458 lines
60 KiB
XML
458 lines
60 KiB
XML
<document id="629234EF9689FAD081ACF7D7E3CCD0CA" ID-DOI="10.1093/zoolinnean/zlad169" ID-ISSN="0024-4082" ID-Zenodo-Dep="14284228" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="GgImagineBatch" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1732842600575" checkinUser="plazi" docAuthor="Słowiak, Justyna, Brusatte, Stephen L. & Szczygielski, Tomasz" docDate="2024" docId="03836047994DFFDDB90003AAFED1FFFC" docLanguage="en" docName="zlad169.pdf" docOrigin="Zoological Journal of the Linnean Society 202 (3)" docSource="https://doi.org/10.1093/zoolinnean/zlad169" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.9:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="7" docTitle="Bagaraatan ostromi Osmolska 1996" docType="treatment" docVersion="3" lastPageNumber="34" masterDocId="FFBA183F996DFFFFB83E033AFFE1FF3B" masterDocTitle="Reassessment of the enigmatic Late Cretaceous theropod dinosaur, Bagaraatan ostromi" masterLastPageNumber="39" masterPageNumber="1" pageNumber="33" updateTime="1733416834927" updateUser="ExternalLinkService">
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<mods:title id="259A0CA21765C8C1A97729ADE1334172">Reassessment of the enigmatic Late Cretaceous theropod dinosaur, Bagaraatan ostromi</mods:title>
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<mods:namePart id="372731EFCC1E8B41EAA44C5867BB53D3">Słowiak, Justyna</mods:namePart>
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<mods:affiliation id="7579D2B425769A1307A6DDC41F00E411">Institute of Paleobiology, Polish Academy of Sciences, Warsaw, Poland</mods:affiliation>
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<mods:nameIdentifier id="5FC93E29D629F14B179492FBB8C4B988" type="email">justyna.slowiak@twarda.pan.pl</mods:nameIdentifier>
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<mods:namePart id="4886FFAF9FF248C9799CF785921B1C85">Brusatte, Stephen L.</mods:namePart>
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<mods:affiliation id="CB8864AD6FFF3667EB54CCEBDC3EE4B8">School of GeoSciences, University of Edinburgh, Edinburgh, UK</mods:affiliation>
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<mods:namePart id="6887DACA1660F33C402D7D980604E4F6">Szczygielski, Tomasz</mods:namePart>
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<mods:affiliation id="C32395F873CFA599DE9877AC33E1EAFA">Institute of Paleobiology, Polish Academy of Sciences, Warsaw, Poland</mods:affiliation>
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<mods:title id="414F11F6C7404D1F4C6789667E161AE2">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="964FDFB66C3AE40FF08B425EFA61308C">2024</mods:date>
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<mods:number id="87BF049D463B22DF50F9F1C288DC602E">2024-02-16</mods:number>
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<mods:number id="598502BE606921351F39E3C804055F09">202</mods:number>
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<treatment id="03836047994DFFDDB90003AAFED1FFFC" LSID="urn:lsid:plazi:treatment:03836047994DFFDDB90003AAFED1FFFC" httpUri="http://treatment.plazi.org/id/03836047994DFFDDB90003AAFED1FFFC" lastPageId="34" lastPageNumber="34" pageId="32" pageNumber="33">
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<subSubSection id="C33082DA994DFFDFB90003AAFDADFF91" box="[318,588,144,170]" pageId="32" pageNumber="33" type="nomenclature">
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<paragraph id="8B95D151994DFFDFB90003AAFDADFF91" blockId="32.[318,588,144,170]" box="[318,588,144,170]" pageId="32" pageNumber="33">
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<heading id="D0DD663D994DFFDFB90003AAFDADFF91" box="[318,588,144,170]" centered="true" fontSize="9" level="2" pageId="32" pageNumber="33" reason="2">
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<emphasis id="B95E0D43994DFFDFB90003AAFDADFF91" bold="true" box="[318,588,144,170]" pageId="32" pageNumber="33">
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Is
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<taxonomicName id="4C2AAAD2994DFFDFB96B03AAFE5BFF91" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[341,442,144,170]" family="Tyrannosauridae" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
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<emphasis id="B95E0D43994DFFDFB96B03AAFE5BFF91" bold="true" box="[341,442,144,170]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
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</taxonomicName>
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a valid taxon?
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</emphasis>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="C33082DA994DFFDDB8BF038CFED1FFFC" lastPageId="34" lastPageNumber="35" pageId="32" pageNumber="33" type="discussion">
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<paragraph id="8B95D151994DFFDFB8BF038CFD34FDBE" blockId="32.[129,778,182,833]" pageId="32" pageNumber="33">
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Osmólska (1996) listed eight diagnostic features for
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<taxonomicName id="4C2AAAD2994DFFDFBAA1038DFCE3FFF5" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[671,770,183,206]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
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<emphasis id="B95E0D43994DFFDFBAA1038DFCE3FFF5" box="[671,770,183,206]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
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</taxonomicName>
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: (i) two surangular foramina [also considered by Holtz (2004) as an autapomorphy of
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<taxonomicName id="4C2AAAD2994DFFDFB9BE03CCFE07FE36" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[384,486,246,269]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
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<emphasis id="B95E0D43994DFFDFB9BE03CCFE07FE36" box="[384,486,246,269]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
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</taxonomicName>
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]; (ii) articular with an oblique posterior surface and a short retroarticular surface; (iii) caudal vertebrae with thin-walled centra; (iv) hyposphenehypantrum articulations in at least the first 16 caudal vertebrae [also listed by Holtz (2004) as an autapomorphy of
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<taxonomicName id="4C2AAAD2994DFFDFBAAB0249FD16FEB1" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[661,759,371,394]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
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<emphasis id="B95E0D43994DFFDFBAAB0249FD16FEB1" box="[661,759,371,394]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
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</taxonomicName>
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]; (v) prezygapophyses in proximal caudal vertebrae with ridges on the lateral surfaces; (vi) ilium with two deep depressions; (vii) femur with the anterior trochanter (anterior crest
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<emphasis id="B95E0D43994DFFDFBAE802EBFCE8FED3" box="[726,777,465,488]" italics="true" pageId="32" pageNumber="33">sensu</emphasis>
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Osmólska); and (viii) tibia and fibula fused distally. The status of those features is briefly discussed below. Given that we have now re-identified the hindlimb bones as belonging to other non-tyrannosaurid taxa, those features regarding the hindlimb were discussed above, hence they will be omitted in this section.
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</paragraph>
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<paragraph id="8B95D151994DFFDFB8A201B6FBABFFDD" blockId="32.[129,778,182,833]" lastBlockId="32.[825,1475,144,826]" pageId="32" pageNumber="33">
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The ilium with a distinct ridge on the lateral surface of the postacetabular process, demarcated anteriorly, medially, and posteriorly by depressions, is striking (
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<figureCitation id="1311CDD4994DFFDFBA2B01F1FDBDFDD8" box="[533,604,715,739]" captionStart="Figure 16" captionStartId="20.[129,194,1516,1540]" captionTargetBox="[131,1471,146,1486]" captionTargetId="figure-232@20.[129,1473,144,1488]" captionTargetPageId="20" captionText="Figure 16. Ilium of Bagaraatan ostromi ZPAL MgD-I/108. A, B, fragment of left preacetabular process in lateral (A) and medial (B) view. C–F, left postacetabular process in lateral (C), medial (D), dorsal (E), and ventral (F) view.G–J, right postacetabular process in dorsal (G), ventral (H), lateral (I), and medial (J) view.K–N, two fragments of ilium blade in lateral (K, M) and medial (L, N) view." figureDoi="http://doi.org/10.5281/zenodo.14284263" httpUri="https://zenodo.org/record/14284263/files/figure.png" pageId="32" pageNumber="33">Fig. 16</figureCitation>
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). It occurs symmetrically on both ilia, and better preserved on the left, which is less compressed. Such ridges are not found in other theropods, to our knowledge, and are not present in juvenile
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<emphasis id="B95E0D43994DFFDFBAD40010FCEBFC7A" box="[746,778,810,833]" italics="true" pageId="32" pageNumber="33">Ta.</emphasis>
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<taxonomicName id="4C2AAAD2994DFFDFBB0703AAFC9BFF93" box="[825,890,144,168]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="bataar">
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<emphasis id="B95E0D43994DFFDFBB0703AAFC9BFF93" box="[825,890,144,168]" italics="true" pageId="32" pageNumber="33">bataar</emphasis>
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</taxonomicName>
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(MPC-D 107/7) nor the other tyrannosaurid juvenile,
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<emphasis id="B95E0D43994DFFDFBD9203ABFA22FF93" box="[1452,1475,145,168]" italics="true" pageId="32" pageNumber="33">R.</emphasis>
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<taxonomicName id="4C2AAAD2994DFFDFBB070395FB8EFFFC" authority="(Sereno et al. 2009)" baseAuthorityName="Sereno" baseAuthorityYear="2009" box="[825,1135,175,199]" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="kriegsteini">
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<emphasis id="B95E0D43994DFFDFBB070395FC7AFFFC" box="[825,923,175,199]" italics="true" pageId="32" pageNumber="33">kriegsteini</emphasis>
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(Sereno
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<emphasis id="B95E0D43994DFFDFBBC4038AFBC8FFFC" box="[1018,1065,175,199]" italics="true" pageId="32" pageNumber="33">et al.</emphasis>
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2009)
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</taxonomicName>
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. Therefore, they might be a diagnostic feature of
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<taxonomicName id="4C2AAAD2994DFFDFBBDC03F5FBA2FFDD" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[994,1091,207,230]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
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<emphasis id="B95E0D43994DFFDFBBDC03F5FBA2FFDD" box="[994,1091,207,230]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
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</taxonomicName>
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.
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</paragraph>
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<paragraph id="8B95D151994DFFDEBB6B03D4FEACFE3D" blockId="32.[825,1475,144,826]" lastBlockId="33.[113,766,144,1985]" lastPageId="33" lastPageNumber="34" pageId="32" pageNumber="33">
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The ridges on the lateral surfaces of the prezygapophyses are found also in the proximal caudal vertebrae of
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<emphasis id="B95E0D43994DFFDFBD780234FA6EFE1E" box="[1350,1423,270,293]" italics="true" pageId="32" pageNumber="33">
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<taxonomicName id="4C2AAAD2994DFFDFBD780234FA63FE1E" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[1350,1410,270,293]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="rex">Ty. rex</taxonomicName>
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&
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</emphasis>
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running from the anterior margin of the transverse process to the prezygapopysis (
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<bibRefCitation id="EFBBACA0994DFFDFBC1F0276FB51FE5F" author="Brochu CA" box="[1057,1200,332,356]" pageId="32" pageNumber="33" pagination="1 - 138" refId="ref25263" refString="Brochu CA. Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleonotology 2003; 7: 1 - 138. https: // doi. org / 10.2307 / 3889334" year="2003">Brochu 2003</bibRefCitation>
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). Similar ridges on the prezygapophyses of anterior caudal vertebrae are also present in
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<taxonomicName id="4C2AAAD2994DFFDFBBB702B1FC10FE99" box="[905,1009,394,418]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="bataar">
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<emphasis id="B95E0D43994DFFDFBBB702B1FC10FE99" box="[905,1009,394,418]" italics="true" pageId="32" pageNumber="33">Ta. bataar</emphasis>
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</taxonomicName>
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(ZPAL MgD-I/176). Osmólska (1996) did not quantify how thin-walled the caudal vertebrae centra of
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<taxonomicName id="4C2AAAD2994DFFDFBD900290FC60FEDA" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
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<emphasis id="B95E0D43994DFFDFBD900290FC60FEDA" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
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</taxonomicName>
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are in comparison to other theropods. We do not recognize any clear difference between the centrum thickness of
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<taxonomicName id="4C2AAAD2994DFFDFBD9002D3FC60FD24" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
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<emphasis id="B95E0D43994DFFDFBD9002D3FC60FD24" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
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</taxonomicName>
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and other theropods. The stout hyposphene–hypantrum articulations in at least the first 16 caudal vertebrae were considered an autapomorphy of
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<taxonomicName id="4C2AAAD2994DFFDFBC62017DFB5CFD65" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1116,1213,583,606]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="ostromi">
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<emphasis id="B95E0D43994DFFDFBC62017DFB5CFD65" box="[1116,1213,583,606]" italics="true" pageId="32" pageNumber="33">B. ostromi</emphasis>
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</taxonomicName>
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by Osmólska (1996) and Holtz (2004). The presence of the hyposphene–hypantrum articulation is seen in many archosaurs, is strongly correlated with body size, and is often already present at a young age, before the articulation is necessary to support the large mass of the fully grown animal (Stefanic and
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<bibRefCitation id="EFBBACA0994DFFDFBC5B01D9FB0CFDC0" author="Nesbitt S & Loewen M" box="[1125,1261,739,763]" pageId="32" pageNumber="33" pagination="892 - 9" refId="ref28234" refString="Nesbitt S, Denton RK, Loewen M et al. A mid-Cretaceous tyrannosauroid and the origin of North American end-Cretaceous dinosaur assemblages. Nature Ecology & Evolution 2019; 3: 892 - 9." year="2019">Nesbitt 2019</bibRefCitation>
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). The hyposphenehypantrum articulations are common in theropods, and for instance, are present in the caudal vertebrae of medium-sized
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<emphasis id="B95E0D43994CFFDEB84F03ABFF3EFF93" box="[113,223,144,168]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
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(ZPAL MgD-I/176). The oblique posterior surface and short retroarticular surface of the articular, also listed by Osmólska (1996), are tyrannosauroid synapomorphies (
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<bibRefCitation id="EFBBACA0994CFFDEB84203D4FEDCFE3D" author="Brusatte SL & Lloyd GT & Wang SC" box="[124,317,238,262]" pageId="33" pageNumber="34" pagination="2386 - 92" refId="ref25593" refString="Brusatte SL, Lloyd GT, Wang SC et al. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Current Biology: CB 2014; 24: 2386 - 92. https: // doi. org / 10.1016 / j. cub. 2014.08.034" year="2014">
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Brusatte
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<emphasis id="B95E0D43994CFFDEB8E803D5FEE4FE3D" box="[214,261,238,262]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
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2014
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</bibRefCitation>
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||
).
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</paragraph>
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<caption id="DF5581D9994DFFDFB8BF04BEFDB4F883" ID-DOI="http://doi.org/10.5281/zenodo.14284283" ID-Zenodo-Dep="14284283" httpUri="https://zenodo.org/record/14284283/files/figure.png" pageId="32" pageNumber="33" startId="32.[129,194,1924,1948]" targetBox="[136,1471,887,1868]" targetPageId="32">
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<paragraph id="8B95D151994DFFDFB8BF04BEFDB4F883" blockId="32.[129,1409,1924,1976]" pageId="32" pageNumber="33">
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<emphasis id="B95E0D43994DFFDFB8BF04BEFF04F8A7" bold="true" box="[129,229,1924,1948]" pageId="32" pageNumber="33">Figure 25.</emphasis>
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Reconstruction of
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<emphasis id="B95E0D43994DFFDFB9A404BFFD1BF8A6" box="[410,762,1925,1949]" italics="true" pageId="32" pageNumber="33">Bagaraatan ostromi ZPAL MgD-I/108</emphasis>
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including only tyrannosaurid bones found in the assemblage and life reconstruction of the dinosaur by Jakub Zalewski.
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</paragraph>
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</caption>
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<paragraph id="8B95D151994CFFDEB8B30234FD49FC62" blockId="33.[113,766,144,1985]" pageId="33" pageNumber="34">
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The presence of two surangular foramina and the ridge on the lateral surface of the postacetabular process of ilium seem to be the only two features listed by Osmólska (1996) that distinguish
|
||
<taxonomicName id="4C2AAAD2994CFFDEB8910256FEF3FEB8" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[175,274,364,387]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
|
||
<emphasis id="B95E0D43994CFFDEB8910256FEF3FEB8" box="[175,274,364,387]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
|
||
</taxonomicName>
|
||
from other tyrannosaurids. The two surangular foramina were also later listed by Holtz (2004) as unique for
|
||
<taxonomicName id="4C2AAAD2994CFFDEBAD802B1FF58FEF9" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
|
||
<emphasis id="B95E0D43994CFFDEBAD802B1FF58FEF9" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
|
||
</taxonomicName>
|
||
in comparison to other theropods (
|
||
<bibRefCitation id="EFBBACA0994CFFDEBA0D0290FD08FEF9" author="Currie PJ & Hurum JH & Sabath K" box="[563,745,426,450]" pageId="33" pageNumber="34" pagination="227 - 34" refId="ref26465" refString="Currie PJ, Hurum JH, Sabath K. Skull structure and evolution in tyrannosaurid dinosaurs. Acta Palaeontologica Polonica 2003; 48: 227 - 34." year="2003">
|
||
Currie
|
||
<emphasis id="B95E0D43994CFFDEBA400291FD4EFEF9" box="[638,687,426,450]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
|
||
2003
|
||
</bibRefCitation>
|
||
). There is some confusion in the literature about the size of the surangular foramen in tyrannosauroids and its phylogenetic significance and ontogenetic and individual variation. In their phylogenetic dataset of tyrannosauroids,
|
||
<bibRefCitation id="EFBBACA0994CFFDEBA0B0112FF5DFD65" author="Brusatte SL & Carr TD" pageId="33" pageNumber="34" refId="ref25346" refString="Brusatte SL, Carr TD. The phylogeny and evolutionary history of tyrannosauroid dinosaurs. Scientific Reports 2016; 6: 20252." year="2016">Brusatte and Carr (2016)</bibRefCitation>
|
||
used a character that simply divided the size of the foramen into two states: those with a dorsoventral depth <30% or> 30% of the depth of the posterior end of the surangular. This was based on earlier characters used by Sereno
|
||
<emphasis id="B95E0D43994CFFDEBA78019FFD95FD87" box="[582,628,676,700]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
|
||
(2009), Carr and Williamson (2010), and
|
||
<bibRefCitation id="EFBBACA0994CFFDEB9A901FEFD8CFDE7" author="Brusatte SL & Norell MA & Carr TD" box="[407,621,708,732]" pageId="33" pageNumber="34" pagination="1481 - 5" refId="ref25425" refString="Brusatte SL, Norell MA, Carr TD et al. Tyrannosaur paleobiology: new research on ancient exemplar organisms. Science 2010; 329: 1481 - 5. https: // doi. org / 10.1126 / science. 1193304" year="2010">
|
||
Brusatte
|
||
<emphasis id="B95E0D43994CFFDEB9CE01FFFDFFFDE7" box="[496,542,708,732]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
|
||
(2010)
|
||
</bibRefCitation>
|
||
. The enlarged condition was found to be synapomorphic of a clade consisting of
|
||
<taxonomicName id="4C2AAAD2994CFFDEB8A40039FEFEFC21" authorityName="Marsh" authorityYear="1877" box="[154,287,771,794]" class="Reptilia" family="Dryptosauridae" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95E0D43994CFFDEB8A40039FEFEFC21" box="[154,287,771,794]" italics="true" pageId="33" pageNumber="34">Dryptosaurus</emphasis>
|
||
</taxonomicName>
|
||
+ Tyrannosauridae, whereas