, Su-Hwan, Barsbold, Rinchen & Tsogtbaatar, Khishigjav" docDate="2022" docId="972687861A5EFFD65269FC21FBA58A60" docLanguage="en" docName="CommunBiol.5.1.1185.1-9.pdf" docOrigin="Communications Biology 5 (1)" docTitle="Natovenator polydontus Lee & Lee & Currie & Sissons & Park & Kim & Barsbold & Tsogtbaatar 2022, gen. et sp. nov." docType="treatment" docVersion="3" lastPageNumber="6" masterDocId="6B1FFFFE1A5FFFD3512AFFA7FF8B8B02" masterDocTitle="A non-avian dinosaur with a streamlined body exhibits potential adaptations for swimming" masterLastPageNumber="9" masterPageNumber="1" pageNumber="2" updateTime="1669988030008" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>A non-avian dinosaur with a streamlined body exhibits potential adaptations for swimming</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Lee, Sungjin</mods:namePart>
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<mods:affiliation>School of Earth and Environmental Sciences, Seoul National University, Seoul, Korea.</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Lee, Yuong-Nam</mods:namePart>
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<mods:affiliation>School of Earth and Environmental Sciences, Seoul National University, Seoul, Korea.</mods:affiliation>
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<mods:nameIdentifier type="email">ynlee@snu.ac.kr</mods:nameIdentifier>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Currie, Philip J.</mods:namePart>
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<mods:affiliation>Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada.</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Sissons, Robin</mods:namePart>
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<mods:affiliation>Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada.</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Park, Jin-Young</mods:namePart>
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<mods:affiliation>School of Earth and Environmental Sciences, Seoul National University, Seoul, Korea.</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Kim, Su-Hwan</mods:namePart>
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<mods:affiliation>School of Earth and Environmental Sciences, Seoul National University, Seoul, Korea.</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Barsbold, Rinchen</mods:namePart>
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<mods:affiliation>Institute of Paleontology, Mongolian Academy of Sciences, Ulaanbaatar, Mongolia.</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Tsogtbaatar, Khishigjav</mods:namePart>
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<mods:affiliation>Institute of Paleontology, Mongolian Academy of Sciences, Ulaanbaatar, Mongolia.</mods:affiliation>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>Communications Biology</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2022</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2022-12-01</mods:number>
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</mods:detail>
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<mods:detail type="volume">
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<mods:number>5</mods:number>
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</mods:detail>
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<mods:detail type="issue">
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<mods:number>1</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>1</mods:start>
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<mods:end>9</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1038/s42003-022-04119-9</mods:identifier>
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<mods:identifier type="GBIF-Dataset">b49b6ede-730b-460a-a2c1-f1b7f221399d</mods:identifier>
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<mods:identifier type="Zenodo-Dep">7390392</mods:identifier>
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</mods:mods>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.7390400" ID-Zenodo-Dep="7390400" LSID="urn:lsid:plazi:treatment:972687861A5EFFD65269FC21FBA58A60" httpUri="http://treatment.plazi.org/id/972687861A5EFFD65269FC21FBA58A60" lastPageId="5" lastPageNumber="6" pageId="1" pageNumber="2">
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<subSubSection box="[835,1241,901,923]" pageId="1" pageNumber="2" type="nomenclature">
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<paragraph blockId="1.[810,1472,482,923]" box="[835,1241,901,923]" pageId="1" pageNumber="2">
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<taxonomicName authority="Lee & Lee & Currie & Sissons & Park & Kim & Barsbold & Tsogtbaatar, 2022" authorityName="Lee & Lee & Currie & Sissons & Park & Kim & Barsbold & Tsogtbaatar" authorityYear="2022" box="[835,1076,901,923]" class="Reptilia" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="polydontus" status="gen. et sp. nov.">
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<emphasis box="[835,1076,901,923]" italics="true" pageId="1" pageNumber="2">Natovenator polydontus</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[1085,1241,901,922]" pageId="1" pageNumber="2" rank="species">gen. et sp. nov.</taxonomicNameLabel>
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2" type="materials_examined">
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<paragraph blockId="1.[810,1471,976,1026]" lastBlockId="2.[116,778,1372,1450]" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">
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<materialsCitation country="Mongolia" lastPageId="2" lastPageNumber="3" location="Hermiin Tsav" pageId="1" pageNumber="2" specimenCode="MPC-D 102/114" specimenCount="1" stateProvince="Omnogovi Province" typeStatus="holotype">
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<emphasis bold="true" box="[810,907,976,997]" pageId="1" pageNumber="2">
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<typeStatus box="[810,907,976,997]" pageId="1" pageNumber="2">Holotype</typeStatus>
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</emphasis>
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.
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<specimenCode box="[919,1086,976,997]" pageId="1" pageNumber="2">MPC-D 102/114</specimenCode>
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(Institute of Paleontology, Mongolian Academy of Sciences, Ulaanbaatar, Mongolia) is a mostly articulated skeleton with a nearly complete skull (See Supplementary
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<tableCitation box="[209,285,1307,1329]" captionStart="Table 1" captionStartId="5.[127,184,1521,1543]" captionTargetBox="[127,1393,1583,1858]" captionTargetId="graphics-966@5.[116,1471,1499,1966]" captionTargetPageId="5" captionText="Table 1 Dorsal rib angles of Natovenator polydontus, various diving birds, and terrestrial taxa measured in ventral view." httpUri="http://table.plazi.org/id/4BF066181A5AFFD65155FA56FAEA8D05" pageId="2" pageNumber="3" tableUuid="4BF066181A5AFFD65155FA56FAEA8D05">Table 1</tableCitation>
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for measurements).
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<emphasis bold="true" box="[116,343,1372,1394]" pageId="2" pageNumber="3">Locality and horizon</emphasis>
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.
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<geologicalTimeScale pageId="2" pageNumber="3">Baruungoyot Formation (Upper Cretaceous)</geologicalTimeScale>
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,
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<location LSID="urn:lsid:plazi:treatment:972687861A5EFFD65269FC21FBA58A60:1A50604B1A5DFFD15190FADFFEC08E8C" box="[186,331,1400,1422]" country="Mongolia" name="Hermiin Tsav" pageId="2" pageNumber="3" stateProvince="Omnogovi Province">Hermiin Tsav</location>
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,
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<collectingRegion box="[344,553,1400,1422]" country="Mongolia" name="Omnogovi" pageId="2" pageNumber="3">Omnogovi Province</collectingRegion>
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,
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<collectingCountry box="[566,663,1400,1422]" name="Mongolia" pageId="2" pageNumber="3">Mongolia</collectingCountry>
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<bibRefCitation author="Eberth, D. A." box="[663,682,1396,1415]" journalOrPublisher="Palaeogeogr. Palaeoclimatol. Palaeoecol." pageId="2" pageNumber="3" pagination="29 - 50" part="494" refId="ref5774" refString="13. Eberth, D. A. Stratigraphy and paleoenvironmental evolution of the dinosaurrich Baruungoyot-Nemegt succession (Upper Cretaceous), Nemegt Basin, southern Mongolia. Palaeogeogr. Palaeoclimatol. Palaeoecol. 494, 29 - 50 (2018)." title="Stratigraphy and paleoenvironmental evolution of the dinosaurrich Baruungoyot-Nemegt succession (Upper Cretaceous), Nemegt Basin, southern Mongolia" type="journal article" year="2018">13</bibRefCitation>
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(Supplementary
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<figureCitation box="[209,268,1428,1450]" captionStart="Fig" captionStartId="6.[116,146,1092,1111]" captionTargetBox="[208,686,415,1066]" captionTargetId="figure-950@6.[207,687,414,1067]" captionTargetPageId="6" captionText="Fig. 5 Life reconstruction of Natovenator polydontus (Artwork by Yusik Choi). The reconstruction shows the proposed swimming behaviour of Natovenator polydontus." figureDoi="http://doi.org/10.5281/zenodo.7390408" httpUri="https://zenodo.org/record/7390408/files/figure.png" pageId="2" pageNumber="3">Fig. 5</figureCitation>
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)
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</materialsCitation>
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.
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</paragraph>
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</subSubSection>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.7390396" ID-Zenodo-Dep="7390396" httpUri="https://zenodo.org/record/7390396/files/figure.png" pageId="1" pageNumber="2" startId="1.[116,146,1895,1914]" targetBox="[315,1272,1080,1870]" targetPageId="1">
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<paragraph blockId="1.[116,1471,1894,1970]" pageId="1" pageNumber="2">
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<emphasis bold="true" box="[116,644,1894,1914]" pageId="1" pageNumber="2">
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Fig. 1
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<taxonomicName authorityName="Lee & Lee & Currie & Sissons & Park & Kim & Barsbold & Tsogtbaatar" authorityYear="2022" box="[168,380,1894,1914]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="polydontus">
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<emphasis bold="true" box="[168,380,1894,1914]" italics="true" pageId="1" pageNumber="2">Natovenator polydontus</emphasis>
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</taxonomicName>
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(MPC-D 102/114, holotype).
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</emphasis>
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Photographs (
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<emphasis bold="true" box="[772,783,1895,1914]" pageId="1" pageNumber="2">a</emphasis>
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,
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<emphasis bold="true" box="[793,803,1895,1914]" pageId="1" pageNumber="2">c</emphasis>
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) and line drawings (
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<emphasis bold="true" box="[983,994,1895,1914]" pageId="1" pageNumber="2">b</emphasis>
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,
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<emphasis bold="true" box="[1004,1015,1895,1914]" pageId="1" pageNumber="2">d</emphasis>
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) of the main block containing most of the specimen in opposite views. cav caudal vertebra, co coracoid, cv cervical vertebra, d dentary, dc distal carpal, dv dorsal vertebra, fem femur, fu furcula, h humerus, mx maxilla, ph phalanx, pm premaxilla, r radius, ul ulna.
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</paragraph>
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</caption>
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<footnote box="[678,1378,2007,2023]" pageId="1" pageNumber="2">
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<paragraph blockId="1.[678,1378,2007,2023]" box="[678,1378,2007,2023]" pageId="1" pageNumber="2">(2022) 5:1185 | https://doi.org/10.1038/s42003-022-04119-9 | www.nature.com/commsbio</paragraph>
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</footnote>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.7390398" ID-Zenodo-Dep="7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="2" pageNumber="3" startId="2.[116,146,1093,1112]" targetBox="[155,1432,136,1068]" targetPageId="2">
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<paragraph blockId="2.[116,1471,1093,1224]" pageId="2" pageNumber="3">
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<emphasis bold="true" box="[116,761,1093,1113]" pageId="2" pageNumber="3">
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Fig. 2 Skull of
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<taxonomicName authorityName="Lee & Lee & Currie & Sissons & Park & Kim & Barsbold & Tsogtbaatar" authorityYear="2022" box="[247,459,1093,1113]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="polydontus">
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<emphasis bold="true" box="[247,459,1093,1113]" italics="true" pageId="2" pageNumber="3">Natovenator polydontus</emphasis>
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</taxonomicName>
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(MPC-D 102/114, holotype). a
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<emphasis bold="true" box="[741,750,1094,1113]" italics="true" pageId="2" pageNumber="3">–</emphasis>
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d
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</emphasis>
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Skull in left lateral (
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<emphasis bold="true" box="[940,951,1093,1112]" pageId="2" pageNumber="3">a</emphasis>
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), right lateral (
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<emphasis bold="true" box="[1085,1096,1093,1112]" pageId="2" pageNumber="3">b</emphasis>
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), dorsal (
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<emphasis bold="true" box="[1183,1193,1093,1112]" pageId="2" pageNumber="3">c</emphasis>
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), and ventral (
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<emphasis bold="true" box="[1325,1336,1093,1112]" pageId="2" pageNumber="3">d</emphasis>
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) views.
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<emphasis bold="true" box="[1411,1422,1093,1112]" pageId="2" pageNumber="3">e</emphasis>
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<emphasis box="[1427,1438,1094,1113]" italics="true" pageId="2" pageNumber="3">µ</emphasis>
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CTrendered image sliced at the point marked on
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<emphasis bold="true" box="[521,532,1121,1140]" pageId="2" pageNumber="3">a</emphasis>
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, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view.
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<emphasis bold="true" box="[1311,1318,1121,1140]" pageId="2" pageNumber="3">f</emphasis>
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Micro-computed tomography (
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<emphasis box="[236,247,1149,1168]" italics="true" pageId="2" pageNumber="3">µ</emphasis>
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CT) rendered image of the occipital region in posterior view.
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<emphasis bold="true" box="[777,788,1149,1168]" pageId="2" pageNumber="3">g</emphasis>
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<emphasis box="[793,804,1149,1168]" italics="true" pageId="2" pageNumber="3">µ</emphasis>
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CT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital.
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</paragraph>
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</caption>
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<subSubSection pageId="2" pageNumber="3" type="etymology">
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<paragraph blockId="2.[116,778,1494,1627]" pageId="2" pageNumber="3">
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<emphasis bold="true" box="[116,228,1494,1515]" pageId="2" pageNumber="3">Etymology</emphasis>
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.
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<taxonomicName box="[241,366,1494,1515]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
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<emphasis box="[241,366,1494,1515]" italics="true" pageId="2" pageNumber="3">Natovenator</emphasis>
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</taxonomicName>
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, from the Latin
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<emphasis box="[530,575,1494,1515]" italics="true" pageId="2" pageNumber="3">nato</emphasis>
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(swim) and
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<emphasis box="[702,777,1494,1515]" italics="true" pageId="2" pageNumber="3">venator</emphasis>
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(hunter), in reference to the hypothesized swimming behaviour and piscivorous diet of the new taxon;
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<taxonomicName authorityName="Lee & Lee & Currie & Sissons & Park & Kim & Barsbold & Tsogtbaatar" authorityYear="2022" box="[504,612,1550,1571]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="polydontus">
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<emphasis box="[504,612,1550,1571]" italics="true" pageId="2" pageNumber="3">polydontus</emphasis>
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</taxonomicName>
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, from the Greek
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<emphasis box="[116,165,1577,1598]" italics="true" pageId="2" pageNumber="3">polys</emphasis>
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(many) and
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<emphasis box="[314,372,1577,1598]" italics="true" pageId="2" pageNumber="3">odous</emphasis>
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(tooth) in reference to the unusually many teeth.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="2" pageNumber="3" type="diagnosis">
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<paragraph blockId="2.[116,778,1670,1972]" lastBlockId="2.[810,1471,1279,1497]" pageId="2" pageNumber="3">
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<emphasis bold="true" box="[116,220,1670,1692]" pageId="2" pageNumber="3">Diagnosis</emphasis>
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. A small halszkaraptorine dromaeosaurid with the following autapomorphies: wide groove delimited by a pair of ridges on the anterodorsal surface of the premaxilla, premaxilla with an elongated internarial process that overlies nasal and extends posterior to the external naris, 13 premaxillary teeth with large and incisiviform crowns, first three anteriormost maxillary teeth are greatly reduced and are clustered together with the following tooth without any separations by interdental septa, anteroposteriorly long external naris (about 30% of the preorbital skull length), paroccipital process with a anteroposteriorly broad dorsal surface, elongate maxillary process of the palatine that extends anteriorly beyond the middle of the antorbital fenestra, pterygoid with a deep fossa on the medial surface of the quadrate ramus, distinct posterolaterally oriented projection on the lateral surface of atlas, absence of pleurocoels in cervical vertebrae (not confirmed in the missing fifth cervical centrum), posterolaterally oriented and nearly horizontal proximal shafts in the dorsal ribs, hourglass-shaped metacarpal II with distinctly concave medial and lateral surfaces.
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="5" lastPageNumber="6" pageId="2" pageNumber="3" type="description">
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<paragraph blockId="2.[810,1472,1531,1971]" lastBlockId="4.[116,779,1643,1972]" lastPageId="4" lastPageNumber="5" pageId="2" pageNumber="3">
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<emphasis box="[810,923,1531,1552]" italics="true" pageId="2" pageNumber="3">Description</emphasis>
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. The skull of
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<taxonomicName box="[1113,1238,1531,1552]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
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<emphasis box="[1113,1238,1531,1552]" italics="true" pageId="2" pageNumber="3">Natovenator</emphasis>
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</taxonomicName>
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is nearly complete, although the preorbital region has been affected by compression and is slightly offset from the rest of the skull (
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<figureCitation box="[1301,1409,1586,1608]" captionStart="Fig" captionStartId="1.[116,146,1895,1914]" captionTargetBox="[315,1272,1080,1870]" captionTargetId="figure-513@1.[314,1274,1079,1871]" captionTargetPageId="1" captionText="Fig. 1 Natovenator polydontus (MPC-D 102/114, holotype). Photographs (a, c) and line drawings (b, d) of the main block containing most of the specimen in opposite views. cav caudal vertebra, co coracoid, cv cervical vertebra, d dentary, dc distal carpal, dv dorsal vertebra, fem femur, fu furcula, h humerus, mx maxilla, ph phalanx, pm premaxilla, r radius, ul ulna." figureDoi="http://doi.org/10.5281/zenodo.7390396" httpUri="https://zenodo.org/record/7390396/files/figure.png" pageId="2" pageNumber="3">Figs. 1c, d</figureCitation>
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,
|
||
<figureCitation box="[1422,1471,1585,1608]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="2" pageNumber="3">2a–d</figureCitation>
|
||
and Supplementary
|
||
<figureCitation box="[1023,1093,1614,1636]" captionStart="Fig" captionStartId="1.[116,146,1895,1914]" captionTargetBox="[315,1272,1080,1870]" captionTargetId="figure-513@1.[314,1274,1079,1871]" captionTargetPageId="1" captionText="Fig. 1 Natovenator polydontus (MPC-D 102/114, holotype). Photographs (a, c) and line drawings (b, d) of the main block containing most of the specimen in opposite views. cav caudal vertebra, co coracoid, cv cervical vertebra, d dentary, dc distal carpal, dv dorsal vertebra, fem femur, fu furcula, h humerus, mx maxilla, ph phalanx, pm premaxilla, r radius, ul ulna." figureDoi="http://doi.org/10.5281/zenodo.7390396" httpUri="https://zenodo.org/record/7390396/files/figure.png" pageId="2" pageNumber="3">Figs. 1</figureCitation>
|
||
,
|
||
<figureCitation box="[1110,1124,1614,1636]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="2" pageNumber="3">2</figureCitation>
|
||
). Near the tip of the snout, the premaxilla is marked by a broad groove. The body of the premaxilla is also dorsoventrally low and is perforated by numerous foramina that lead into a complex network of neurovascular chambers (Supplementary
|
||
<figureCitation box="[1091,1163,1726,1748]" captionStart="Fig" captionStartId="1.[116,146,1895,1914]" captionTargetBox="[315,1272,1080,1870]" captionTargetId="figure-513@1.[314,1274,1079,1871]" captionTargetPageId="1" captionText="Fig. 1 Natovenator polydontus (MPC-D 102/114, holotype). Photographs (a, c) and line drawings (b, d) of the main block containing most of the specimen in opposite views. cav caudal vertebra, co coracoid, cv cervical vertebra, d dentary, dc distal carpal, dv dorsal vertebra, fem femur, fu furcula, h humerus, mx maxilla, ph phalanx, pm premaxilla, r radius, ul ulna." figureDoi="http://doi.org/10.5281/zenodo.7390396" httpUri="https://zenodo.org/record/7390396/files/figure.png" pageId="2" pageNumber="3">Fig. 1b</figureCitation>
|
||
) as in
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[1248,1390,1726,1747]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1248,1390,1726,1747]" italics="true" pageId="2" pageNumber="3">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
4. Similarly, the external naris is positioned posteriorly and is level with the premaxilla-maxilla contact (
|
||
<figureCitation box="[1143,1246,1781,1803]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="2" pageNumber="3">Fig. 2a, b</figureCitation>
|
||
), although it is marginally behind this position in
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[1129,1271,1809,1830]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1129,1271,1809,1830]" italics="true" pageId="2" pageNumber="3">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Cau, A." box="[1271,1281,1804,1823]" journalOrPublisher="Nature" pageId="2" pageNumber="3" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
. It is also dorsally placed compared to those of other non-avian theropods and faces dorsolaterally. The exceptionally long external naris and accordingly elongated internarial process of
|
||
<taxonomicName box="[1208,1333,1894,1915]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1208,1333,1894,1915]" italics="true" pageId="2" pageNumber="3">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation box="[1350,1423,1893,1915]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="2" pageNumber="3">Fig. 2c</figureCitation>
|
||
) are unique among dromaeosaurids but comparable to those in aquatic toothed birds
|
||
<bibRefCitation author="Field, D. J." box="[1038,1057,1944,1963]" journalOrPublisher="Nature" pageId="2" pageNumber="3" pagination="96 - 100" part="557" refId="ref5817" refString="14. Field, D. J. et al. Complete Ichthyornis skull illuminates mosaic assembly of the avian head. Nature 557, 96 - 100 (2018)." title="Complete Ichthyornis skull illuminates mosaic assembly of the avian head" type="journal article" year="2018">14</bibRefCitation>
|
||
as well as in therizinosaurs
|
||
<bibRefCitation author="Clark, J. M. & Altangerel, P. & Norell, M. A." box="[1369,1388,1944,1963]" journalOrPublisher="Am. Mus. Novit." pageId="2" pageNumber="3" pagination="1 - 39" part="3115" refId="ref5849" refString="15. Clark, J. M., Altangerel, P. & Norell, M. A. The skull of Erlicosaurus andrewsi, a Late Cretaceous " Segnosaur " (Theropoda: Therizinosauridae) from Mongolia. Am. Mus. Novit. 3115, 1 - 39 (1994)." title="The skull of Erlicosaurus andrewsi, a Late Cretaceous " Segnosaur " (Theropoda: Therizinosauridae) from Mongolia" type="journal article" year="1994">15</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Pu, H." box="[1393,1412,1944,1963]" journalOrPublisher="PLoS ONE" pageId="2" pageNumber="3" pagination="63423" part="8" refId="ref5904" refString="16. Pu, H. et al. An unusual basal therizinosaur dinosaur with an ornithischian dental arrangement from Northeastern China. PLoS ONE 8, e 63423 (2013)." title="An unusual basal therizinosaur dinosaur with an ornithischian dental arrangement from Northeastern China" type="journal article" year="2013">16</bibRefCitation>
|
||
. The frontal is similar to those of other halszkaraptorines
|
||
<bibRefCitation author="Cau, A." box="[647,657,1498,1517]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Turner, A. H. & Pol, D. & Norell, M. A." box="[661,680,1498,1517]" journalOrPublisher="Am. Mus. Novit." pageId="3" pageNumber="4" pagination="1 - 66" part="3722" refId="ref5937" refString="17. Turner, A. H., Pol, D. & Norell, M. A. Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia. Am. Mus. Novit. 3722, 1 - 66 (2011)." title="Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia" type="journal article" year="2011">17</bibRefCitation>
|
||
in that it is vaulted to accommodate a large orbit and has little contribution to the supratemporal fossa. A sharp nuchal crest is formed by the parietal and the squamosal (Supplementary
|
||
<figureCitation box="[566,659,1586,1609]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="3" pageNumber="4">Fig. 2a–e</figureCitation>
|
||
). The latter also produces a shelf that extends over the quadrate head as in other dromaeosaurids
|
||
<bibRefCitation author="Norell, M. A. & Makovicky, P. J." box="[342,361,1637,1656]" journalOrPublisher="University of California Press" pageId="3" pageNumber="4" refId="ref5987" refString="18. Norell, M. A. & Makovicky, P. J. Dromaeosauridae in The Dinosauria (eds Weishampel, D. B., Dodson, P. & Osmolska, H.) Ch. 10 (University of California Press, 2004)." title="Dromaeosauridae in The Dinosauria (eds Weishampel, D. B., Dodson, P. & Osmolska, H.) Ch. 10" type="book" year="2004">18</bibRefCitation>
|
||
. The paroccipital process curves gently on the occiput and has a broad dorsal surface that tapers laterally (
|
||
<figureCitation box="[123,190,1698,1720]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="3" pageNumber="4">Fig. 2f</figureCitation>
|
||
and Supplementary
|
||
<figureCitation box="[408,506,1698,1720]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="3" pageNumber="4">Fig. 2b, e</figureCitation>
|
||
). Its ventrolateral orientation is reminiscent of
|
||
<taxonomicName authorityName="Turner, Pol, Clarke, Erickson & Norell" authorityYear="2007" box="[341,443,1726,1747]" family="Dromaeosauridae" genus="Mahakala" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[341,443,1726,1747]" italics="true" pageId="3" pageNumber="4">Mahakala</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Turner, A. H. & Pol, D. & Norell, M. A." box="[444,463,1721,1740]" journalOrPublisher="Am. Mus. Novit." pageId="3" pageNumber="4" pagination="1 - 66" part="3722" refId="ref5937" refString="17. Turner, A. H., Pol, D. & Norell, M. A. Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia. Am. Mus. Novit. 3722, 1 - 66 (2011)." title="Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia" type="journal article" year="2011">17</bibRefCitation>
|
||
but is different from the more horizontal paroccipital process of
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[475,617,1754,1775]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[475,617,1754,1775]" italics="true" pageId="3" pageNumber="4">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Cau, A." box="[617,627,1749,1768]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
. The occipital condyle is long and constricted at its base. A shallow dorsal tympanic recess on the lateral wall of the braincase is different from the deep one of
|
||
<taxonomicName authorityName="Turner, Pol, Clarke, Erickson & Norell" authorityYear="2007" box="[356,458,1838,1859]" family="Dromaeosauridae" genus="Mahakala" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[356,458,1838,1859]" italics="true" pageId="3" pageNumber="4">Mahakala</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Turner, A. H. & Pol, D. & Norell, M. A." box="[459,478,1833,1852]" journalOrPublisher="Am. Mus. Novit." pageId="3" pageNumber="4" pagination="1 - 66" part="3722" refId="ref5937" refString="17. Turner, A. H., Pol, D. & Norell, M. A. Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia. Am. Mus. Novit. 3722, 1 - 66 (2011)." title="Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia" type="journal article" year="2011">17</bibRefCitation>
|
||
. The palatine is tetraradiate with a greatly elongated maxillary process, which extends anteriorly beyond the level of the mid-antorbital fenestra. The pterygoid is missing its anterior portion (
|
||
<figureCitation box="[501,572,1921,1943]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="3" pageNumber="4">Fig. 2g</figureCitation>
|
||
and Supplementary
|
||
<figureCitation box="[116,208,1948,1971]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="3" pageNumber="4">Fig. 2a–e</figureCitation>
|
||
). A deep fossa on the medial surface of the thin quadrate ramus is not seen in any other dromaeosaurids. The mandibles of
|
||
<taxonomicName box="[810,935,1532,1553]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[810,935,1532,1553]" italics="true" pageId="3" pageNumber="4">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
preserve most of the elements, especially those on the left side (
|
||
<figureCitation box="[938,1054,1559,1581]" captionStart="Fig" captionStartId="1.[116,146,1895,1914]" captionTargetBox="[315,1272,1080,1870]" captionTargetId="figure-513@1.[314,1274,1079,1871]" captionTargetPageId="1" captionText="Fig. 1 Natovenator polydontus (MPC-D 102/114, holotype). Photographs (a, c) and line drawings (b, d) of the main block containing most of the specimen in opposite views. cav caudal vertebra, co coracoid, cv cervical vertebra, d dentary, dc distal carpal, dv dorsal vertebra, fem femur, fu furcula, h humerus, mx maxilla, ph phalanx, pm premaxilla, r radius, ul ulna." figureDoi="http://doi.org/10.5281/zenodo.7390396" httpUri="https://zenodo.org/record/7390396/files/figure.png" pageId="3" pageNumber="4">Fig. 1a, b, d</figureCitation>
|
||
and Supplementary
|
||
<figureCitation box="[1262,1338,1559,1581]" captionStart="Fig" captionStartId="1.[116,146,1895,1914]" captionTargetBox="[315,1272,1080,1870]" captionTargetId="figure-513@1.[314,1274,1079,1871]" captionTargetPageId="1" captionText="Fig. 1 Natovenator polydontus (MPC-D 102/114, holotype). Photographs (a, c) and line drawings (b, d) of the main block containing most of the specimen in opposite views. cav caudal vertebra, co coracoid, cv cervical vertebra, d dentary, dc distal carpal, dv dorsal vertebra, fem femur, fu furcula, h humerus, mx maxilla, ph phalanx, pm premaxilla, r radius, ul ulna." figureDoi="http://doi.org/10.5281/zenodo.7390396" httpUri="https://zenodo.org/record/7390396/files/figure.png" pageId="3" pageNumber="4">Figs. 1a</figureCitation>
|
||
,
|
||
<figureCitation box="[1349,1363,1559,1581]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="3" pageNumber="4">2</figureCitation>
|
||
). Each jaw is characterized by a slender dentary with nearly parallel dorsal and ventral margins, a surangular partially fused with the articular, a distinctive surangular shelf, and a fan-shaped retroarticular process that protrudes dorsomedially. The upper dentition of
|
||
<taxonomicName box="[901,1026,1699,1720]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[901,1026,1699,1720]" italics="true" pageId="3" pageNumber="4">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
is heterodont as the premaxillary teeth are morphologically distinct from the maxillary teeth (
|
||
<figureCitation box="[1344,1471,1726,1748]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="3" pageNumber="4">Fig. 2a, b, e</figureCitation>
|
||
and Supplementary
|
||
<figureCitation box="[1023,1124,1754,1776]" captionStart="Fig" captionStartId="1.[116,146,1895,1914]" captionTargetBox="[315,1272,1080,1870]" captionTargetId="figure-513@1.[314,1274,1079,1871]" captionTargetPageId="1" captionText="Fig. 1 Natovenator polydontus (MPC-D 102/114, holotype). Photographs (a, c) and line drawings (b, d) of the main block containing most of the specimen in opposite views. cav caudal vertebra, co coracoid, cv cervical vertebra, d dentary, dc distal carpal, dv dorsal vertebra, fem femur, fu furcula, h humerus, mx maxilla, ph phalanx, pm premaxilla, r radius, ul ulna." figureDoi="http://doi.org/10.5281/zenodo.7390396" httpUri="https://zenodo.org/record/7390396/files/figure.png" pageId="3" pageNumber="4">Fig. 1a, c</figureCitation>
|
||
). There are unusually numerous premaxillary teeth tightly packed without any separation of the alveoli by bony septa. The roots of the teeth are long, and the crowns are tall and incisiviform as in
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[1201,1343,1838,1859]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1201,1343,1838,1859]" italics="true" pageId="3" pageNumber="4">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Cau, A." box="[1344,1354,1833,1852]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
. Moreover, the large replacement teeth in the premaxilla suggest that the replacement of the premaxillary teeth was delayed as in
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[810,952,1921,1942]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[810,952,1921,1942]" italics="true" pageId="3" pageNumber="4">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Cau, A." box="[952,962,1916,1935]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
. However, the number of teeth in each premaxilla is 13 in
|
||
<taxonomicName box="[904,1029,1950,1971]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[904,1029,1950,1971]" italics="true" pageId="3" pageNumber="4">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
, whereas it is only 11 in
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[1313,1455,1949,1970]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1313,1455,1949,1970]" italics="true" pageId="3" pageNumber="4">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Cau, A." box="[1456,1466,1944,1963]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
. In the maxilla, the three most anterior maxillary teeth are markedly shorter than the premaxillary teeth and the more posterior maxillary teeth. This pattern is also observed in
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[116,258,1726,1747]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[116,258,1726,1747]" italics="true" pageId="4" pageNumber="5">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
, although the number of shorter maxillary teeth differs as it has two reduced ones
|
||
<bibRefCitation author="Cau, A." box="[462,472,1749,1768]" journalOrPublisher="PeerJ" pageId="4" pageNumber="5" pagination="8672" part="8" refId="ref5542" refString="7. Cau, A. The body plan of Halszkaraptor escuilliei (Dinosauria, Theropoda) is not a transitional form along the evolution of dromaeosaurid hypercarnivory. PeerJ 8, e 8672 (2020)." title="The body plan of Halszkaraptor escuilliei (Dinosauria, Theropoda) is not a transitional form along the evolution of dromaeosaurid hypercarnivory" type="journal article" year="2020">7</bibRefCitation>
|
||
. Both the maxillary and dentary teeth have sharp fang-like crowns that lack serrations. Although posteriormost parts are poorly preserved, there are at least 23 alveoli in each of the maxilla and dentary, which suggests high numbers of teeth in both elements.
|
||
</paragraph>
|
||
<footnote box="[424,1123,2007,2023]" pageId="2" pageNumber="3">
|
||
<paragraph blockId="2.[424,1123,2007,2023]" box="[424,1123,2007,2023]" pageId="2" pageNumber="3">(2022) 5:1185 | https://doi.org/10.1038/s42003-022-04119-9 | www.nature.com/commsbio</paragraph>
|
||
</footnote>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.7390404" ID-Zenodo-Dep="7390404" httpUri="https://zenodo.org/record/7390404/files/figure.png" pageId="3" pageNumber="4" startId="3.[116,146,1298,1317]" targetBox="[155,1432,136,1271]" targetPageId="3">
|
||
<paragraph blockId="3.[116,1472,1297,1456]" pageId="3" pageNumber="4">
|
||
<emphasis bold="true" box="[116,1126,1297,1317]" pageId="3" pageNumber="4">
|
||
Fig. 3 Postcranial elements and phylogenetic position of
|
||
<taxonomicName authorityName="Lee & Lee & Currie & Sissons & Park & Kim & Barsbold & Tsogtbaatar" authorityYear="2022" box="[632,844,1297,1317]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="polydontus">
|
||
<emphasis bold="true" box="[632,844,1297,1317]" italics="true" pageId="3" pageNumber="4">Natovenator polydontus</emphasis>
|
||
</taxonomicName>
|
||
(MPC-D 102/114, holotype). a
|
||
</emphasis>
|
||
Anterior cervical vertebrae in left lateral view.
|
||
<emphasis bold="true" box="[165,176,1326,1345]" pageId="3" pageNumber="4">b</emphasis>
|
||
Axis and third cervical vertebra in dorsal view.
|
||
<emphasis bold="true" box="[576,586,1326,1345]" pageId="3" pageNumber="4">c</emphasis>
|
||
Fourth cervical vertebra in dorsal view.
|
||
<emphasis bold="true" box="[921,932,1326,1345]" pageId="3" pageNumber="4">d</emphasis>
|
||
Posterior cervical vertebrae in right lateral view.
|
||
<emphasis bold="true" box="[1345,1356,1326,1345]" pageId="3" pageNumber="4">e</emphasis>
|
||
Dorsal series in right lateral view.
|
||
<emphasis bold="true" box="[291,298,1353,1372]" pageId="3" pageNumber="4">f</emphasis>
|
||
Anterior caudal vertebrae in right lateral view.
|
||
<emphasis bold="true" box="[705,716,1353,1372]" pageId="3" pageNumber="4">g</emphasis>
|
||
Left forearm elements in medial view and manus in ventral view.
|
||
<emphasis bold="true" box="[1284,1295,1353,1372]" pageId="3" pageNumber="4">h</emphasis>
|
||
Right foot in ventral view.
|
||
<emphasis bold="true" box="[166,172,1381,1400]" pageId="3" pageNumber="4">i</emphasis>
|
||
Phylogenetic position of
|
||
<taxonomicName box="[388,488,1381,1401]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis box="[388,488,1381,1401]" italics="true" pageId="3" pageNumber="4">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
in
|
||
<taxonomicName authorityName="Matthew and Brown" authorityYear="1922" box="[514,668,1381,1400]" family="Dromaeosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Dromaeosauridae</taxonomicName>
|
||
. Numbers at each node indicate Bremer support values. at atlas, c3 third cervical vertebra, c4 fourth cervical vertebra, c7 seventh cervical vertebra, c9 ninth cervical vertebra, ch chevron, d7 seventh dorsal vertebra, fem femur, mc I metacarpal I, mt III metatarsal III, mt IV metatarsal IV, poz postzygapophysis, prz prezygapophysis, r radius, r7 seventh dorsal rib, ul ulna, I-2 pedal phalanx I-2.
|
||
</paragraph>
|
||
</caption>
|
||
<footnote box="[678,1378,2007,2023]" pageId="3" pageNumber="4">
|
||
<paragraph blockId="3.[678,1378,2007,2023]" box="[678,1378,2007,2023]" pageId="3" pageNumber="4">(2022) 5:1185 | https://doi.org/10.1038/s42003-022-04119-9 | www.nature.com/commsbio</paragraph>
|
||
</footnote>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.7390406" ID-Zenodo-Dep="7390406" httpUri="https://zenodo.org/record/7390406/files/figure.png" pageId="4" pageNumber="5" startId="4.[116,146,1473,1492]" targetBox="[155,1433,141,1448]" targetPageId="4">
|
||
<paragraph blockId="4.[116,1471,1472,1604]" pageId="4" pageNumber="5">
|
||
<emphasis bold="true" box="[116,1409,1472,1492]" pageId="4" pageNumber="5">
|
||
Fig. 4 Body plan of
|
||
<taxonomicName authorityName="Lee & Lee & Currie & Sissons & Park & Kim & Barsbold & Tsogtbaatar" authorityYear="2022" box="[290,501,1472,1492]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="polydontus">
|
||
<emphasis bold="true" box="[290,501,1472,1492]" italics="true" pageId="4" pageNumber="5">Natovenator polydontus</emphasis>
|
||
</taxonomicName>
|
||
(MPC-D 102/114, holotype) and dorsal rib morphology of various diving birds and terrestrial taxa. a
|
||
</emphasis>
|
||
Dorsal series of
|
||
<taxonomicName box="[199,302,1500,1520]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[199,302,1500,1520]" italics="true" pageId="4" pageNumber="5">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
in ventral view.
|
||
<emphasis bold="true" box="[451,462,1501,1520]" pageId="4" pageNumber="5">b</emphasis>
|
||
Reconstruction of dorsal vertebrae and ribs of
|
||
<taxonomicName box="[888,991,1500,1520]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[888,991,1500,1520]" italics="true" pageId="4" pageNumber="5">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
in left lateral view.
|
||
<emphasis bold="true" box="[1170,1180,1501,1520]" pageId="4" pageNumber="5">c</emphasis>
|
||
Skeletal reconstruction of
|
||
<taxonomicName box="[116,219,1528,1548]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[116,219,1528,1548]" italics="true" pageId="4" pageNumber="5">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
with missing parts in dark grey.
|
||
<emphasis bold="true" box="[515,542,1529,1548]" pageId="4" pageNumber="5">
|
||
d
|
||
<emphasis bold="true" box="[527,537,1529,1548]" italics="true" pageId="4" pageNumber="5">–</emphasis>
|
||
i
|
||
</emphasis>
|
||
Dorsal rib morphology of
|
||
<taxonomicName box="[780,883,1528,1548]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[780,883,1528,1548]" italics="true" pageId="4" pageNumber="5">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<emphasis bold="true" box="[898,909,1529,1548]" pageId="4" pageNumber="5">d</emphasis>
|
||
), diving birds (
|
||
<emphasis bold="true" box="[1048,1073,1529,1548]" pageId="4" pageNumber="5">
|
||
e
|
||
<emphasis bold="true" box="[1059,1069,1529,1548]" italics="true" pageId="4" pageNumber="5">–</emphasis>
|
||
i
|
||
</emphasis>
|
||
), common ostrich (
|
||
<emphasis bold="true" box="[1254,1259,1529,1548]" pageId="4" pageNumber="5">j</emphasis>
|
||
), and
|
||
<taxonomicName box="[1319,1390,1528,1548]" family="Dromaeosauridae" genus="Shri" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="devi">
|
||
<emphasis box="[1319,1390,1528,1548]" italics="true" pageId="4" pageNumber="5">Shri devi</emphasis>
|
||
</taxonomicName>
|
||
, a likely terrestrial dromaeosaurid from the Baruungoyot Formation (
|
||
<emphasis bold="true" box="[655,666,1557,1576]" pageId="4" pageNumber="5">k</emphasis>
|
||
) in ventral view (not to scale).
|
||
<emphasis bold="true" box="[953,959,1557,1576]" pageId="4" pageNumber="5">l</emphasis>
|
||
Reconstruction of the fourth dorsal vertebra with corresponding ribs in anterior view. d2 second dorsal vertebra, r2 second dorsal rib, r3 third dorsal rib, r4 fourth dorsal rib.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="4.[116,779,1643,1972]" lastBlockId="5.[116,779,136,856]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
|
||
The neck of
|
||
<taxonomicName box="[271,396,1895,1916]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[271,396,1895,1916]" italics="true" pageId="4" pageNumber="5">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
, as preserved, is twisted and includes ten elongated cervical vertebrae, although most of the 5th cervical is missing (
|
||
<figureCitation box="[232,300,1950,1972]" captionStart="Fig" captionStartId="1.[116,146,1895,1914]" captionTargetBox="[315,1272,1080,1870]" captionTargetId="figure-513@1.[314,1274,1079,1871]" captionTargetPageId="1" captionText="Fig. 1 Natovenator polydontus (MPC-D 102/114, holotype). Photographs (a, c) and line drawings (b, d) of the main block containing most of the specimen in opposite views. cav caudal vertebra, co coracoid, cv cervical vertebra, d dentary, dc distal carpal, dv dorsal vertebra, fem femur, fu furcula, h humerus, mx maxilla, ph phalanx, pm premaxilla, r radius, ul ulna." figureDoi="http://doi.org/10.5281/zenodo.7390396" httpUri="https://zenodo.org/record/7390396/files/figure.png" pageId="4" pageNumber="5">Figs. 1</figureCitation>
|
||
,
|
||
<figureCitation box="[314,365,1949,1972]" captionStart="Fig" captionStartId="3.[116,146,1298,1317]" captionTargetBox="[155,1432,136,1271]" captionTargetId="figure-371@3.[154,1434,135,1273]" captionTargetPageId="3" captionText="Fig. 3 Postcranial elements and phylogenetic position of Natovenator polydontus (MPC-D 102/114, holotype). a Anterior cervical vertebrae in left lateral view. b Axis and third cervical vertebra in dorsal view. c Fourth cervical vertebra in dorsal view.d Posterior cervical vertebrae in right lateral view. e Dorsal series in right lateral view. f Anterior caudal vertebrae in right lateral view. g Left forearm elements in medial view and manus in ventral view. h Right foot in ventral view. i Phylogenetic position of Natovenator in Dromaeosauridae. Numbers at each node indicate Bremer support values. at atlas, c3 third cervical vertebra, c4 fourth cervical vertebra, c7 seventh cervical vertebra, c9 ninth cervical vertebra, ch chevron, d7 seventh dorsal vertebra, fem femur, mc I metacarpal I, mt III metatarsal III, mt IV metatarsal IV, poz postzygapophysis, prz prezygapophysis, r radius, r7 seventh dorsal rib, ul ulna, I-2 pedal phalanx I-2." figureDoi="http://doi.org/10.5281/zenodo.7390404" httpUri="https://zenodo.org/record/7390404/files/figure.png" pageId="4" pageNumber="5">3a–d</figureCitation>
|
||
). This elongation of the cervicals results in a noticeably longer neck than those of most dromaeosaurids and is estimated to be longer than the dorsal series. It is, however, proportionately shorter than that of
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[1190,1332,1698,1719]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1190,1332,1698,1719]" italics="true" pageId="4" pageNumber="5">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
, which has a neck as long as its dorsal and sacral vertebra combined
|
||
<bibRefCitation author="Cau, A." box="[1365,1375,1721,1740]" journalOrPublisher="Nature" pageId="4" pageNumber="5" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
. Another peculiarity in the neck of the
|
||
<taxonomicName box="[1154,1279,1755,1776]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1154,1279,1755,1776]" italics="true" pageId="4" pageNumber="5">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
is a pronounced posterolaterally extending projection on the neurapophysis of the atlas (
|
||
<figureCitation box="[871,941,1810,1832]" captionStart="Fig" captionStartId="3.[116,146,1298,1317]" captionTargetBox="[155,1432,136,1271]" captionTargetId="figure-371@3.[154,1434,135,1273]" captionTargetPageId="3" captionText="Fig. 3 Postcranial elements and phylogenetic position of Natovenator polydontus (MPC-D 102/114, holotype). a Anterior cervical vertebrae in left lateral view. b Axis and third cervical vertebra in dorsal view. c Fourth cervical vertebra in dorsal view.d Posterior cervical vertebrae in right lateral view. e Dorsal series in right lateral view. f Anterior caudal vertebrae in right lateral view. g Left forearm elements in medial view and manus in ventral view. h Right foot in ventral view. i Phylogenetic position of Natovenator in Dromaeosauridae. Numbers at each node indicate Bremer support values. at atlas, c3 third cervical vertebra, c4 fourth cervical vertebra, c7 seventh cervical vertebra, c9 ninth cervical vertebra, ch chevron, d7 seventh dorsal vertebra, fem femur, mc I metacarpal I, mt III metatarsal III, mt IV metatarsal IV, poz postzygapophysis, prz prezygapophysis, r radius, r7 seventh dorsal rib, ul ulna, I-2 pedal phalanx I-2." figureDoi="http://doi.org/10.5281/zenodo.7390404" httpUri="https://zenodo.org/record/7390404/files/figure.png" pageId="4" pageNumber="5">Fig. 3a</figureCitation>
|
||
and Supplementary
|
||
<figureCitation box="[1158,1280,1810,1832]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="4" pageNumber="5">Fig. 2b, c, e</figureCitation>
|
||
). The postzygapophyses of each anterior cervical are fused into a single lobe-like process as in
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[972,1114,1866,1887]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[972,1114,1866,1887]" italics="true" pageId="4" pageNumber="5">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Cau, A." box="[1115,1125,1861,1880]" journalOrPublisher="Nature" pageId="4" pageNumber="5" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
. Pleurocoels are absent in the cervical vertebrae. In contrast,
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[1128,1270,1894,1915]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1128,1270,1894,1915]" italics="true" pageId="4" pageNumber="5">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
has pleurocoels on its 7th–9th cervicals
|
||
<bibRefCitation author="Cau, A." box="[1011,1021,1917,1936]" journalOrPublisher="Nature" pageId="4" pageNumber="5" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
. A total of 12 dorsal vertebrae are preserved (
|
||
<figureCitation box="[817,924,1950,1972]" captionStart="Fig" captionStartId="1.[116,146,1895,1914]" captionTargetBox="[315,1272,1080,1870]" captionTargetId="figure-513@1.[314,1274,1079,1871]" captionTargetPageId="1" captionText="Fig. 1 Natovenator polydontus (MPC-D 102/114, holotype). Photographs (a, c) and line drawings (b, d) of the main block containing most of the specimen in opposite views. cav caudal vertebra, co coracoid, cv cervical vertebra, d dentary, dc distal carpal, dv dorsal vertebra, fem femur, fu furcula, h humerus, mx maxilla, ph phalanx, pm premaxilla, r radius, ul ulna." figureDoi="http://doi.org/10.5281/zenodo.7390396" httpUri="https://zenodo.org/record/7390396/files/figure.png" pageId="4" pageNumber="5">Figs. 1a, b</figureCitation>
|
||
,
|
||
<figureCitation box="[937,961,1950,1971]" captionStart="Fig" captionStartId="3.[116,146,1298,1317]" captionTargetBox="[155,1432,136,1271]" captionTargetId="figure-371@3.[154,1434,135,1273]" captionTargetPageId="3" captionText="Fig. 3 Postcranial elements and phylogenetic position of Natovenator polydontus (MPC-D 102/114, holotype). a Anterior cervical vertebrae in left lateral view. b Axis and third cervical vertebra in dorsal view. c Fourth cervical vertebra in dorsal view.d Posterior cervical vertebrae in right lateral view. e Dorsal series in right lateral view. f Anterior caudal vertebrae in right lateral view. g Left forearm elements in medial view and manus in ventral view. h Right foot in ventral view. i Phylogenetic position of Natovenator in Dromaeosauridae. Numbers at each node indicate Bremer support values. at atlas, c3 third cervical vertebra, c4 fourth cervical vertebra, c7 seventh cervical vertebra, c9 ninth cervical vertebra, ch chevron, d7 seventh dorsal vertebra, fem femur, mc I metacarpal I, mt III metatarsal III, mt IV metatarsal IV, poz postzygapophysis, prz prezygapophysis, r radius, r7 seventh dorsal rib, ul ulna, I-2 pedal phalanx I-2." figureDoi="http://doi.org/10.5281/zenodo.7390404" httpUri="https://zenodo.org/record/7390404/files/figure.png" pageId="4" pageNumber="5">3e</figureCitation>
|
||
,
|
||
<figureCitation box="[974,997,1950,1971]" captionStart="Fig" captionStartId="4.[116,146,1473,1492]" captionTargetBox="[155,1433,141,1448]" captionTargetId="figure-263@4.[154,1434,135,1449]" captionTargetPageId="4" captionText="Fig. 4 Body plan of Natovenator polydontus (MPC-D 102/114, holotype) and dorsal rib morphology of various diving birds and terrestrial taxa. a Dorsal series of Natovenator in ventral view. b Reconstruction of dorsal vertebrae and ribs of Natovenator in left lateral view. c Skeletal reconstruction of Natovenator with missing parts in dark grey. d–i Dorsal rib morphology of Natovenator (d), diving birds (e–i), common ostrich (j), and Shri devi, a likely terrestrial dromaeosaurid from the Baruungoyot Formation (k) in ventral view (not to scale). l Reconstruction of the fourth dorsal vertebra with corresponding ribs in anterior view. d2 second dorsal vertebra, r2 second dorsal rib, r3 third dorsal rib, r4 fourth dorsal rib." figureDoi="http://doi.org/10.5281/zenodo.7390406" httpUri="https://zenodo.org/record/7390406/files/figure.png" pageId="4" pageNumber="5">4a</figureCitation>
|
||
and Supplementary
|
||
<figureCitation box="[1213,1318,1949,1972]" captionStart="Fig" captionStartId="3.[116,146,1298,1317]" captionTargetBox="[155,1432,136,1271]" captionTargetId="figure-371@3.[154,1434,135,1273]" captionTargetPageId="3" captionText="Fig. 3 Postcranial elements and phylogenetic position of Natovenator polydontus (MPC-D 102/114, holotype). a Anterior cervical vertebrae in left lateral view. b Axis and third cervical vertebra in dorsal view. c Fourth cervical vertebra in dorsal view.d Posterior cervical vertebrae in right lateral view. e Dorsal series in right lateral view. f Anterior caudal vertebrae in right lateral view. g Left forearm elements in medial view and manus in ventral view. h Right foot in ventral view. i Phylogenetic position of Natovenator in Dromaeosauridae. Numbers at each node indicate Bremer support values. at atlas, c3 third cervical vertebra, c4 fourth cervical vertebra, c7 seventh cervical vertebra, c9 ninth cervical vertebra, ch chevron, d7 seventh dorsal vertebra, fem femur, mc I metacarpal I, mt III metatarsal III, mt IV metatarsal IV, poz postzygapophysis, prz prezygapophysis, r radius, r7 seventh dorsal rib, ul ulna, I-2 pedal phalanx I-2." figureDoi="http://doi.org/10.5281/zenodo.7390404" httpUri="https://zenodo.org/record/7390404/files/figure.png" pageId="4" pageNumber="5">Figs. 3a–d</figureCitation>
|
||
). They all lack pleurocoels, and their parapophyses on the anterior and middorsals are placed high on the anterodorsal end of each centrum. Interestingly, the positions of the parapophyses are similar to those of hesperornithiforms
|
||
<bibRefCitation author="Marsh, O. C." box="[399,418,215,234]" journalOrPublisher="Government Printing Office" pageId="5" pageNumber="6" refId="ref6037" refString="19. Marsh, O. C. Odontornithes: A Monograph on the Extinct Toothed Birds of North America (Government Printing Office, 1880)." title="Odontornithes: A Monograph on the Extinct Toothed Birds of North America" type="book" year="1880">19</bibRefCitation>
|
||
<bibRefCitation author="Tokaryk, T. T. & Harington, C. R." box="[418,428,215,233]" journalOrPublisher="J. Paleontol." pageId="5" pageNumber="6" pagination="1010 - 1012" part="66" refId="ref6065" refString="20. Tokaryk, T. T. & Harington, C. R. Baptornis sp. (Aves: Hesperornithiformes) from the Judith River Formation (Campanian) of Saskatchewan, Canada. J. Paleontol. 66, 1010 - 1012 (1992)." title="Baptornis sp. (Aves: Hesperornithiformes) from the Judith River Formation (Campanian) of Saskatchewan, Canada" type="journal article" year="1992">–</bibRefCitation>
|
||
<bibRefCitation author="Rees, J. & Lindgren, J." box="[428,447,215,234]" journalOrPublisher="Palaeontology" pageId="5" pageNumber="6" pagination="1321 - 1329" part="48" refId="ref6114" refString="21. Rees, J. & Lindgren, J. Aquatic birds from the Upper Cretaceous (Lower Campanian) of Sweden and the biology and distribution of hesperornithiforms. Palaeontology 48, 1321 - 1329 (2005)." title="Aquatic birds from the Upper Cretaceous (Lower Campanian) of Sweden and the biology and distribution of hesperornithiforms" type="journal article" year="2005">21</bibRefCitation>
|
||
rather than other dromaeosaurids such as
|
||
<taxonomicName authorityName="Ostrom" authorityYear="1969" box="[231,359,248,269]" class="Reptilia" family="Dromaeosauridae" genus="Deinonychus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[231,359,248,269]" italics="true" pageId="5" pageNumber="6">Deinonychus</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Ostrom, J. H." box="[359,378,243,262]" journalOrPublisher="B. Peabody Mus. Nat. Hist." pageId="5" pageNumber="6" pagination="1 - 165" part="30" refId="ref6155" refString="22. Ostrom, J. H. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. B. Peabody Mus. Nat. Hist. 30, 1 - 165 (1969)." title="Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana" type="journal article" year="1969">22</bibRefCitation>
|
||
or
|
||
<taxonomicName authorityName="Osborn" authorityYear="1924" box="[413,536,248,269]" class="Reptilia" family="Dromaeosauridae" genus="Velociraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[413,536,248,269]" italics="true" pageId="5" pageNumber="6">Velociraptor</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Norell, M. A. & Makovicky, P. J." box="[536,555,243,262]" journalOrPublisher="Am. Mus. Novit." pageId="5" pageNumber="6" pagination="1 - 45" part="3282" refId="ref6196" refString="23. Norell, M. A. & Makovicky, P. J. Important features of the dromaeosaurid skeleton II: information from newly collected specimens of Velociraptor mongoliensis. Am. Mus. Novit. 3282, 1 - 45 (1999)." title="Important features of the dromaeosaurid skeleton II: information from newly collected specimens of Velociraptor mongoliensis" type="journal article" year="1999">23</bibRefCitation>
|
||
. The preserved dorsal ribs, articulated with the second to seventh dorsals, are flattened and posteriorly oriented (
|
||
<figureCitation box="[380,448,304,326]" captionStart="Fig" captionStartId="1.[116,146,1895,1914]" captionTargetBox="[315,1272,1080,1870]" captionTargetId="figure-513@1.[314,1274,1079,1871]" captionTargetPageId="1" captionText="Fig. 1 Natovenator polydontus (MPC-D 102/114, holotype). Photographs (a, c) and line drawings (b, d) of the main block containing most of the specimen in opposite views. cav caudal vertebra, co coracoid, cv cervical vertebra, d dentary, dc distal carpal, dv dorsal vertebra, fem femur, fu furcula, h humerus, mx maxilla, ph phalanx, pm premaxilla, r radius, ul ulna." figureDoi="http://doi.org/10.5281/zenodo.7390396" httpUri="https://zenodo.org/record/7390396/files/figure.png" pageId="5" pageNumber="6">Figs. 1</figureCitation>
|
||
,
|
||
<figureCitation box="[463,487,304,325]" captionStart="Fig" captionStartId="3.[116,146,1298,1317]" captionTargetBox="[155,1432,136,1271]" captionTargetId="figure-371@3.[154,1434,135,1273]" captionTargetPageId="3" captionText="Fig. 3 Postcranial elements and phylogenetic position of Natovenator polydontus (MPC-D 102/114, holotype). a Anterior cervical vertebrae in left lateral view. b Axis and third cervical vertebra in dorsal view. c Fourth cervical vertebra in dorsal view.d Posterior cervical vertebrae in right lateral view. e Dorsal series in right lateral view. f Anterior caudal vertebrae in right lateral view. g Left forearm elements in medial view and manus in ventral view. h Right foot in ventral view. i Phylogenetic position of Natovenator in Dromaeosauridae. Numbers at each node indicate Bremer support values. at atlas, c3 third cervical vertebra, c4 fourth cervical vertebra, c7 seventh cervical vertebra, c9 ninth cervical vertebra, ch chevron, d7 seventh dorsal vertebra, fem femur, mc I metacarpal I, mt III metatarsal III, mt IV metatarsal IV, poz postzygapophysis, prz prezygapophysis, r radius, r7 seventh dorsal rib, ul ulna, I-2 pedal phalanx I-2." figureDoi="http://doi.org/10.5281/zenodo.7390404" httpUri="https://zenodo.org/record/7390404/files/figure.png" pageId="5" pageNumber="6">3e</figureCitation>
|
||
,
|
||
<figureCitation box="[500,551,303,326]" captionStart="Fig" captionStartId="4.[116,146,1473,1492]" captionTargetBox="[155,1433,141,1448]" captionTargetId="figure-263@4.[154,1434,135,1449]" captionTargetPageId="4" captionText="Fig. 4 Body plan of Natovenator polydontus (MPC-D 102/114, holotype) and dorsal rib morphology of various diving birds and terrestrial taxa. a Dorsal series of Natovenator in ventral view. b Reconstruction of dorsal vertebrae and ribs of Natovenator in left lateral view. c Skeletal reconstruction of Natovenator with missing parts in dark grey. d–i Dorsal rib morphology of Natovenator (d), diving birds (e–i), common ostrich (j), and Shri devi, a likely terrestrial dromaeosaurid from the Baruungoyot Formation (k) in ventral view (not to scale). l Reconstruction of the fourth dorsal vertebra with corresponding ribs in anterior view. d2 second dorsal vertebra, r2 second dorsal rib, r3 third dorsal rib, r4 fourth dorsal rib." figureDoi="http://doi.org/10.5281/zenodo.7390406" httpUri="https://zenodo.org/record/7390406/files/figure.png" pageId="5" pageNumber="6">4a–d</figureCitation>
|
||
). The proximal shafts are also nearly horizontal, which is indicative of a dorsoventrally compressed ribcage. Each proximal caudal vertebra has a long centrum and horizontal zygapophyses with expanded laminae (
|
||
<figureCitation box="[123,188,415,437]" captionStart="Fig" captionStartId="3.[116,146,1298,1317]" captionTargetBox="[155,1432,136,1271]" captionTargetId="figure-371@3.[154,1434,135,1273]" captionTargetPageId="3" captionText="Fig. 3 Postcranial elements and phylogenetic position of Natovenator polydontus (MPC-D 102/114, holotype). a Anterior cervical vertebrae in left lateral view. b Axis and third cervical vertebra in dorsal view. c Fourth cervical vertebra in dorsal view.d Posterior cervical vertebrae in right lateral view. e Dorsal series in right lateral view. f Anterior caudal vertebrae in right lateral view. g Left forearm elements in medial view and manus in ventral view. h Right foot in ventral view. i Phylogenetic position of Natovenator in Dromaeosauridae. Numbers at each node indicate Bremer support values. at atlas, c3 third cervical vertebra, c4 fourth cervical vertebra, c7 seventh cervical vertebra, c9 ninth cervical vertebra, ch chevron, d7 seventh dorsal vertebra, fem femur, mc I metacarpal I, mt III metatarsal III, mt IV metatarsal IV, poz postzygapophysis, prz prezygapophysis, r radius, r7 seventh dorsal rib, ul ulna, I-2 pedal phalanx I-2." figureDoi="http://doi.org/10.5281/zenodo.7390404" httpUri="https://zenodo.org/record/7390404/files/figure.png" pageId="5" pageNumber="6">Fig. 3f</figureCitation>
|
||
and Supplementary
|
||
<figureCitation box="[402,489,415,438]" captionStart="Fig" captionStartId="3.[116,146,1298,1317]" captionTargetBox="[155,1432,136,1271]" captionTargetId="figure-371@3.[154,1434,135,1273]" captionTargetPageId="3" captionText="Fig. 3 Postcranial elements and phylogenetic position of Natovenator polydontus (MPC-D 102/114, holotype). a Anterior cervical vertebrae in left lateral view. b Axis and third cervical vertebra in dorsal view. c Fourth cervical vertebra in dorsal view.d Posterior cervical vertebrae in right lateral view. e Dorsal series in right lateral view. f Anterior caudal vertebrae in right lateral view. g Left forearm elements in medial view and manus in ventral view. h Right foot in ventral view. i Phylogenetic position of Natovenator in Dromaeosauridae. Numbers at each node indicate Bremer support values. at atlas, c3 third cervical vertebra, c4 fourth cervical vertebra, c7 seventh cervical vertebra, c9 ninth cervical vertebra, ch chevron, d7 seventh dorsal vertebra, fem femur, mc I metacarpal I, mt III metatarsal III, mt IV metatarsal IV, poz postzygapophysis, prz prezygapophysis, r radius, r7 seventh dorsal rib, ul ulna, I-2 pedal phalanx I-2." figureDoi="http://doi.org/10.5281/zenodo.7390404" httpUri="https://zenodo.org/record/7390404/files/figure.png" pageId="5" pageNumber="6">Fig. 3e–i</figureCitation>
|
||
), all of which are characters shared with other halszkaraptorines
|
||
<bibRefCitation author="Cau, A." box="[491,501,438,457]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Turner, A. H. & Pol, D. & Norell, M. A." box="[505,524,438,457]" journalOrPublisher="Am. Mus. Novit." pageId="5" pageNumber="6" pagination="1 - 66" part="3722" refId="ref5937" refString="17. Turner, A. H., Pol, D. & Norell, M. A. Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia. Am. Mus. Novit. 3722, 1 - 66 (2011)." title="Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia" type="journal article" year="2011">17</bibRefCitation>
|
||
. The forelimb elements are partially exposed (
|
||
<figureCitation box="[364,486,471,493]" captionStart="Fig" captionStartId="1.[116,146,1895,1914]" captionTargetBox="[315,1272,1080,1870]" captionTargetId="figure-513@1.[314,1274,1079,1871]" captionTargetPageId="1" captionText="Fig. 1 Natovenator polydontus (MPC-D 102/114, holotype). Photographs (a, c) and line drawings (b, d) of the main block containing most of the specimen in opposite views. cav caudal vertebra, co coracoid, cv cervical vertebra, d dentary, dc distal carpal, dv dorsal vertebra, fem femur, fu furcula, h humerus, mx maxilla, ph phalanx, pm premaxilla, r radius, ul ulna." figureDoi="http://doi.org/10.5281/zenodo.7390396" httpUri="https://zenodo.org/record/7390396/files/figure.png" pageId="5" pageNumber="6">Figs. 1a, b</figureCitation>
|
||
,
|
||
<figureCitation box="[505,555,470,493]" captionStart="Fig" captionStartId="2.[116,146,1093,1112]" captionTargetBox="[155,1432,136,1068]" captionTargetId="figure-418@2.[154,1434,135,1069]" captionTargetPageId="2" captionText="Fig. 2 Skull of Natovenator polydontus (MPC-D 102/114, holotype). a–d Skull in left lateral (a), right lateral (b), dorsal (c), and ventral (d) views. e µCTrendered image sliced at the point marked on a, showing a cross-section of the premaxillary and anterior maxillary teeth in dorsal view. f Micro-computed tomography (µCT) rendered image of the occipital region in posterior view. g µCT-rendered image of the pterygoid and quadrate.?bm possible bite mark, d dentary, f frontal, h humerus, l lacrimal, m5 5th maxillary tooth, mx maxilla, na nasal p parietal, p13 13th premaxillary tooth, pl palatine, pm premaxilla, pop paroccipital process, pt pterygoid, q quadrate, rt replacement tooth, sq squamosal, so supraoccipital." figureDoi="http://doi.org/10.5281/zenodo.7390398" httpUri="https://zenodo.org/record/7390398/files/figure.png" pageId="5" pageNumber="6">2a–d</figureCitation>
|
||
,
|
||
<figureCitation box="[576,633,471,493]" captionStart="Fig" captionStartId="3.[116,146,1298,1317]" captionTargetBox="[155,1432,136,1271]" captionTargetId="figure-371@3.[154,1434,135,1273]" captionTargetPageId="3" captionText="Fig. 3 Postcranial elements and phylogenetic position of Natovenator polydontus (MPC-D 102/114, holotype). a Anterior cervical vertebrae in left lateral view. b Axis and third cervical vertebra in dorsal view. c Fourth cervical vertebra in dorsal view.d Posterior cervical vertebrae in right lateral view. e Dorsal series in right lateral view. f Anterior caudal vertebrae in right lateral view. g Left forearm elements in medial view and manus in ventral view. h Right foot in ventral view. i Phylogenetic position of Natovenator in Dromaeosauridae. Numbers at each node indicate Bremer support values. at atlas, c3 third cervical vertebra, c4 fourth cervical vertebra, c7 seventh cervical vertebra, c9 ninth cervical vertebra, ch chevron, d7 seventh dorsal vertebra, fem femur, mc I metacarpal I, mt III metatarsal III, mt IV metatarsal IV, poz postzygapophysis, prz prezygapophysis, r radius, r7 seventh dorsal rib, ul ulna, I-2 pedal phalanx I-2." figureDoi="http://doi.org/10.5281/zenodo.7390404" httpUri="https://zenodo.org/record/7390404/files/figure.png" pageId="5" pageNumber="6">3e, g</figureCitation>
|
||
). The nearly complete right humerus is proportionately short and distally flattened like that of
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[348,490,527,548]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[348,490,527,548]" italics="true" pageId="5" pageNumber="6">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Cau, A." box="[490,500,522,541]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
. The shaft of the ulna is mediolaterally compressed to produce a sharp posterior margin as in
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[178,320,583,604]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[178,320,583,604]" italics="true" pageId="5" pageNumber="6">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Cau, A." box="[321,331,578,597]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
and
|
||
<taxonomicName authorityName="Turner, Pol, Clarke, Erickson & Norell" authorityYear="2007" box="[390,492,583,604]" family="Dromaeosauridae" genus="Mahakala" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[390,492,583,604]" italics="true" pageId="5" pageNumber="6">Mahakala</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Turner, A. H. & Pol, D. & Norell, M. A." box="[492,511,578,597]" journalOrPublisher="Am. Mus. Novit." pageId="5" pageNumber="6" pagination="1 - 66" part="3722" refId="ref5937" refString="17. Turner, A. H., Pol, D. & Norell, M. A. Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia. Am. Mus. Novit. 3722, 1 - 66 (2011)." title="Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia" type="journal article" year="2011">17</bibRefCitation>
|
||
. Metacarpal III is robust and is only slightly longer than metacarpal II. Similarly, metacarpal III is almost as thick and long as other second metacarpals of other halszkaraptorines
|
||
<bibRefCitation author="Cau, A." box="[510,520,661,680]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Turner, A. H. & Pol, D. & Norell, M. A." box="[524,543,661,680]" journalOrPublisher="Am. Mus. Novit." pageId="5" pageNumber="6" pagination="1 - 66" part="3722" refId="ref5937" refString="17. Turner, A. H., Pol, D. & Norell, M. A. Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia. Am. Mus. Novit. 3722, 1 - 66 (2011)." title="Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia" type="journal article" year="2011">17</bibRefCitation>
|
||
. The femur has a long ridge on its posterior surface, which is another characteristic shared among halszkaraptorines
|
||
<bibRefCitation author="Cau, A." box="[443,453,717,736]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
. Typically for a dromaeosaurid, metatarsals II and III have ginglymoid distal articular surfaces (
|
||
<figureCitation box="[123,198,778,800]" captionStart="Fig" captionStartId="3.[116,146,1298,1317]" captionTargetBox="[155,1432,136,1271]" captionTargetId="figure-371@3.[154,1434,135,1273]" captionTargetPageId="3" captionText="Fig. 3 Postcranial elements and phylogenetic position of Natovenator polydontus (MPC-D 102/114, holotype). a Anterior cervical vertebrae in left lateral view. b Axis and third cervical vertebra in dorsal view. c Fourth cervical vertebra in dorsal view.d Posterior cervical vertebrae in right lateral view. e Dorsal series in right lateral view. f Anterior caudal vertebrae in right lateral view. g Left forearm elements in medial view and manus in ventral view. h Right foot in ventral view. i Phylogenetic position of Natovenator in Dromaeosauridae. Numbers at each node indicate Bremer support values. at atlas, c3 third cervical vertebra, c4 fourth cervical vertebra, c7 seventh cervical vertebra, c9 ninth cervical vertebra, ch chevron, d7 seventh dorsal vertebra, fem femur, mc I metacarpal I, mt III metatarsal III, mt IV metatarsal IV, poz postzygapophysis, prz prezygapophysis, r radius, r7 seventh dorsal rib, ul ulna, I-2 pedal phalanx I-2." figureDoi="http://doi.org/10.5281/zenodo.7390404" httpUri="https://zenodo.org/record/7390404/files/figure.png" pageId="5" pageNumber="6">Fig. 3h</figureCitation>
|
||
and Supplementary
|
||
<figureCitation box="[421,522,778,800]" captionStart="Fig" captionStartId="4.[116,146,1473,1492]" captionTargetBox="[155,1433,141,1448]" captionTargetId="figure-263@4.[154,1434,135,1449]" captionTargetPageId="4" captionText="Fig. 4 Body plan of Natovenator polydontus (MPC-D 102/114, holotype) and dorsal rib morphology of various diving birds and terrestrial taxa. a Dorsal series of Natovenator in ventral view. b Reconstruction of dorsal vertebrae and ribs of Natovenator in left lateral view. c Skeletal reconstruction of Natovenator with missing parts in dark grey. d–i Dorsal rib morphology of Natovenator (d), diving birds (e–i), common ostrich (j), and Shri devi, a likely terrestrial dromaeosaurid from the Baruungoyot Formation (k) in ventral view (not to scale). l Reconstruction of the fourth dorsal vertebra with corresponding ribs in anterior view. d2 second dorsal vertebra, r2 second dorsal rib, r3 third dorsal rib, r4 fourth dorsal rib." figureDoi="http://doi.org/10.5281/zenodo.7390406" httpUri="https://zenodo.org/record/7390406/files/figure.png" pageId="5" pageNumber="6">Fig. 4f, h</figureCitation>
|
||
). The ventral surface of metatarsal III is invaded by a ridge near the distal end, unlike other halszkaraptorines (
|
||
<figureCitation box="[370,443,834,856]" captionStart="Fig" captionStartId="3.[116,146,1298,1317]" captionTargetBox="[155,1432,136,1271]" captionTargetId="figure-371@3.[154,1434,135,1273]" captionTargetPageId="3" captionText="Fig. 3 Postcranial elements and phylogenetic position of Natovenator polydontus (MPC-D 102/114, holotype). a Anterior cervical vertebrae in left lateral view. b Axis and third cervical vertebra in dorsal view. c Fourth cervical vertebra in dorsal view.d Posterior cervical vertebrae in right lateral view. e Dorsal series in right lateral view. f Anterior caudal vertebrae in right lateral view. g Left forearm elements in medial view and manus in ventral view. h Right foot in ventral view. i Phylogenetic position of Natovenator in Dromaeosauridae. Numbers at each node indicate Bremer support values. at atlas, c3 third cervical vertebra, c4 fourth cervical vertebra, c7 seventh cervical vertebra, c9 ninth cervical vertebra, ch chevron, d7 seventh dorsal vertebra, fem femur, mc I metacarpal I, mt III metatarsal III, mt IV metatarsal IV, poz postzygapophysis, prz prezygapophysis, r radius, r7 seventh dorsal rib, ul ulna, I-2 pedal phalanx I-2." figureDoi="http://doi.org/10.5281/zenodo.7390404" httpUri="https://zenodo.org/record/7390404/files/figure.png" pageId="5" pageNumber="6">Fig. 3h</figureCitation>
|
||
)
|
||
<bibRefCitation author="Cau, A." box="[451,461,829,848]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Osmolska, H." box="[465,475,829,848]" journalOrPublisher="Neues Jahrb. Geol. P. M." pageId="5" pageNumber="6" pagination="440 - 448" part="1982" refId="ref5437" refString="5. Osmolska, H. Hulsanpes perlei n. g. n. sp. (Deinonychosauria, Saurischia, Dinosauria) from the upper Cretaceous Barun Goyot Formation of Mongolia. Neues Jahrb. Geol. P. M. 1982, 440 - 448 (1982)." title="Hulsanpes perlei n. g. n. sp. (Deinonychosauria, Saurischia, Dinosauria) from the upper Cretaceous Barun Goyot Formation of Mongolia" type="journal article" year="1982">5</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Turner, A. H. & Pol, D. & Norell, M. A." box="[479,498,829,848]" journalOrPublisher="Am. Mus. Novit." pageId="5" pageNumber="6" pagination="1 - 66" part="3722" refId="ref5937" refString="17. Turner, A. H., Pol, D. & Norell, M. A. Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia. Am. Mus. Novit. 3722, 1 - 66 (2011)." title="Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Togrogiin Shiree, Mongolia" type="journal article" year="2011">17</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Cau, A. & Madzia, D." box="[503,522,829,848]" journalOrPublisher="PeerJ" pageId="5" pageNumber="6" pagination="4868" part="6" refId="ref6243" refString="24. Cau, A. & Madzia, D. Redescription and affinities of Hulsanpes perlei (Dinosauria, Theropoda) from the Upper Cretaceous of Mongolia. PeerJ 6, e 4868 (2018)." title="Redescription and affinities of Hulsanpes perlei (Dinosauria, Theropoda) from the Upper Cretaceous of Mongolia" type="journal article" year="2018">24</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<footnote box="[424,1123,2007,2023]" pageId="4" pageNumber="5">
|
||
<paragraph blockId="4.[424,1123,2007,2023]" box="[424,1123,2007,2023]" pageId="4" pageNumber="5">(2022) 5:1185 | https://doi.org/10.1038/s42003-022-04119-9 | www.nature.com/commsbio</paragraph>
|
||
</footnote>
|
||
<subSubSection pageId="5" pageNumber="6" type="discussion">
|
||
<paragraph blockId="5.[116,778,890,1470]" lastBlockId="5.[810,1472,136,354]" pageId="5" pageNumber="6">
|
||
<emphasis box="[116,326,890,911]" italics="true" pageId="5" pageNumber="6">Phylogenetic analysis</emphasis>
|
||
. The phylogenetic analysis found more than 99,999 most parsimonious trees (CI = 0.23, RI = 0.55) with 6574 steps. Deinonychosaurian monophyly is not supported by the strict consensus tree (Supplementary Fig. 6). Instead,
|
||
<taxonomicName authorityName="Matthew and Brown" authorityYear="1922" family="Dromaeosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Dromaeosauridae</taxonomicName>
|
||
was recovered as a sister clade to a monophyletic clade formed by
|
||
<taxonomicName box="[295,433,1029,1051]" family="Troodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Troodontidae</taxonomicName>
|
||
and Avialae, which is consistent with the results of Cau et al.
|
||
<bibRefCitation author="Cau, A." box="[428,438,1052,1071]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
and Cau
|
||
<bibRefCitation author="Cau, A." box="[540,550,1052,1071]" journalOrPublisher="PeerJ" pageId="5" pageNumber="6" pagination="8672" part="8" refId="ref5542" refString="7. Cau, A. The body plan of Halszkaraptor escuilliei (Dinosauria, Theropoda) is not a transitional form along the evolution of dromaeosaurid hypercarnivory. PeerJ 8, e 8672 (2020)." title="The body plan of Halszkaraptor escuilliei (Dinosauria, Theropoda) is not a transitional form along the evolution of dromaeosaurid hypercarnivory" type="journal article" year="2020">7</bibRefCitation>
|
||
. Halszkaraptorinae is positioned at the base of
|
||
<taxonomicName authorityName="Matthew and Brown" authorityYear="1922" box="[399,578,1085,1107]" family="Dromaeosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Dromaeosauridae</taxonomicName>
|
||
as in Cau et al.
|
||
<bibRefCitation author="Cau, A." box="[763,773,1080,1099]" journalOrPublisher="Nature" pageId="5" pageNumber="6" pagination="395 - 399" part="552" refId="ref5405" refString="4. Cau, A. et al. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature 552, 395 - 399 (2017)." title="Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs" type="journal article" year="2017">4</bibRefCitation>
|
||
, although there are claims that dromaeosaurid affinities of halszkaraptorines are not well supported
|
||
<bibRefCitation author="Agnolin, F. L. & Motta, M. J. & Brisson Egli, F. & Lo Coco, G. & Novas, F. E." box="[495,514,1136,1155]" journalOrPublisher="Front. Earth Sci." pageId="5" pageNumber="6" pagination="1 - 28" part="6" refId="ref6281" refString="25. Agnolin, F. L., Motta, M. J., Brisson Egli, F., Lo Coco, G. & Novas, F. E. Paravian phylogeny and the dinosaur-bird transition: an overview. Front. Earth Sci. 6, 1 - 28 (2019)." title="Paravian phylogeny and the dinosaur-bird transition: an overview" type="journal article" year="2019">25</bibRefCitation>
|
||
. Nine (seven ambiguous and two unambiguous) synapomorphies support the inclusion of Halszkaraptorinae in
|
||
<taxonomicName authorityName="Matthew and Brown" authorityYear="1922" box="[344,525,1196,1218]" family="Dromaeosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="family">Dromaeosauridae</taxonomicName>
|
||
. The two unambiguous synapomorphies are the anterior tympanic recess at the same level as the basipterygoid process and the presence of a ventral flange on the paroccipital process. A total of 20 synapomorphies (including one unambiguous synapomorphy) unite the four halszkaraptorines, including
|
||
<taxonomicName box="[408,533,1337,1358]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[408,533,1337,1358]" italics="true" pageId="5" pageNumber="6">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
(Supplementary Fig. 7). In Halszkaraptorinae,
|
||
<taxonomicName authorityName="Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie & Godefroit" authorityYear="2017" box="[357,499,1364,1385]" class="Reptilia" family="Dromaeosauridae" genus="Halszkaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[357,499,1364,1385]" italics="true" pageId="5" pageNumber="6">Halszkaraptor</emphasis>
|
||
</taxonomicName>
|
||
is the earliest branching taxon, and the remaining three taxa form an unresolved clade supported by three ambiguous synapomorphies (characters 121/ 1, 569/0, and 1153/1). Two of these synapomorphies are related to the paroccipital process (characters 121 and 569), which is not preserved in
|
||
<taxonomicName authorityName="Osmolska" authorityYear="1982" box="[956,1059,164,185]" class="Reptilia" family="Dromaeosauridae" genus="Hulsanpes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[956,1059,164,185]" italics="true" pageId="5" pageNumber="6">Hulsanpes</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Osmolska, H." box="[1059,1069,159,178]" journalOrPublisher="Neues Jahrb. Geol. P. M." pageId="5" pageNumber="6" pagination="440 - 448" part="1982" refId="ref5437" refString="5. Osmolska, H. Hulsanpes perlei n. g. n. sp. (Deinonychosauria, Saurischia, Dinosauria) from the upper Cretaceous Barun Goyot Formation of Mongolia. Neues Jahrb. Geol. P. M. 1982, 440 - 448 (1982)." title="Hulsanpes perlei n. g. n. sp. (Deinonychosauria, Saurischia, Dinosauria) from the upper Cretaceous Barun Goyot Formation of Mongolia" type="journal article" year="1982">5</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Cau, A. & Madzia, D." box="[1073,1092,159,178]" journalOrPublisher="PeerJ" pageId="5" pageNumber="6" pagination="4868" part="6" refId="ref6243" refString="24. Cau, A. & Madzia, D. Redescription and affinities of Hulsanpes perlei (Dinosauria, Theropoda) from the Upper Cretaceous of Mongolia. PeerJ 6, e 4868 (2018)." title="Redescription and affinities of Hulsanpes perlei (Dinosauria, Theropoda) from the Upper Cretaceous of Mongolia" type="journal article" year="2018">24</bibRefCitation>
|
||
. The other is the presence of an expansion on the medial margin of the distal half of metatarsal III, which is not entirely preserved in the
|
||
<taxonomicName box="[1267,1392,221,242]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1267,1392,221,242]" italics="true" pageId="5" pageNumber="6">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
. When scored as 0 for this character,
|
||
<taxonomicName box="[1117,1242,249,270]" family="Dromaeosauridae" genus="Natovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1117,1242,249,270]" italics="true" pageId="5" pageNumber="6">Natovenator</emphasis>
|
||
</taxonomicName>
|
||
branches off from the unresolved clade. It suggests that the medial expansion of the dorsal surface of metatarsal III could be a derived character among halszkaraptorines.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |