468 lines
84 KiB
XML
468 lines
84 KiB
XML
<document id="4EA57F8A91F0AA455E6D2DE8CF2BC818" ID-CLB-Dataset="3804" ID-DOI="10.1206/0003-0082(2002)364<0001:CAOCOT>2.0.CO;2" ID-GBIF-Dataset="b8965a84-547b-4855-ac3d-2a9286b327d5" ID-ISSN="0003-0082" ID-Zenodo-Dep="4734832" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1620067791992" checkinUser="felipe" docAuthor="CLARK, JAMES M., NORELL, MARK A. & ROWE, TIMOTHY" docDate="2002" docId="64443907284A7E67FD5E9DBCFCA7AC13" docLanguage="en" docName="AmMusNovit.2002.3364.1-24.pdf" docOrigin="American Museum Novitates 3364" docSource="http://www.bioone.org/doi/abs/10.1206/0003-0082%282002%29364%3C0001%3ACAOCOT%3E2.0.CO%3B2" docStyle="DocumentStyle:70DDF1BDF6846024993F05262BAE3001.3:AmMusNovit.2000-2010.journal_article.type1" docStyleId="70DDF1BDF6846024993F05262BAE3001" docStyleName="AmMusNovit.2000-2010.journal_article.type1" docStyleVersion="3" docTitle="Citipati osmolskae Clark 2001" docType="treatment" docVersion="6" lastPageNumber="14" masterDocId="987D417F28487E6AFFB79F22FF97A906" masterDocTitle="Cranial Anatomy of Citipati osmolskae (Theropoda, Oviraptorosauria), and a Reinterpretation of the Holotype of Oviraptor philoceratops" masterLastPageNumber="24" masterPageNumber="1" pageNumber="3" updateTime="1698938106598" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:title id="31D7329C9FC204779B90A8C61DAD0AAC">Cranial Anatomy of Citipati osmolskae (Theropoda, Oviraptorosauria), and a Reinterpretation of the Holotype of Oviraptor philoceratops</mods:title>
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<mods:namePart id="A03910980BED27E7A25CA86B4E64FDA3">CLARK, JAMES M.</mods:namePart>
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<mods:namePart id="B65FB2F16F76BDE02609FA4FCE56B4C0">NORELL, MARK A.</mods:namePart>
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<mods:namePart id="78F83183810AA7D973DE1410F8AE5AFA">ROWE, TIMOTHY</mods:namePart>
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<mods:title id="82D7B51B7E07C23C78213D19E1B7B642">American Museum Novitates</mods:title>
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<mods:date id="F0A6C2A29E09C830B58674998931877E">2002</mods:date>
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<treatment id="64443907284A7E67FD5E9DBCFCA7AC13" ID-DOI="http://doi.org/10.5281/zenodo.5604437" ID-GBIF-Taxon="189534864" ID-Zenodo-Dep="5604437" LSID="urn:lsid:plazi:treatment:64443907284A7E67FD5E9DBCFCA7AC13" httpUri="http://treatment.plazi.org/id/64443907284A7E67FD5E9DBCFCA7AC13" lastPageId="13" lastPageNumber="14" pageId="2" pageNumber="3">
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<subSubSection id="A4F7DB9A284A7E68FD5E9DBCFB08ABB3" box="[745,1183,670,693]" pageId="2" pageNumber="3" type="nomenclature">
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<paragraph id="EC528811284A7E68FD5E9DBCFB08ABB3" blockId="2.[745,1183,670,693]" box="[745,1183,670,693]" pageId="2" pageNumber="3">
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<heading id="B71A3F7D284A7E68FD5E9DBCFB08ABB3" box="[745,1183,670,693]" centered="true" fontSize="36" level="2" pageId="2" pageNumber="3" reason="8">
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<taxonomicName id="2BEDF392284A7E68FD5E9DBCFB08ABB3" authority="Clark et al., 2001" authorityName="Clark" authorityYear="2001" box="[745,1183,670,693]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="osmolskae">
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<emphasis id="DE995403284A7E68FD5E9DBCFC52ABB3" box="[745,965,670,693]" italics="true" pageId="2" pageNumber="3">Citipati osmolskae</emphasis>
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<bibRefCitation id="887CF5E0284A7E68FC799DBCFB08ABB3" author="Clark, J. M. & M. A. Norell & R. Barsbold" box="[974,1183,670,693]" pageId="2" pageNumber="3" pagination="209 - 213" refId="ref11162" refString="Clark, J. M., M. A. Norell, and R. Barsbold. 2001. Two new oviraptorids (Theropoda: Oviraptorosauria) Late Cretaceous Djadoktha Formation, Ukhaa Tolgod, Mongolia. Journal of Vertebrate Paleontology 21: 209 - 213." type="journal article" year="2001">Clark et al., 2001</bibRefCitation>
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</taxonomicName>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="A4F7DB9A284A7E68FD6E9DEDFBCEAD0D" pageId="2" pageNumber="3" type="materials_examined">
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<paragraph id="EC528811284A7E68FD6E9DEDFCF0AA05" blockId="2.[700,1229,719,1742]" pageId="2" pageNumber="3">
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<materialsCitation id="5C85824C284A7E68FD6E9DEDFCF0AA05" ID-GBIF-Occurrence="3394411302" collectionCode="IGM" pageId="2" pageNumber="3" specimenCode="IGM 100" specimenCount="1" typeStatus="holotype">
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<typeStatus id="335636B3284A7E68FD6E9DEDFCD8ABE3" box="[729,847,719,742]" pageId="2" pageNumber="3" type="holotype">HOLOTYPE</typeStatus>
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:
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<specimenCode id="BC4B206A284A7E68FCD29DEDFC4FABE0" box="[869,984,719,742]" collectionCode="IGM" country="Mongolia" httpUri="http://grbio.org/cool/0h5a-qjh4" name="Geological Institute, Mongolian Academy of Sciences" pageId="2" pageNumber="3">IGM 100</specimenCode>
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/978, a nearly complete skeleton.
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</materialsCitation>
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</paragraph>
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<paragraph id="EC528811284A7E68FD6E9C28FB96AA90" blockId="2.[700,1229,719,1742]" pageId="2" pageNumber="3">
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<collectionCode id="8AFC10D4284A7E68FD6E9C28FC05AA26" box="[729,914,778,801]" pageId="2" pageNumber="3">TYPE LOCALITY</collectionCode>
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: Djadokhta Formation at Ankylosaur Flats, Ukhaa Tolgod, Gurvan Tes Somon,
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<collectingRegion id="2E2946F3284A7E68FCA99C66FC7AAA5D" box="[798,1005,836,859]" country="Mongolia" name="Omnogovi" pageId="2" pageNumber="3">Omnogov Aimak</collectingRegion>
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,
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<collectingCountry id="94FAC881284A7E68FC4E9C66FBF9AA5D" box="[1017,1134,836,859]" name="Mongolia" pageId="2" pageNumber="3">Mongolia</collectingCountry>
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. Precise coordinates will be made available to qualified researchers on request.
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</paragraph>
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<paragraph id="EC528811284A7E68FD6E9CBEFBCEAD0D" blockId="2.[700,1229,719,1742]" pageId="2" pageNumber="3">
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REFERRED SPECIMENS:
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<collectionCode id="8AFC10D4284A7E68FC6A9CBEFB81AAB5" box="[989,1046,924,947]" country="Mongolia" httpUri="http://grbio.org/cool/0h5a-qjh4" name="Geological Institute, Mongolian Academy of Sciences" pageId="2" pageNumber="3">IGM</collectionCode>
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100/979, a partial skeleton overlying a nest, from Ankylosaur Flats, Ukhaa Tolgod;
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<collectionCode id="8AFC10D4284A7E68FC4A9CF5FBA1AAE8" box="[1021,1078,983,1006]" country="Mongolia" httpUri="http://grbio.org/cool/0h5a-qjh4" name="Geological Institute, Mongolian Academy of Sciences" pageId="2" pageNumber="3">IGM</collectionCode>
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100/971, an embryonic skeleton within an egg.
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</paragraph>
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</subSubSection>
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<subSubSection id="A4F7DB9A284A7E68FD6E9B30FB5AAF94" pageId="2" pageNumber="3" type="diagnosis">
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<paragraph id="EC528811284A7E68FD6E9B30FB5AAF94" blockId="2.[700,1229,719,1742]" pageId="2" pageNumber="3">
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DIAGNOSIS: Oviraptorid differing from
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<taxonomicName id="2BEDF392284A7E68FD0B9B0DFCF6AD40" authorityName="Barsbold" authorityYear="1986" box="[700,865,1071,1094]" class="Reptilia" family="Oviraptoridae" genus="Conchoraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
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<emphasis id="DE995403284A7E68FD0B9B0DFCF6AD40" box="[700,865,1071,1094]" italics="true" pageId="2" pageNumber="3">Conchoraptor</emphasis>
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</taxonomicName>
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in having a taller and more highly pneumatized nasal, from
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<taxonomicName id="2BEDF392284A7E68FBF89B6EFB3FAD65" authorityName="Barsbold" authorityYear="1981" box="[1103,1192,1100,1123]" class="Adenophorea" family="Tripyloididae" genus="Ingenia" kingdom="Animalia" order="Araeolaimida" pageId="2" pageNumber="3" phylum="Nematoda" rank="genus">
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<emphasis id="DE995403284A7E68FBF89B6EFB3FAD65" box="[1103,1192,1100,1123]" italics="true" pageId="2" pageNumber="3">Ingenia</emphasis>
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</taxonomicName>
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in that metacarpal I is not extremely broad, from
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<taxonomicName id="2BEDF392284A7E68FCB29BA5FBBDAD98" authorityName="Osborn" authorityYear="1924" box="[773,1066,1159,1182]" class="Reptilia" family="Oviraptoridae" genus="Oviraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="philoceratops">
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<emphasis id="DE995403284A7E68FCB29BA5FBBDAD98" box="[773,1066,1159,1182]" italics="true" pageId="2" pageNumber="3">Oviraptor philoceratops</emphasis>
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</taxonomicName>
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in having a shorter skull and mandible, from
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<taxonomicName id="2BEDF392284A7E68FBFA9B86FC91ADDF" baseAuthorityName="Barsbold" baseAuthorityYear="1986" class="Reptilia" family="Oviraptoridae" genus="Oviraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="mongoliensis">
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<emphasis id="DE995403284A7E68FBFA9B86FC91ADDF" italics="true" pageId="2" pageNumber="3">O. mongoliensis</emphasis>
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</taxonomicName>
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in lacking a convex crest on the frontals and parietals, and from
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<taxonomicName id="2BEDF392284A7E68FBB09BFDFB5BADF0" box="[1031,1228,1247,1270]" class="Reptilia" family="Oviraptoridae" genus="Khaan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="mckennai">
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<emphasis id="DE995403284A7E68FBB09BFDFB5BADF0" box="[1031,1228,1247,1270]" italics="true" pageId="2" pageNumber="3">Khaan mckennai</emphasis>
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</taxonomicName>
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in the features listed below. Differs from all other oviraptorids where known in the anterodorsally sloping occiput and quadrate, the parietal being much longer along the midline than the frontal and reaching nearly to the level of the anterior end of the orbit, the ascending process of the jugal being perpendicular to the horizontal ramus (rather than extending posterodorsally), the narial opening being more nearly circular and the ascending process of the premaxilla being vertical rather than sloping posterodorsally, and the cervical vertebrae being elongate (approximately twice as long as they are wide).
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</paragraph>
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</subSubSection>
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<subSubSection id="A4F7DB9A284A7E69FD6E99BBFD8CAAD5" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3" type="discussion">
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<paragraph id="EC528811284A7E69FD6E99BBFD8CAAD5" blockId="2.[700,1229,719,1742]" lastBlockId="3.[123,652,193,1741]" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3">
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REMARKS: This specimen was identified preliminarily as
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<taxonomicName id="2BEDF392284A7E68FCC99995FBD4AFC8" authorityName="Osborn" authorityYear="1924" box="[894,1091,1719,1742]" class="Reptilia" family="Oviraptoridae" genus="Oviraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="philoceratops">
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<emphasis id="DE995403284A7E68FCC99995FBD4AFC8" box="[894,1091,1719,1742]" italics="true" pageId="2" pageNumber="3">O. philoceratops</emphasis>
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</taxonomicName>
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(e.g., Webster, 1996), but comparison with the
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<typeStatus id="335636B3284B7E69FD929FE3FD1CA9DE" box="[549,651,193,216]" pageId="3" pageNumber="4" type="holotype">holotype</typeStatus>
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of the latter revealed several significant differences (see below). A postcranial skeleton that is preserved on top of a nest, IGM 100/ 979, is referred to the new species and was described by
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<bibRefCitation id="887CF5E0284B7E69FEA89E76FD93A86D" author="Clark, J. M. & M. Norell & L. Chiappe" box="[287,516,340,363]" pageId="3" pageNumber="4" pagination="1 - 36" refId="ref11210" refString="Clark, J. M., M. Norell, and L. Chiappe. 1999. An oviraptorid skeleton from the Late Cretaceous of Ukhaa Tolgod, Mongolia, preserved in an avian-like brooding position over an oviraptorid nest. American Museum Novitates 3265: 1 - 36." type="journal article" year="1999">Clark et al. (1999)</bibRefCitation>
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. IGM 100/ 979 is referred to
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<taxonomicName id="2BEDF392284B7E69FEC69E53FD82A88E" box="[369,533,369,392]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="osmolskae">
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<emphasis id="DE995403284B7E69FEC69E53FD82A88E" box="[369,533,369,392]" italics="true" pageId="3" pageNumber="4">C. osmolskae</emphasis>
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</taxonomicName>
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, pending preparation of the postcranial skeleton of the
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<typeStatus id="335636B3284B7E69FFCC9E8EFF73A8C5" box="[123,228,428,451]" pageId="3" pageNumber="4" type="holotype">holotype</typeStatus>
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, primarily on the basis of its large size and its differences from the skeleton of
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<taxonomicName id="2BEDF392284B7E69FFCC9EC5FED7A8F8" box="[123,320,487,510]" class="Reptilia" family="Oviraptoridae" genus="Khaan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="mckennai">
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<emphasis id="DE995403284B7E69FFCC9EC5FED7A8F8" box="[123,320,487,510]" italics="true" pageId="3" pageNumber="4">Khaan mckennai</emphasis>
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</taxonomicName>
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. An embryonic skeleton referred to this species was given a preliminary description by
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<bibRefCitation id="887CF5E0284B7E69FE8D9D03FDA4AB3E" author="Norell, M. & A, J. M. & Clark, D. & Dashzeveg, R. & Barsbold, L. M. & Chiappe, A. R. & Davidson, M. C. & M. J. Novacek" box="[314,563,545,568]" pageId="3" pageNumber="4" pagination="779 - 782" refId="ref12515" refString="Norell, M. A, J. M. Clark, D. Dashzeveg, R. Barsbold, L. M. Chiappe, A. R. Davidson, M. C. McKenna, and M. J. Novacek. 1994. A theropod dinosaur embryo, and the affinities of the Flaming Cliffs dinosaur eggs. Science 266: 779 - 782." type="journal article" year="1994">Norell et al. (1994)</bibRefCitation>
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and a more detailed description by Norell et al. (2001). This specimen is referred to
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<taxonomicName id="2BEDF392284B7E69FD859D7EFD1CAB75" box="[562,651,604,627]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="DE995403284B7E69FD859D7EFD1CAB75" box="[562,651,604,627]" italics="true" pageId="3" pageNumber="4">Citipati</emphasis>
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</taxonomicName>
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on the basis of its premaxilla, which is nearly vertical rather than sloping posteriorly as in
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<taxonomicName id="2BEDF392284B7E69FFCC9D96FF5FABCD" box="[123,200,692,715]" class="Reptilia" family="Oviraptoridae" genus="Khaan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="DE995403284B7E69FFCC9D96FF5FABCD" box="[123,200,692,715]" italics="true" pageId="3" pageNumber="4">Khaan</emphasis>
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</taxonomicName>
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. An oviraptorid collected from the Djadokhta Formation at Dzamin Khong, IGM 100/42, was previously identified as
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<taxonomicName id="2BEDF392284B7E69FDD89DCDFE8CAA25" authorityName="Osborn" authorityYear="1924" class="Reptilia" family="Oviraptoridae" genus="Oviraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="philoceratops">
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<emphasis id="DE995403284B7E69FDD89DCDFE8CAA25" italics="true" pageId="3" pageNumber="4">O. philoceratops</emphasis>
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</taxonomicName>
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(
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<bibRefCitation id="887CF5E0284B7E69FE879C2EFE7AAA25" author="Barsbold, R." box="[304,493,780,803]" pageId="3" pageNumber="4" pagination="28 - 39" refId="ref10938" refString="Barsbold, R. 1981. Toothless dinosaurs of Mon- golia. Joint Soviet - Mongolian Paleontological Expedition Transactions 15: 28 - 39. [in Russian]" type="journal article" year="1981">Barsbold, 1981</bibRefCitation>
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||
;
|
||
<bibRefCitation id="887CF5E0284B7E69FDB79C2EFF70AA46" author="Barsbold, R. & T. Maryanska & H. Osmolska" pageId="3" pageNumber="4" pagination="249 - 258" refId="ref11058" refString="Barsbold, R., T. Maryanska, and H. Osmolska. 1990. Oviraptorosauria. In D. B. Weishampel, P. Dodson, and H. Osmolska (editors), The Dinosauria: 249 - 258. Berkeley: University of California Press." type="book chapter" year="1990">Barsbold et al., 1990</bibRefCitation>
|
||
,
|
||
<figureCitation id="74D69494284B7E69FF409C0BFE5EAA46" box="[247,457,809,832]" captionStart="Fig" captionStartId="13.[149,187,827,848]" captionTargetBox="[123,1228,185,805]" captionTargetId="figure-13@13.[123,1228,185,805]" captionTargetPageId="13" captionText="Fig. 10. Ventral view of the holotype cranium of Citipati osmolskae IGM 100/978 showing a left hyoid element in natural articulation. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734868" httpUri="https://zenodo.org/record/4734868/files/figure.png" pageId="3" pageNumber="4">fig. 10.1A and B</figureCitation>
|
||
), but it is more similar to
|
||
<taxonomicName id="2BEDF392284B7E69FF489C65FECFAA58" box="[255,344,839,862]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="DE995403284B7E69FF489C65FECFAA58" box="[255,344,839,862]" italics="true" pageId="3" pageNumber="4">Citipati</emphasis>
|
||
</taxonomicName>
|
||
in the shape of its premaxilla and circumnarial region and differs from
|
||
<taxonomicName id="2BEDF392284B7E69FF769CA3FE1DAA9E" authorityName="Osborn" authorityYear="1924" box="[193,394,897,920]" class="Reptilia" family="Oviraptoridae" genus="Oviraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="philoceratops">
|
||
<emphasis id="DE995403284B7E69FF769CA3FE1DAA9E" box="[193,394,897,920]" italics="true" pageId="3" pageNumber="4">O. philoceratops</emphasis>
|
||
</taxonomicName>
|
||
in the length of the maxilla and dentary (see below). It may represent a second species of
|
||
<taxonomicName id="2BEDF392284B7E69FE0C9C9EFD83AAD5" box="[443,532,956,979]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="DE995403284B7E69FE0C9C9EFD83AAD5" box="[443,532,956,979]" italics="true" pageId="3" pageNumber="4">Citipati</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="A4F7DB9A284B7E67FF2F9CFBFCA7AC13" lastPageId="13" lastPageNumber="14" pageId="3" pageNumber="4" type="description">
|
||
<paragraph id="EC528811284B7E69FF2F9CFBFD8EAF3D" blockId="3.[123,652,193,1741]" pageId="3" pageNumber="4">
|
||
DESCRIPTION: The skull of the
|
||
<typeStatus id="335636B3284B7E69FD929CFBFD1CAAF6" box="[549,651,985,1008]" pageId="3" pageNumber="4" type="holotype">holotype</typeStatus>
|
||
(
|
||
<figureCitation id="74D69494284B7E69FF349CD4FF6CAD0B" box="[131,251,1014,1037]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStart-4="Fig" captionStart-5="Fig" captionStart-6="Fig" captionStart-7="Fig" captionStart-8="Fig" captionStart-9="Fig" captionStartId-0="4.[150,188,1280,1301]" captionStartId-1="5.[149,187,936,957]" captionStartId-2="6.[150,188,1125,1146]" captionStartId-3="6.[726,764,920,941]" captionStartId-4="7.[149,187,1152,1173]" captionStartId-5="7.[725,763,755,776]" captionStartId-6="8.[150,188,1435,1456]" captionStartId-7="9.[149,187,1064,1085]" captionStartId-8="12.[150,188,1229,1250]" captionStartId-9="13.[149,187,827,848]" captionTargetBox-0="[124,1229,187,1259]" captionTargetBox-1="[123,1228,185,914]" captionTargetBox-2="[124,653,188,1105]" captionTargetBox-3="[702,1229,186,897]" captionTargetBox-4="[123,651,186,1130]" captionTargetBox-5="[699,1227,186,732]" captionTargetBox-6="[135,1219,189,1414]" captionTargetBox-7="[123,1227,187,1043]" captionTargetBox-8="[124,1230,186,1206]" captionTargetBox-9="[123,1228,185,805]" captionTargetId-0="figure-16@4.[124,1229,187,1259]" captionTargetId-1="figure-13@5.[123,1228,185,914]" captionTargetId-2="figure-18@6.[124,653,188,1105]" captionTargetId-3="figure-37@6.[702,1229,186,898]" captionTargetId-4="figure-15@7.[123,651,186,1130]" captionTargetId-5="figure-34@7.[699,1227,186,733]" captionTargetId-6="figure-16@8.[135,1219,189,1415]" captionTargetId-7="figure-13@9.[123,1227,187,1043]" captionTargetId-8="figure-16@12.[123,1230,186,1207]" captionTargetId-9="figure-13@13.[123,1228,185,805]" captionTargetPageId-0="4" captionTargetPageId-1="5" captionTargetPageId-2="6" captionTargetPageId-3="6" captionTargetPageId-4="7" captionTargetPageId-5="7" captionTargetPageId-6="8" captionTargetPageId-7="9" captionTargetPageId-8="12" captionTargetPageId-9="13" captionText-0="Fig. 1. The holotype skull and mandible of Citipati osmolskae (IGM 100/978) in right lateral view before final preparation. The mandible, scleral ossicles, and hyoid elements were later separated from the skull. Abbreviations in appendix 1." captionText-1="Fig. 2. Right lateral view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." captionText-2="Fig. 3. Palatal view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." captionText-3="Fig. 4. Anterior view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." captionText-4="Fig. 5. Dorsal view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." captionText-5="Fig. 6. Occipital view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." captionText-6="Fig. 7. CT scan transverse sections through the holotype skull of Citipati osmolskae IGM 100/978 at three levels showing a large, presumably pneumatized space dorsal to the endocranial cavity and its connection to the dorsal tympanic recess. Arrow indicates apparent connection medially from dorsal tympanic recess to dorsal space. Abbreviations in appendix 1." captionText-7="Fig. 8. CT scan parasagittal section through the right side of the holotype cranium of Citipati osmolskae IGM 100/978, showing the boundaries of the endocranial cavity and the large, presumably pneumatized space dorsal to it. Abbreviations in appendix 1." captionText-8="Fig. 9. (A) Lateral and (B) medial views of the left hemimandible of the holotype of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." captionText-9="Fig. 10. Ventral view of the holotype cranium of Citipati osmolskae IGM 100/978 showing a left hyoid element in natural articulation. Abbreviations in appendix 1." figureDoi-0="http://doi.org/10.5281/zenodo.4734838" figureDoi-1="http://doi.org/10.5281/zenodo.4734840" figureDoi-2="http://doi.org/10.5281/zenodo.4734848" figureDoi-3="http://doi.org/10.5281/zenodo.4734845" figureDoi-4="http://doi.org/10.5281/zenodo.4734856" figureDoi-5="http://doi.org/10.5281/zenodo.4734852" figureDoi-6="http://doi.org/10.5281/zenodo.4734858" figureDoi-7="http://doi.org/10.5281/zenodo.4734862" figureDoi-8="http://doi.org/10.5281/zenodo.4734866" figureDoi-9="http://doi.org/10.5281/zenodo.4734868" httpUri-0="https://zenodo.org/record/4734838/files/figure.png" httpUri-1="https://zenodo.org/record/4734840/files/figure.png" httpUri-2="https://zenodo.org/record/4734848/files/figure.png" httpUri-3="https://zenodo.org/record/4734845/files/figure.png" httpUri-4="https://zenodo.org/record/4734856/files/figure.png" httpUri-5="https://zenodo.org/record/4734852/files/figure.png" httpUri-6="https://zenodo.org/record/4734858/files/figure.png" httpUri-7="https://zenodo.org/record/4734862/files/figure.png" httpUri-8="https://zenodo.org/record/4734866/files/figure.png" httpUri-9="https://zenodo.org/record/4734868/files/figure.png" pageId="3" pageNumber="4">figs. 1–10</figureCitation>
|
||
) is complete, including both stapes and the paired elements of the hyoid apparatus. In lateral view (
|
||
<figureCitation id="74D69494284B7E69FE119B13FE7DAD4E" box="[422,490,1073,1096]" captionStart="Fig" captionStartId="5.[149,187,936,957]" captionTargetBox="[123,1228,185,914]" captionTargetId="figure-13@5.[123,1228,185,914]" captionTargetPageId="5" captionText="Fig. 2. Right lateral view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734840" httpUri="https://zenodo.org/record/4734840/files/figure.png" pageId="3" pageNumber="4">fig. 2</figureCitation>
|
||
), the skull is almost rectangular, with a vertical premaxilla anteriorly. The right and left premaxillae are fused and edentulous. Ventrally (
|
||
<figureCitation id="74D69494284B7E69FDBD9BABFDDAADA6" box="[522,589,1161,1184]" captionStart="Fig" captionStartId="6.[150,188,1125,1146]" captionTargetBox="[124,653,188,1105]" captionTargetId="figure-18@6.[124,653,188,1105]" captionTargetPageId="6" captionText="Fig. 3. Palatal view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734848" httpUri="https://zenodo.org/record/4734848/files/figure.png" pageId="3" pageNumber="4">fig. 3</figureCitation>
|
||
), the premaxilla expands transversely to form a curved, Ushaped triturating surface in ventral view that supports a series of parallel ridges and troughs. The oral margin of the premaxilla forms a sharp edge that bears a series of large bony denticles, five on each side. Posterior to these, the palatal surface of the premaxilla supports a pair of robust, longitudinal, parasagittal ridges that extend posteriorly along either side of the midline to the posterior edge of the palate. The palatal surface broadens toward the back of the mouth, where it bears two additional ridges that lie lateral and parallel to the first pair.
|
||
</paragraph>
|
||
<paragraph id="EC528811284B7E69FF2F9963FC54AB75" blockId="3.[123,652,193,1741]" lastBlockId="3.[699,1228,193,1741]" pageId="3" pageNumber="4">The large, elliptical nares are bordered ventrally, anteriorly, and anterodorsally by the premaxilla and posteriorly and posterodorsally by the nasal. Owing to profound foreshortening of the face, the naris lies almost entirely dorsal to the antorbital fossa, its rear margin lying nearly as far back as the posterior edge of the antorbital fenestra. The posteriorly fused nasals form a complex array of pockets along the posterodorsal edge of the narial opening. The pockets lie dorsal to a horizontal roof to the nares, and another, narrower horizontal roof overlies these pockets anterodorsally. A broad elliptical opening in the horizontal narial roof communicates between the nares and the dorsal nasal pockets. Dorsal to the naris the nasal forms a vertical lamina, and a lip along its dorsal edge was presumably for the attachment of circumnarial soft tissues.</paragraph>
|
||
<paragraph id="EC528811284B7E69FD6F9D5BFD6AAC35" blockId="3.[699,1228,193,1741]" pageId="3" pageNumber="4">
|
||
The large premaxilla forms much of the anterodorsal margin of the antorbital fossa. A fossa on the lateral surface of the ascending ramus anterior to the naris (
|
||
<figureCitation id="74D69494284B7E69FB9A9DF3FBFBABEE" box="[1069,1132,721,744]" captionStart="Fig" captionStartId="6.[726,764,920,941]" captionTargetBox="[702,1229,186,897]" captionTargetId="figure-37@6.[702,1229,186,898]" captionTargetPageId="6" captionText="Fig. 4. Anterior view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734845" httpUri="https://zenodo.org/record/4734845/files/figure.png" pageId="3" pageNumber="4">fig. 4</figureCitation>
|
||
) is similar to an accessory opening of IGM 100/42 described by
|
||
<bibRefCitation id="887CF5E0284B7E69FCE19C2EFBCFAA25" author="Barsbold, R. & T. Maryanska & H. Osmolska" box="[854,1112,780,803]" pageId="3" pageNumber="4" pagination="249 - 258" refId="ref11058" refString="Barsbold, R., T. Maryanska, and H. Osmolska. 1990. Oviraptorosauria. In D. B. Weishampel, P. Dodson, and H. Osmolska (editors), The Dinosauria: 249 - 258. Berkeley: University of California Press." type="book chapter" year="1990">Barsbold et al. (1990)</bibRefCitation>
|
||
. Posterior to the naris, the premaxilla extends dorsal to the maxilla to meet the nasal, separating the maxilla from the narial opening. The antorbital fossa is triangular with a vertical posterior edge and horizontal ventral edge. The tall, oval antorbital fenestra is separated from the large, triangular accessory antorbital fenestra by an hourglassshaped, inset interfenestral bar formed by the maxilla. Dorsal to the accessory fenestra are several smaller fenestrae near the dorsal margin of the fossa. The anteroventral margin of the fossa is composed of the maxilla, and a small splint of maxilla extends posterodorsally above the anterior edge of the fossa. Posteriorly, ventral to the antorbital fenestra, the fossa is bordered by the slender anterior process of the jugal.
|
||
</paragraph>
|
||
<paragraph id="EC528811284B7E69FD6F9A1BFBF1AFCB" blockId="3.[699,1228,193,1741]" pageId="3" pageNumber="4">
|
||
The lacrimal is transversely broad and forms a concave, posteriorly facing surface along the anterior edge of the orbit. It is perforated by a large lacrimal foramen. The lacrimal canal passed through this foramen, traversing a laterally open trough above the internal antorbital fenestra (
|
||
<bibRefCitation id="887CF5E0284B7E69FC589ACBFB03AF06" author="Witmer, L. M." box="[1007,1172,1513,1536]" pageId="3" pageNumber="4" pagination="1 - 73" refId="ref13170" refString="Witmer, L. M. 1997. The evolution of the antorbital cavity in archosaurs: a study in soft-tissue reconstruction in the fossil record with an analysis of the function of pneumaticity. Journal of Vertebrate Paleontology Memoir 3: 1 - 73." type="journal article" year="1997">Witmer, 1997</bibRefCitation>
|
||
) before penetrating the side of the face to enter the nasopharyngeal cavity just behind the posterodorsal extremity of the naris. The anterior part of the lacrimal dorsal to the antorbital fossa houses a broad pneumatic pocket opening anteriorly, within which lies several recessed pneumatic pockets.
|
||
</paragraph>
|
||
<caption id="B892D899284C7E6EFF219A22FD88AC4C" ID-DOI="http://doi.org/10.5281/zenodo.4734838" ID-Zenodo-Dep="4734838" httpUri="https://zenodo.org/record/4734838/files/figure.png" pageId="4" pageNumber="5" startId="4.[150,188,1280,1301]" targetBox="[124,1229,187,1259]" targetPageId="4">
|
||
<paragraph id="EC528811284C7E6EFF219A22FD88AC4C" blockId="4.[124,1229,1280,1354]" pageId="4" pageNumber="5">
|
||
Fig. 1. The holotype skull and mandible of
|
||
<taxonomicName id="2BEDF392284C7E6EFDC89A22FCD2AC13" box="[639,837,1280,1301]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE995403284C7E6EFDC89A22FCD2AC13" box="[639,837,1280,1301]" italics="true" pageId="4" pageNumber="5">Citipati osmolskae</emphasis>
|
||
</taxonomicName>
|
||
(IGM 100/978) in right lateral view before final preparation. The mandible, scleral ossicles, and hyoid elements were later separated from the skull. Abbreviations in appendix 1.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="EC528811284C7E6EFF2E9AB3FCAAAF5E" blockId="4.[124,653,1425,1741]" lastBlockId="4.[700,1229,1425,1741]" pageId="4" pageNumber="5">
|
||
There is no evidence of a prefrontal. The paired frontals contact the nasals in a simple suture (
|
||
<figureCitation id="74D69494284C7E6EFF639AEEFE82ACE5" box="[212,277,1484,1507]" captionStart="Fig" captionStartId="7.[149,187,1152,1173]" captionTargetBox="[123,651,186,1130]" captionTargetId="figure-15@7.[123,651,186,1130]" captionTargetPageId="7" captionText="Fig. 5. Dorsal view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734856" httpUri="https://zenodo.org/record/4734856/files/figure.png" pageId="4" pageNumber="5">fig. 5</figureCitation>
|
||
). The joint between the frontals is short but highly complex. Posteriorly, the fused parietals divide the frontals for over onehalf of their length. The frontal participates in the supratemporal fenestra, but it lacks the mandibular adductor fossa that deeply excavates the dorsal surface of the frontal in many other nonavian theropods. Laterally, it is overlapped by an anteromedial process of the postorbital. The frontals contact one another along the midline to form the dorsal surface of the anterior end of the braincase. The ventral surface of the frontal forming the dorsal edge of the orbit houses several small pockets that may have been pneumatic.
|
||
</paragraph>
|
||
<paragraph id="EC528811284C7E6FFD6E997CFDADAF73" blockId="4.[700,1229,1425,1741]" lastBlockId="5.[123,652,1044,1741]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
|
||
The orbit is bounded ventrally by the jugal and posteriorly by the postorbital, which meets the ascending process of the jugal to form a postorbital bar. The infratemporal fenestra is subrectangular, with straight anterior and ventral margins and rounded dorsal and posterior margins. The anterior process of the quadratojugal forms most of the ventral border, and the ascending process forms most of the posterior border of the fenestra. The anterior process is longer than the ascending process. In addition, the quadratojugal possesses a distinct posterior process, as in dromaeosaurids. A large quadrate foramen lies between the quadrate and quadratojugal just dorsal to the posterior angle of the infratemporal fenestra (
|
||
<figureCitation id="74D69494284D7E6FFE869A56FEE3AC8D" box="[305,372,1396,1419]" captionStart="Fig" captionStartId="7.[725,763,755,776]" captionTargetBox="[699,1227,186,732]" captionTargetId="figure-34@7.[699,1227,186,733]" captionTargetPageId="7" captionText="Fig. 6. Occipital view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734852" httpUri="https://zenodo.org/record/4734852/files/figure.png" pageId="5" pageNumber="6">fig. 6</figureCitation>
|
||
). The quadrate is massive and tightly sutured to the squamosal, quadratojugal, pterygoid, and braincase. The distal articular surface forms two condyles of approximately equal surface and curvature that are separated by a longitudinal groove. The pterygoid flange of the quadrate is tall and extends posterodorsally to meet the short descending process of the squamosal.
|
||
</paragraph>
|
||
<caption id="B892D899284D7E6FFF229C8AFE82AADE" ID-DOI="http://doi.org/10.5281/zenodo.4734840" ID-Zenodo-Dep="4734840" httpUri="https://zenodo.org/record/4734840/files/figure.png" pageId="5" pageNumber="6" startId="5.[149,187,936,957]" targetBox="[123,1228,185,914]" targetPageId="5">
|
||
<paragraph id="EC528811284D7E6FFF229C8AFE82AADE" blockId="5.[123,1227,936,984]" pageId="5" pageNumber="6">
|
||
Fig. 2. Right lateral view of the holotype cranium of
|
||
<taxonomicName id="2BEDF392284D7E6FFD659C8AFC58AABB" authority="IGM" authorityName="IGM" box="[722,975,936,957]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE995403284D7E6FFD659C8AFC02AABB" box="[722,917,936,957]" italics="true" pageId="5" pageNumber="6">Citipati osmolskae</emphasis>
|
||
IGM
|
||
</taxonomicName>
|
||
100/978. Abbreviations in appendix 1.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="EC528811284D7E6FFF2F995EFBA9AC8D" blockId="5.[123,652,1044,1741]" lastBlockId="5.[699,1227,1044,1741]" pageId="5" pageNumber="6">The squamosal is a complex bone that forms the entire dorsal margin of the infratemporal fenestra. Medially, it underlies the parietal, extending nearly to the medial edge of the supratemporal fossa. Anteriorly, it lies medial to the postorbital and ventral to its posterior process. The squamosal curves posterolaterally, and its distal end is forked around the external auditory meatus and downturned. A small posttemporal fenestra may be present on the occipital surface between the squamosal and the paroccipital process as seen on the right side, although this may be an artifact. The articulation between the squamosal and the quadrate and quadratojugal is poorly exposed.</paragraph>
|
||
<paragraph id="EC528811284D7E6FFD6F9AB3FB03AFB6" blockId="5.[699,1227,1044,1741]" pageId="5" pageNumber="6">The postorbital is a triradiate bone forming most of the postorbital bar. It overlies the frontal anteriorly and the squamosal posteriorly, and it extends lateral to the jugal on the postorbital bar. The laterosphenoid has a short contact with the postorbital at the anterior end of the supratemporal fenestra. The postorbital forms the posterior border of a fossa dorsal to the orbit extending into the pneumatic recesses of the narial region.</paragraph>
|
||
<paragraph id="EC528811284D7E6CFD6F9994FC76AD85" blockId="5.[699,1227,1044,1741]" lastBlockId="6.[700,1229,1044,1741]" lastPageId="6" lastPageNumber="7" pageId="5" pageNumber="6">
|
||
The parietals are fused and form the dorsal surface of nearly half of the skull along the midline (
|
||
<figureCitation id="74D69494284E7E6CFF5E9BDCFEB9AC13" box="[233,302,1278,1301]" captionStart="Fig" captionStartId="7.[149,187,1152,1173]" captionTargetBox="[123,651,186,1130]" captionTargetId="figure-15@7.[123,651,186,1130]" captionTargetPageId="7" captionText="Fig. 5. Dorsal view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734856" httpUri="https://zenodo.org/record/4734856/files/figure.png" pageId="6" pageNumber="7">fig. 5</figureCitation>
|
||
). The fused parietals narrow anteriorly except where they expand slightly at their anterior end. The occipital portion of the parietal is very broad and is oriented obliquely at an angle of nearly 45
|
||
<emphasis id="DE995403284E7E6CFD939A51FDBAAC8D" bold="true" box="[548,557,1395,1419]" pageId="6" pageNumber="7">°</emphasis>
|
||
to face posterodorsally. The parietal forms little of the supratemporal fossa except posteriorly, where a descending flange borders the opening of a dorsal extension of the dorsal tympanic recess. This recess extends anteromedially parallel to the occipital margin immediately posterior to it, and lies almost entirely within the parietal. The CT scans of the skull suggest that the recess connects with a large space within the parietal and frontal dorsal to the brain cavity (
|
||
<figureCitation id="74D69494284E7E6CFDA59994FDC9AFCB" box="[530,606,1718,1741]" captionStart="Fig" captionStartId="8.[150,188,1435,1456]" captionTargetBox="[135,1219,189,1414]" captionTargetId="figure-16@8.[135,1219,189,1415]" captionTargetPageId="8" captionText="Fig. 7. CT scan transverse sections through the holotype skull of Citipati osmolskae IGM 100/978 at three levels showing a large, presumably pneumatized space dorsal to the endocranial cavity and its connection to the dorsal tympanic recess. Arrow indicates apparent connection medially from dorsal tympanic recess to dorsal space. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734858" httpUri="https://zenodo.org/record/4734858/files/figure.png" pageId="6" pageNumber="7">figs. 7</figureCitation>
|
||
,
|
||
<figureCitation id="74D69494284E7E6CFDDA9994FDEAAFCB" box="[621,637,1718,1741]" captionStart="Fig" captionStartId="9.[149,187,1064,1085]" captionTargetBox="[123,1227,187,1043]" captionTargetId="figure-13@9.[123,1227,187,1043]" captionTargetPageId="9" captionText="Fig. 8. CT scan parasagittal section through the right side of the holotype cranium of Citipati osmolskae IGM 100/978, showing the boundaries of the endocranial cavity and the large, presumably pneumatized space dorsal to it. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734862" httpUri="https://zenodo.org/record/4734862/files/figure.png" pageId="6" pageNumber="7">8</figureCitation>
|
||
). The supraoccipital is poorly preserved, but has a low vertical midline crest and may have been separated from the foramen magnum by the exoccipitals.
|
||
</paragraph>
|
||
<caption id="B892D899284E7E6CFD619CBAFC05AAE4" ID-DOI="http://doi.org/10.5281/zenodo.4734845" ID-Zenodo-Dep="4734845" httpUri="https://zenodo.org/record/4734845/files/figure.png" pageId="6" pageNumber="7" startId="6.[726,764,920,941]" targetBox="[702,1229,186,897]" targetPageId="6">
|
||
<paragraph id="EC528811284E7E6CFD619CBAFC05AAE4" blockId="6.[700,1229,920,994]" pageId="6" pageNumber="7">
|
||
Fig. 4. Anterior view of the holotype cranium of
|
||
<taxonomicName id="2BEDF392284E7E6CFD579C91FC7CAACE" authority="IGM" authorityName="IGM" box="[736,1003,947,968]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE995403284E7E6CFD579C91FC3CAACE" box="[736,939,947,968]" italics="true" pageId="6" pageNumber="7">Citipati osmolskae</emphasis>
|
||
IGM
|
||
</taxonomicName>
|
||
100/978. Abbreviations in appendix 1.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="B892D899284E7E6CFF219B47FEC5ADB6" ID-DOI="http://doi.org/10.5281/zenodo.4734848" ID-Zenodo-Dep="4734848" httpUri="https://zenodo.org/record/4734848/files/figure.png" pageId="6" pageNumber="7" startId="6.[150,188,1125,1146]" targetBox="[124,653,188,1105]" targetPageId="6">
|
||
<paragraph id="EC528811284E7E6CFF219B47FEC5ADB6" blockId="6.[124,653,1125,1200]" pageId="6" pageNumber="7">
|
||
Fig. 3. Palatal view of the holotype cranium of
|
||
<taxonomicName id="2BEDF392284E7E6CFF179BA2FE3CAD93" authority="IGM" authorityName="IGM" box="[160,427,1152,1173]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE995403284E7E6CFF179BA2FEFCAD93" box="[160,363,1152,1173]" italics="true" pageId="6" pageNumber="7">Citipati osmolskae</emphasis>
|
||
IGM
|
||
</taxonomicName>
|
||
100/978. Abbreviations in appendix 1.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="EC528811284E7E6CFD6E9BABFBE7AF3D" blockId="6.[700,1229,1044,1741]" pageId="6" pageNumber="7">
|
||
Within the parietal and frontals lies an extensive space dorsal to the endocranial cavity, as revealed by CT scans (
|
||
<figureCitation id="74D69494284E7E6CFB939BE6FBE6ADDD" box="[1060,1137,1220,1243]" captionStart="Fig" captionStartId="8.[150,188,1435,1456]" captionTargetBox="[135,1219,189,1414]" captionTargetId="figure-16@8.[135,1219,189,1415]" captionTargetPageId="8" captionText="Fig. 7. CT scan transverse sections through the holotype skull of Citipati osmolskae IGM 100/978 at three levels showing a large, presumably pneumatized space dorsal to the endocranial cavity and its connection to the dorsal tympanic recess. Arrow indicates apparent connection medially from dorsal tympanic recess to dorsal space. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734858" httpUri="https://zenodo.org/record/4734858/files/figure.png" pageId="6" pageNumber="7">figs. 7</figureCitation>
|
||
,
|
||
<figureCitation id="74D69494284E7E6CFB369BE6FB06ADDD" box="[1153,1169,1220,1243]" captionStart="Fig" captionStartId="9.[149,187,1064,1085]" captionTargetBox="[123,1227,187,1043]" captionTargetId="figure-13@9.[123,1227,187,1043]" captionTargetPageId="9" captionText="Fig. 8. CT scan parasagittal section through the right side of the holotype cranium of Citipati osmolskae IGM 100/978, showing the boundaries of the endocranial cavity and the large, presumably pneumatized space dorsal to it. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734862" httpUri="https://zenodo.org/record/4734862/files/figure.png" pageId="6" pageNumber="7">8</figureCitation>
|
||
). Indeed, this space is larger in volume than the endocranial cavity itself. It is continuous anteriorly with the circumnarial pneumatic cavities and is largest anteriorly, dorsal to the anterior part of the orbit. There is no apparent partitioning of the space, but in the anterior part of the space radioopaque areas laterally indicate what may be internal struts. The space attenuates posteriorly and extends nearly to the occiput (
|
||
<figureCitation id="74D69494284E7E6CFC7F9ACBFB9DAF06" box="[968,1034,1513,1536]" captionStart="Fig" captionStartId="9.[149,187,1064,1085]" captionTargetBox="[123,1227,187,1043]" captionTargetId="figure-13@9.[123,1227,187,1043]" captionTargetPageId="9" captionText="Fig. 8. CT scan parasagittal section through the right side of the holotype cranium of Citipati osmolskae IGM 100/978, showing the boundaries of the endocranial cavity and the large, presumably pneumatized space dorsal to it. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734862" httpUri="https://zenodo.org/record/4734862/files/figure.png" pageId="6" pageNumber="7">fig. 8</figureCitation>
|
||
), and it appears to connect with the dorsal tympanic recess above the otic regions of both sides.
|
||
</paragraph>
|
||
<paragraph id="EC528811284E7E6DFD6E9963FE70AF5E" blockId="6.[700,1229,1044,1741]" lastBlockId="7.[123,652,1278,1741]" lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7">The jugal is extremely slender. Posteriorly, it terminates about halfway along the infratemporal fenestra, where it lies lateral to the more robust quadratojugal. It bends anteroventrally anterior to the orbit and extends anteriorly beneath the anterior end of the antorbital fenestra. The right element was damaged beneath the orbit during the life of the animal. The postorbital process of the jugal is robust and forms a broad posterior surface of the orbit. The quadratojugal forms much of the bar beneath the infratemporal fenestra, where it is nearly circular in cross section, and extends anteriorly to beneath the postorbital bar. Dorsally, the quadratojugal extends to meet the squamosal, and it borders the quadrate foramen laterally.</paragraph>
|
||
<caption id="B892D899284F7E6DFD629DD1FC26AA3B" ID-DOI="http://doi.org/10.5281/zenodo.4734852" ID-Zenodo-Dep="4734852" httpUri="https://zenodo.org/record/4734852/files/figure.png" pageId="7" pageNumber="8" startId="7.[725,763,755,776]" targetBox="[699,1227,186,732]" targetPageId="7">
|
||
<paragraph id="EC528811284F7E6DFD629DD1FC26AA3B" blockId="7.[699,1227,755,829]" pageId="7" pageNumber="8">
|
||
Fig. 6. Occipital view of the holotype cranium of
|
||
<taxonomicName id="2BEDF392284F7E6DFCBF9C2CFB98AA24" authority="IGM" authorityName="IGM" box="[776,1039,781,803]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE995403284F7E6DFCBF9C2CFC47AA25" box="[776,976,782,803]" italics="true" pageId="7" pageNumber="8">Citipati osmolskae</emphasis>
|
||
IGM
|
||
</taxonomicName>
|
||
100/978. Abbreviations in appendix 1.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="B892D899284F7E6DFF229BA2FEC6ADCC" ID-DOI="http://doi.org/10.5281/zenodo.4734856" ID-Zenodo-Dep="4734856" httpUri="https://zenodo.org/record/4734856/files/figure.png" pageId="7" pageNumber="8" startId="7.[149,187,1152,1173]" targetBox="[123,651,186,1130]" targetPageId="7">
|
||
<paragraph id="EC528811284F7E6DFF229BA2FEC6ADCC" blockId="7.[123,651,1152,1226]" pageId="7" pageNumber="8">
|
||
Fig. 5. Dorsal view of the holotype cranium of
|
||
<taxonomicName id="2BEDF392284F7E6DFF299BB9FE3EADB6" authority="IGM" authorityName="IGM" box="[158,425,1179,1200]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE995403284F7E6DFF299BB9FEFEADB6" box="[158,361,1179,1200]" italics="true" pageId="7" pageNumber="8">Citipati osmolskae</emphasis>
|
||
IGM
|
||
</taxonomicName>
|
||
100/978. Abbreviations in appendix 1.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="EC528811284F7E6DFF2F997CFC4FAD85" blockId="7.[123,652,1278,1741]" lastBlockId="7.[699,1228,897,1741]" pageId="7" pageNumber="8">The facial portion of the maxilla is very small and edentulous but includes a single osseous denticle in line with the premaxillary denticles. Posterior to the facial portion, the maxilla forms a narrow shelf ventral to the antorbital fossa. Below this shelf are a variable number of large, anteroposteriorly elongate openings in the ventrolateral surface of the maxilla, two on the right side and one on the left. Posteriorly, the maxilla contacts first the palatine and then the ectopterygoid along its ventral edge. Posterolaterally, the maxilla is overlain by the jugal.</paragraph>
|
||
<paragraph id="EC528811284F7E6DFD6F9BABFC9FAC8D" blockId="7.[699,1228,897,1741]" pageId="7" pageNumber="8">Within the antorbital fossa, the maxilla forms the medial wall. A slender posterodorsal process underlies the premaxilla and forms the dorsal edge of the accessory antorbital fenestra. The nasal overlies the posterodorsal part of the maxilla laterally, and the premaxilla in turn overlies the lacrimal laterally. The interfenestral bar was described above.</paragraph>
|
||
<paragraph id="EC528811284F7E62FD6F9AB3FF7FAFCB" blockId="7.[699,1228,897,1741]" lastBlockId="8.[124,653,1601,1741]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
|
||
The palate (
|
||
<figureCitation id="74D69494284F7E6DFCDD9AB3FC3BACAE" box="[874,940,1425,1448]" captionStart="Fig" captionStartId="6.[150,188,1125,1146]" captionTargetBox="[124,653,188,1105]" captionTargetId="figure-18@6.[124,653,188,1105]" captionTargetPageId="6" captionText="Fig. 3. Palatal view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734848" httpUri="https://zenodo.org/record/4734848/files/figure.png" pageId="7" pageNumber="8">fig. 3</figureCitation>
|
||
) is generally similar to that of an oviraptorid described by
|
||
<bibRefCitation id="887CF5E0284F7E6DFB399A8CFCFFACE5" author="Elzanowski, A." pageId="7" pageNumber="8" pagination="311 - 323" refId="ref11642" refString="Elzanowski, A. 1999. A comparison of the jaw skeleton in theropods and birds, with a description of the palate in the Oviraptoridae. Smithsonian Contributions to Paleobiology 89: 311 - 323." type="journal article" year="1999">Elzanowski (1999)</bibRefCitation>
|
||
. The lateral portion of the palate is oriented ventrolaterally, and medially the palatal portion forms a distinct posteroventrally directed midline process that surrounds the anterior end of the vomer. The maxilla forms most of the palate anterior to this, which bears four stout longitudinal ridges. A deep groove borders the lateral edge of the lateral ridge. The large choanae lie almost directly beneath the naris and antorbital fossa, such that the nasopharyngeal passageway was oriented almost vertically. The choanae are divided by the vomer posterior to the maxillae.
|
||
</paragraph>
|
||
<caption id="B892D89928407E62FF219AB9FC81AF06" ID-DOI="http://doi.org/10.5281/zenodo.4734858" ID-Zenodo-Dep="4734858" httpUri="https://zenodo.org/record/4734858/files/figure.png" pageId="8" pageNumber="9" startId="8.[150,188,1435,1456]" targetBox="[135,1219,189,1414]" targetPageId="8">
|
||
<paragraph id="EC52881128407E62FF219AB9FC81AF06" blockId="8.[124,1229,1435,1536]" pageId="8" pageNumber="9">
|
||
Fig. 7. CT scan transverse sections through the holotype skull of
|
||
<taxonomicName id="2BEDF39228407E62FCDB9AB9FBF9ACB6" authority="IGM" authorityName="IGM" box="[876,1134,1435,1456]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE99540328407E62FCDB9AB9FBA5ACB6" box="[876,1074,1435,1456]" italics="true" pageId="8" pageNumber="9">Citipati osmolskae</emphasis>
|
||
IGM
|
||
</taxonomicName>
|
||
100/978 at three levels showing a large, presumably pneumatized space dorsal to the endocranial cavity and its connection to the dorsal tympanic recess. Arrow indicates apparent connection medially from dorsal tympanic recess to dorsal space. Abbreviations in appendix 1.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="EC52881128407E63FD6E9963FE39AF1B" blockId="8.[700,1230,1601,1741]" lastBlockId="9.[123,652,1190,1741]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
|
||
The palatines are anteroposteriorly short and highly modified. They meet on the midline along the posterior edge of the choanae, where their anterior edge curves dorsally to become vertical and transversely oriented. Laterally, the palatine curves dorsally to meet the maxilla and is separated from the anterior margin of the ectopterygoid by a small, circular, nearly laterally facing suborbital fenestra (the ‘‘postpalatine fenestra’’ of
|
||
<bibRefCitation id="887CF5E028417E63FFCC9A1BFEC6AC56" author="Elzanowski, A." box="[123,337,1337,1360]" pageId="9" pageNumber="10" pagination="311 - 323" refId="ref11642" refString="Elzanowski, A. 1999. A comparison of the jaw skeleton in theropods and birds, with a description of the palate in the Oviraptoridae. Smithsonian Contributions to Paleobiology 89: 311 - 323." type="journal article" year="1999">Elzanowski, 1999</bibRefCitation>
|
||
). The ectopterygoid is vertical, meeting the maxilla laterally and the pterygoid medially and abutting the palatine throughout most of its length. The vomers are fused into a single short, solid bone that does not extend posterior beyond its contact with the pterygoid and palatine at the posterior edge of the choana.
|
||
</paragraph>
|
||
<caption id="B892D89928417E63FF229B0AFC90AD74" ID-DOI="http://doi.org/10.5281/zenodo.4734862" ID-Zenodo-Dep="4734862" httpUri="https://zenodo.org/record/4734862/files/figure.png" pageId="9" pageNumber="10" startId="9.[149,187,1064,1085]" targetBox="[123,1227,187,1043]" targetPageId="9">
|
||
<paragraph id="EC52881128417E63FF229B0AFC90AD74" blockId="9.[123,1227,1064,1138]" pageId="9" pageNumber="10">
|
||
Fig. 8. CT scan parasagittal section through the right side of the holotype cranium of
|
||
<taxonomicName id="2BEDF39228417E63FBCC9B0AFEB3AD5E" authority="IGM" authorityName="IGM" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE99540328417E63FBCC9B0AFF7FAD5E" italics="true" pageId="9" pageNumber="10">Citipati osmolskae</emphasis>
|
||
IGM
|
||
</taxonomicName>
|
||
100/978, showing the boundaries of the endocranial cavity and the large, presumably pneumatized space dorsal to it. Abbreviations in appendix 1.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="EC52881128417E60FF2F9906FF62A8FB" blockId="9.[123,652,1190,1741]" lastBlockId="10.[124,653,193,1741]" lastPageId="10" lastPageNumber="11" pageId="9" pageNumber="10">
|
||
The pterygoids have elongate palatal processes that do not appear to meet along the midline until they contact the vomers. The palatal surface of each element has a longitudinal concavity, confluent anteriorly with the choana. The pterygoid flange is greatly reduced compared to theropods with teeth. It is anteroposteriorly elongate, and, in lateral view, it has a linear dorsal edge and a ventrally convex, crescentic ventral edge, the ectopterygoid forming the anterior half. The flange does not extend laterally beyond the body of the pterygoid, indicating that the
|
||
<taxonomicName id="2BEDF39228417E63FB1C9A75FCC6AC8D" class="Reptilia" family="Caenagnathidae" genus="Microvenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="pterygoideus">
|
||
<emphasis id="DE99540328417E63FB1C9A75FCC6AC8D" italics="true" pageId="9" pageNumber="10">M. pterygoideus</emphasis>
|
||
</taxonomicName>
|
||
was relatively small. There is no evidence for a subsidiary palatal fenestra between the pterygoid and palatine. Posteriorly, the pterygoids converge towards the midline at the basipterygoid joint, but they do not come into contact, suggesting that if a contact was present it was formed by soft tissue. The posteromedial edge of the pterygoid is expanded near the midline, as described by
|
||
<bibRefCitation id="887CF5E028417E63FCF7995EFBB4AF95" author="Elzanowski, A." box="[832,1059,1660,1683]" pageId="9" pageNumber="10" pagination="311 - 323" refId="ref11642" refString="Elzanowski, A. 1999. A comparison of the jaw skeleton in theropods and birds, with a description of the palate in the Oviraptoridae. Smithsonian Contributions to Paleobiology 89: 311 - 323." type="journal article" year="1999">Elzanowski (1999)</bibRefCitation>
|
||
. The quadrate ramus of the pterygoid is anteroposteriorly short but very tall, and with the quadrate forms the lateral surface of a deep pocket lateral to the braincase that narrows anteriorly. The pterygoid extends posteriorly to end medial to the mandibular articulating surface of the quadrate. A small, dorsomedially opening pocket is formed dorsal to the posterior end at the level of the quadrate foramen. Another pocket on the dorsal surface of the pterygoid lateral to the basipterygoid joint extends anteriorly into the body of the pterygoid.
|
||
</paragraph>
|
||
<paragraph id="EC52881128427E60FF2E9D26FE6FAA7D" blockId="10.[124,653,193,1741]" pageId="10" pageNumber="11">
|
||
An isolated oviraptorid quadrate was described by
|
||
<bibRefCitation id="887CF5E028427E60FF489D03FD1FAB3E" author="Maryanska, T. & H. Osmolska" box="[255,648,545,568]" pageId="10" pageNumber="11" pagination="377 - 387" refId="ref12229" refString="Maryanska, T., and H. Osmolska. 1997. The quadrate of oviraptorid dinosaurs. Acta Palaeontologia Polonica 42: 377 - 387." type="journal article" year="1997">Maryanska and Osmólska (1997)</bibRefCitation>
|
||
, and the quadrate of
|
||
<taxonomicName id="2BEDF39228427E60FEDF9D1DFE56AB50" box="[360,449,575,598]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="10" pageNumber="11" phylum="Chordata" rank="genus">
|
||
<emphasis id="DE99540328427E60FEDF9D1DFE56AB50" box="[360,449,575,598]" italics="true" pageId="10" pageNumber="11">Citipati</emphasis>
|
||
</taxonomicName>
|
||
is generally similar. It is oriented obliquely in lateral view, extending posteroventrally from the squamosal. The quadrate has a mediolaterally biconvex mandibular trochlea. The vertical body of the quadrate narrows dorsally, but its dorsal articulation with the squamosal and, apparently, the paroccipital process is not exposed. The pterygoid ramus of the quadrate is very tall, equal in height to about half that of the occipital region.
|
||
</paragraph>
|
||
<paragraph id="EC52881128427E60FF2E9CA3FF74AC35" blockId="10.[124,653,193,1741]" pageId="10" pageNumber="11">An epipterygoid is present on both sides of the skull. It is much broader anteroposteriorly than it is thick. It has a broad articulation ventrally with the pterygoid, where it faces anteromedially, and it twists laterally as it narrows dorsally. Dorsally, it contacts the ventrolateral surface of the coossified braincase where a vertical ridge on the laterosphenoid converges with a ridge along the capitate process. On the left side it appears to be fused to the braincase, but not on the right. The dorsal tip twists dorsolaterally and is expanded in the transverse plane, aligned with the vertical ridge on the laterosphenoid above it.</paragraph>
|
||
<paragraph id="EC52881128427E60FF2E9A1BFB21A8A3" blockId="10.[124,653,193,1741]" lastBlockId="10.[700,1230,193,1741]" pageId="10" pageNumber="11">
|
||
The elements comprising the braincase are coossified, although some areas of contact between bones are apparent. The basioccipital is notable for the posterior extension of the occipital condyle well beyond the posterior end of the foramen magnum (
|
||
<figureCitation id="74D6949428427E60FD8D9AEEFDE9ACE5" box="[570,638,1484,1507]" captionStart="Fig" captionStartId="7.[149,187,1152,1173]" captionTargetBox="[123,651,186,1130]" captionTargetId="figure-15@7.[123,651,186,1130]" captionTargetPageId="7" captionText="Fig. 5. Dorsal view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734856" httpUri="https://zenodo.org/record/4734856/files/figure.png" pageId="10" pageNumber="11">fig. 5</figureCitation>
|
||
). Indeed, with the floor of the braincase horizontal most if not all of the basioccipital lies posterior to the dorsal edge of the foramen magnum due to the anterodorsal slope of the occiput. The condyle has a nearly flat posterior surface with a central depression. The basioccipital has a midline depression on its ventral surface. The foramen magnum is nearly circular except for a ventral midline depression (
|
||
<figureCitation id="74D6949428427E60FCFD9FFCFC1BA9F3" box="[842,908,222,245]" captionStart="Fig" captionStartId="7.[725,763,755,776]" captionTargetBox="[699,1227,186,732]" captionTargetId="figure-34@7.[699,1227,186,733]" captionTargetPageId="7" captionText="Fig. 6. Occipital view of the holotype cranium of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734852" httpUri="https://zenodo.org/record/4734852/files/figure.png" pageId="10" pageNumber="11">fig. 6</figureCitation>
|
||
). The lateral surface of the basioccipital is concave, and at least three small openings within the concavity presumably enter a pneumatic recess in the body of the bone. A large opening in this region on the right side is presumably due to the breakage of thinwalled bone covering a cavity.
|
||
</paragraph>
|
||
<paragraph id="EC52881128427E60FD6E9E8EFC25AAD5" blockId="10.[700,1230,193,1741]" pageId="10" pageNumber="11">A large vacuity is present on the ventral midline between the basioccipital and basisphenoid. The foramen lies mainly within the basisphenoid, dividing it in two posteriorly. The basisphenoid and basioccipital expand at their contact, and on the right side they appear to be fused. The ventral surface of the basisphenoid descends anteroventrally from this contact. Basipterygoid processes are absent, although a matrixfilled area between the basisphenoid and pterygoids suggests that poorly ossified processes may have been present. The basisphenoid has several pneumatic openings and cavities on the lateral surface of the braincase ventral to the trigeminal foramen. These are poorly exposed, covered laterally by the epipterygoid and beneath matrix ventrally. Several delicate struts span these openings.</paragraph>
|
||
<paragraph id="EC52881128427E60FD6E9CFBFB5AADDD" blockId="10.[700,1230,193,1741]" pageId="10" pageNumber="11">
|
||
A delicate, tall parasphenoid process extends above the length of the interpterygoid vacuity, terminating slightly posterior to the level of the vomer. It descends and tapers anteroventrally from its tall posterior base. The lateral surface of the base has a shallow depression dorsally that opens anterodorsally, presumably the site of origin of an ocular muscle, perhaps the
|
||
<emphasis id="DE99540328427E60FC1D9BE6FB51ADDD" box="[938,1222,1220,1243]" italics="true" pageId="10" pageNumber="11">
|
||
<taxonomicName id="2BEDF39228427E60FC1D9BE6FBCEADDD" box="[938,1113,1220,1243]" class="Reptilia" family="Caenagnathidae" genus="Microvenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="10" pageNumber="11" phylum="Chordata" rank="subSpecies" species="rectus" subSpecies="oculi">M. rectus oculi</taxonomicName>
|
||
ventralis
|
||
</emphasis>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="EC52881128427E61FD6E9BC3FDF2AA03" blockId="10.[700,1230,193,1741]" lastBlockId="11.[123,652,193,1741]" lastPageId="11" lastPageNumber="12" pageId="10" pageNumber="11">
|
||
The laterosphenoid portion of the braincase forms a ventral floor anteriorly, and a discrete orbitosphenoid ossification is present anterior to it. The large, horizontal orbitosphenoids encircle the anterior part of the endocranial cavity, meeting on the midline dorsally and ventrally. Posterior to the orbitosphenoid the laterosphenoid sends a short, ventrally expanding process ventrally along the midline. This process forms the anterior edge of a large foramen, presumably for cranial nerve (CN) II, that is open ventrally. A posterior descending process forms the posterior edge of this foramen and nearly contacts the top of the parasphenoid rostrum. The capitate process of the laterosphenoid is long and slender and extends laterally along the ventral surface of the frontal and contacts the postorbital. A small slit separates the process from the frontal anteriorly. A sharp ridge descends posteromedially from the capitate process to the epipterygoid contact. A vertical ridge within the supratemporal fossa presumably separated portions of the
|
||
<taxonomicName id="2BEDF39228437E61FD8C9E53FECCA8A0" class="Reptilia" family="Caenagnathidae" genus="Microvenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="11" pageNumber="12" phylum="Chordata" rank="subSpecies" species="adductor" subSpecies="mandibulae">
|
||
<emphasis id="DE99540328437E61FD8C9E53FECCA8A0" italics="true" pageId="11" pageNumber="12">M. adductor mandibulae</emphasis>
|
||
</taxonomicName>
|
||
. The dorsal contact with the frontal and parietal is apparent but posteriorly the laterosphenoid is fused to the prootic. The posterodorsal contact with the parietal slopes anterodorsally parallel to the occiput. The trigeminal opening (CN V) is relatively small compared with dromaeosaurids (e.g.,
|
||
<taxonomicName id="2BEDF39228437E61FF5C9D7EFE5EAB75" authority=", IGM" authorityName="IGM" box="[235,457,604,627]" class="Reptilia" family="Dromaeosauridae" genus="Velociraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="11" pageNumber="12" phylum="Chordata" rank="genus">
|
||
<emphasis id="DE99540328437E61FF5C9D7EFE17AB75" box="[235,384,604,627]" italics="true" pageId="11" pageNumber="12">Velociraptor</emphasis>
|
||
, IGM
|
||
</taxonomicName>
|
||
100/976) and is smaller than CN II of this specimen. It is slightly longer than tall and exits ventrolaterally from the braincase. The lateral surface of the laterosphenoid is rugose, presumably indicating an origin of adductor muscles.
|
||
</paragraph>
|
||
<paragraph id="EC52881128437E61FF2F9C2EFE72AD2D" blockId="11.[123,652,193,1741]" pageId="11" pageNumber="12">
|
||
The prootic has a welldeveloped fossa dorsally, homologous to the dorsal tympanic recess described in dromaeosaurids (
|
||
<bibRefCitation id="887CF5E028437E61FD889C64FE1FAA7D" author="Norell, M. A. & P. Makovicky" pageId="11" pageNumber="12" pagination="1 - 28" refId="ref12705" refString="Norell, M. A., and P. Makovicky. 1997. Important features of the dromaeosaur skeleton: information from a new specimen. American Museum Novitates 3215: 1 - 28." type="journal article" year="1997">Norell and Makovicky, 1997</bibRefCitation>
|
||
, 1999). A horizontal swelling forms the ventrolateral border of the fossa. Dorsal to this fossa lies a large pneumatic opening extending medially into the braincase, as described above. The prootic apparently surrounds this opening dorsally, separating it from the parietal.
|
||
</paragraph>
|
||
<paragraph id="EC52881128437E61FF2F9B13FD82ACAE" blockId="11.[123,652,193,1741]" pageId="11" pageNumber="12">The opening in the prootic for CN VII is well preserved on the right side, but on the left it is apparently within a crushed area just ventral to the horizontal swelling. On the right side a depression with at least two, presumably pneumatic, openings lies dorsal to the foramen for CN VII, but this depression is absent on the left side. The otic recess is poorly preserved on both sides, and the fenestrae ovale and rotundum cannot be identified within it. The recess is larger on the right side than the left, and both are small compared with other coelurosaurs.</paragraph>
|
||
<paragraph id="EC52881128437E61FF2F9A8CFC80ABAB" blockId="11.[123,652,193,1741]" lastBlockId="11.[699,1228,193,1741]" pageId="11" pageNumber="12">The contacts between the opisthotic and the prootic and basioccipital appear to be fused. A posterior tympanic recess is present within the anterior surface of the paroccipital process on the right side, but this region is broken on the left. The recess continues medially into the braincase from within a shallow fossa. The fossa penetrates the posterior surface of the paroccipital process via a narrow, mediolaterally elongate opening. In medial view, the jugular foramen and the exits of two hypoglossal foramina are visible within the braincase. The hypoglossal foramina have a short, simple course ventrolaterally from the foramen magnum, opening into a shallow depression. Two ventrolaterally opening foramina on the ventral edge of the paroccipital process presumably originate within the metotic foramen, and likely conveyed CN IX and X and the associated vasculature. The more ventromedial of the two is larger and may be the foramen vagi. The more lateral foramen is small, similar in size to the hypoglossal foramina, and possibly for CN IX. The paroccipital processes are pendant and curve medioventrally, attenuating distally.</paragraph>
|
||
<paragraph id="EC52881128437E61FD6F9D96FC1DAAD5" blockId="11.[699,1228,193,1741]" pageId="11" pageNumber="12">
|
||
Both stapes are preserved, the left element incompletely and the right disarticulated from the skull. The stapes is elongate, straight, and slender, with a footplate that is approximately three times the diameter of the shaft. As implied by the position of the otic notch in the squamosal, the stapes would have projected posterolaterally from the fenestra ovalis at an angle of about 45
|
||
<emphasis id="DE99540328437E61FB3D9CBCFB04AAB0" bold="true" box="[1162,1171,926,950]" pageId="11" pageNumber="12">°</emphasis>
|
||
and slightly ventrally.
|
||
</paragraph>
|
||
<paragraph id="EC52881128437E61FD6F9CFBFCECADBB" blockId="11.[699,1228,193,1741]" pageId="11" pageNumber="12">
|
||
Five scleral ossicles, from a partial scleral ring, preserved in the right orbit and below it were removed during preparation (
|
||
<figureCitation id="74D6949428437E61FBCE9B36FB2EAD2D" box="[1145,1209,1044,1067]" captionStart="Fig" captionStartId="4.[150,188,1280,1301]" captionTargetBox="[124,1229,187,1259]" captionTargetId="figure-16@4.[124,1229,187,1259]" captionTargetPageId="4" captionText="Fig. 1. The holotype skull and mandible of Citipati osmolskae (IGM 100/978) in right lateral view before final preparation. The mandible, scleral ossicles, and hyoid elements were later separated from the skull. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734838" httpUri="https://zenodo.org/record/4734838/files/figure.png" pageId="11" pageNumber="12">fig. 1</figureCitation>
|
||
). They are roughly square in shape with irregular margins and a gently concave medial surface. They are similar to those of other basal coelurosaurs such as troodontids and dromaeosaurids.
|
||
</paragraph>
|
||
<paragraph id="EC52881128437E66FD6F9BE6FDECAF95" blockId="11.[699,1228,193,1741]" lastBlockId="12.[124,653,1337,1741]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
|
||
The edentulous mandible (
|
||
<figureCitation id="74D6949428437E61FBAE9BE6FBCDADDD" box="[1049,1114,1220,1243]" captionStart="Fig" captionStartId="12.[150,188,1229,1250]" captionTargetBox="[124,1230,186,1206]" captionTargetId="figure-16@12.[123,1230,186,1207]" captionTargetPageId="12" captionText="Fig. 9. (A) Lateral and (B) medial views of the left hemimandible of the holotype of Citipati osmolskae IGM 100/978. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734866" httpUri="https://zenodo.org/record/4734866/files/figure.png" pageId="11" pageNumber="12">fig. 9</figureCitation>
|
||
) is dominated by a high, arching coronoid eminence. The symphysis is short and broad, with a transversely oriented, upturned anterior edge. It is slightly below the level of the mandibular articulation. The dorsal surface of the symphysis is concave dorsally behind the anterior edge, and the posterior edge of the symphysis descends posteroventrally. The coronoid eminence rises abruptly from the posterior part of the symphysis and flattens dorsally. The surangular descends gradually from the eminence and extends posteriorly to the end of the mandible, completely covering the lateral surface of the articular. The large mandibular fenestra is divided by an anterior process of the surangular that reaches approximately threequarters of the distance across the fenestra. The lateral surface of the surangular is gently depressed anterior to the mandibular articulation. The adductor fossa on the medial surface of the mandibular ramus is extremely large and bordered ventrally by a slender splenial. The anterior end of the splenial is divided into dorsal and ventral rami in the symphysis. It overlaps the prearticular in the middle of the mandibular ramus, and the prearticular extends posteriorly to the end of the mandible, covering most of the medial surface of the articular.
|
||
</paragraph>
|
||
<caption id="B892D89928447E66FF219BEFFD5DADFB" ID-DOI="http://doi.org/10.5281/zenodo.4734866" ID-Zenodo-Dep="4734866" httpUri="https://zenodo.org/record/4734866/files/figure.png" pageId="12" pageNumber="13" startId="12.[150,188,1229,1250]" targetBox="[124,1230,186,1206]" targetPageId="12">
|
||
<paragraph id="EC52881128447E66FF219BEFFD5DADFB" blockId="12.[124,1229,1229,1277]" pageId="12" pageNumber="13">
|
||
Fig. 9.
|
||
<emphasis id="DE99540328447E66FF4D9BEFFE88ADE4" bold="true" box="[250,287,1229,1250]" pageId="12" pageNumber="13">(A)</emphasis>
|
||
Lateral and
|
||
<emphasis id="DE99540328447E66FE029BEFFE40ADE4" bold="true" box="[437,471,1229,1250]" pageId="12" pageNumber="13">(B)</emphasis>
|
||
medial views of the left hemimandible of the holotype of
|
||
<taxonomicName id="2BEDF39228447E66FBCA9BECFEB2ADFB" authority="IGM" authorityName="IGM" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE99540328447E66FBCA9BECFF7EADFB" italics="true" pageId="12" pageNumber="13">Citipati osmolskae</emphasis>
|
||
IGM
|
||
</taxonomicName>
|
||
100/978. Abbreviations in appendix 1.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="EC52881128447E66FF2E99BBFBEFAFB6" blockId="12.[124,653,1337,1741]" lastBlockId="12.[700,1229,1337,1741]" pageId="12" pageNumber="13">
|
||
The mandibular articulation surface is anteroposteriorly elongate and unbounded anteriorly or posteriorly, suggesting that the mandible was capable of anteroposterior movement (cf.
|
||
<bibRefCitation id="887CF5E028447E66FCD99A56FBA7AC8D" author="Osmolska, H." box="[878,1072,1396,1419]" pageId="12" pageNumber="13" pagination="683 - 684" refId="ref12843" refString="Osmolska, H. 1976. New light on skull anatomy and systematic position of Oviraptor. Nature 262: 683 - 684." type="journal article" year="1976">Osmólska, 1976</bibRefCitation>
|
||
). The surface bears a longitudinal midline ridge and faces posterodorsally. The retroarticular process descends posteroventrally from the articular surface, with which it is coplanar, and its ventral edge descends slightly below the level of the remainder of the mandible. The mandibular articulation is much wider than the remainder of the mandibular ramus, and it is bordered ventrally by the prearticular medially and the surangular laterally.
|
||
</paragraph>
|
||
<paragraph id="EC52881128447E67FD6E9994FE44ADBB" blockId="12.[700,1229,1337,1741]" lastBlockId="13.[123,652,956,1741]" lastPageId="13" lastPageNumber="14" pageId="12" pageNumber="13">
|
||
A very small coronoid element appears to be present. A splint of bone on the medial surface of the posterodorsal ramus of the dentary on both sides is in the position of a coronoid bone, although this bone is reported as absent in other oviraptorids (
|
||
<bibRefCitation id="887CF5E028457E67FDB79B13FF7FAD63" author="Barsbold, R. & T. Maryanska & H. Osmolska" pageId="13" pageNumber="14" pagination="249 - 258" refId="ref11058" refString="Barsbold, R., T. Maryanska, and H. Osmolska. 1990. Oviraptorosauria. In D. B. Weishampel, P. Dodson, and H. Osmolska (editors), The Dinosauria: 249 - 258. Berkeley: University of California Press." type="book chapter" year="1990">Barsbold et al., 1990</bibRefCitation>
|
||
). A bone as small as this could be easily lost or overlooked in poorly preserved specimens, suggesting that it may also be present in other oviraptorids.
|
||
</paragraph>
|
||
<caption id="B892D89928457E67FF229C19FCD0AA6C" ID-DOI="http://doi.org/10.5281/zenodo.4734868" ID-Zenodo-Dep="4734868" httpUri="https://zenodo.org/record/4734868/files/figure.png" pageId="13" pageNumber="14" startId="13.[149,187,827,848]" targetBox="[123,1228,185,805]" targetPageId="13">
|
||
<paragraph id="EC52881128457E67FF229C19FCD0AA6C" blockId="13.[123,1227,827,874]" pageId="13" pageNumber="14">
|
||
Fig. 10. Ventral view of the holotype cranium of
|
||
<taxonomicName id="2BEDF39228457E67FD739C19FC50AA56" authority="IGM" authorityName="IGM" box="[708,967,827,848]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE99540328457E67FD739C19FC1DAA56" box="[708,906,827,848]" italics="true" pageId="13" pageNumber="14">Citipati osmolskae</emphasis>
|
||
IGM
|
||
</taxonomicName>
|
||
100/978 showing a left hyoid element in natural articulation. Abbreviations in appendix 1.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="EC52881128457E67FF2F9BE6FEEDACE5" blockId="13.[123,652,956,1741]" pageId="13" pageNumber="14">
|
||
A pair of elements of the hyoid apparatus were preserved beneath the mandible (
|
||
<figureCitation id="74D6949428457E67FDD39BC3FF0AAC13" captionStart="Fig" captionStartId="13.[149,187,827,848]" captionTargetBox="[123,1228,185,805]" captionTargetId="figure-13@13.[123,1228,185,805]" captionTargetPageId="13" captionText="Fig. 10. Ventral view of the holotype cranium of Citipati osmolskae IGM 100/978 showing a left hyoid element in natural articulation. Abbreviations in appendix 1." figureDoi="http://doi.org/10.5281/zenodo.4734868" httpUri="https://zenodo.org/record/4734868/files/figure.png" pageId="13" pageNumber="14">fig. 10</figureCitation>
|
||
), presumably the ceratohyals (cornua branchiala I). They are simple structures, essentially elongate rods that curve dorsally posteriorly. The anterior and posterior ends are expanded and slightly compressed mediolaterally. They were preserved paralleling the ventral edge of the mandible, possibly their natural position.
|
||
</paragraph>
|
||
<paragraph id="EC52881128457E67FF2F9ACBFBFEAAF6" blockId="13.[123,652,956,1741]" lastBlockId="13.[699,1228,956,1302]" pageId="13" pageNumber="14">
|
||
Most elements of the postcranial skeleton of
|
||
<taxonomicName id="2BEDF39228457E67FF299925FEABAF18" box="[158,316,1543,1566]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE99540328457E67FF299925FEABAF18" box="[158,316,1543,1566]" italics="true" pageId="13" pageNumber="14">C. osmolskae</emphasis>
|
||
</taxonomicName>
|
||
were described by
|
||
<bibRefCitation id="887CF5E028457E67FD9C9924FF63AF3D" author="Clark, J. M. & M. Norell & L. Chiappe" pageId="13" pageNumber="14" pagination="1 - 36" refId="ref11210" refString="Clark, J. M., M. Norell, and L. Chiappe. 1999. An oviraptorid skeleton from the Late Cretaceous of Ukhaa Tolgod, Mongolia, preserved in an avian-like brooding position over an oviraptorid nest. American Museum Novitates 3265: 1 - 36." type="journal article" year="1999">Clark et al. (1999)</bibRefCitation>
|
||
for the referred specimen IGM 100/979. This specimen preserves only a single vertebra, however, and lacks the ilia and most of the scapulae, pubes, and ischia. It differs from
|
||
<taxonomicName id="2BEDF39228457E67FEA399BBFEF5AFB6" box="[276,354,1689,1712]" class="Reptilia" family="Oviraptoridae" genus="Khaan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
|
||
<emphasis id="DE99540328457E67FEA399BBFEF5AFB6" box="[276,354,1689,1712]" italics="true" pageId="13" pageNumber="14">Khaan</emphasis>
|
||
</taxonomicName>
|
||
in that the first metacarpal of the latter is reduced proximally and the bone does not expand, whereas that of
|
||
<taxonomicName id="2BEDF39228457E67FD0C9CFBFC83AAF6" box="[699,788,985,1008]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
|
||
<emphasis id="DE99540328457E67FD0C9CFBFC83AAF6" box="[699,788,985,1008]" italics="true" pageId="13" pageNumber="14">Citipati</emphasis>
|
||
</taxonomicName>
|
||
is unreduced and expanded.
|
||
</paragraph>
|
||
<paragraph id="EC52881128457E67FD6F9CD4FCA7AC13" blockId="13.[699,1228,956,1302]" pageId="13" pageNumber="14">
|
||
The postcranial skeleton of the
|
||
<typeStatus id="335636B328457E67FBF19CD4FB3BAD0B" box="[1094,1196,1014,1037]" pageId="13" pageNumber="14" type="holotype">holotype</typeStatus>
|
||
of
|
||
<taxonomicName id="2BEDF39228457E67FD0C9B36FCCFAD2D" box="[699,856,1044,1067]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="osmolskae">
|
||
<emphasis id="DE99540328457E67FD0C9B36FCCFAD2D" box="[699,856,1044,1067]" italics="true" pageId="13" pageNumber="14">C. osmolskae</emphasis>
|
||
</taxonomicName>
|
||
is very well preserved and includes representatives of nearly every element, but it has not yet been fully prepared. The cervical vertebrae are distinctly more elongate than those of
|
||
<taxonomicName id="2BEDF39228457E67FC7B9BABFB8DADA6" box="[972,1050,1161,1184]" class="Reptilia" family="Oviraptoridae" genus="Khaan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
|
||
<emphasis id="DE99540328457E67FC7B9BABFB8DADA6" box="[972,1050,1161,1184]" italics="true" pageId="13" pageNumber="14">Khaan</emphasis>
|
||
</taxonomicName>
|
||
and other oviraptorids where known. The cervical ribs are also more elongate, longer than the corresponding centrum in
|
||
<taxonomicName id="2BEDF39228457E67FC049BC3FB9BADFE" box="[947,1036,1249,1272]" class="Reptilia" family="Oviraptoridae" genus="Citipati" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
|
||
<emphasis id="DE99540328457E67FC049BC3FB9BADFE" box="[947,1036,1249,1272]" italics="true" pageId="13" pageNumber="14">Citipati</emphasis>
|
||
</taxonomicName>
|
||
but shorter than in
|
||
<taxonomicName id="2BEDF39228457E67FD6C9BDDFCBEAC10" box="[731,809,1279,1302]" class="Reptilia" family="Oviraptoridae" genus="Khaan" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="13" pageNumber="14" phylum="Chordata" rank="genus">
|
||
<emphasis id="DE99540328457E67FD6C9BDDFCBEAC10" box="[731,809,1279,1302]" italics="true" pageId="13" pageNumber="14">Khaan</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |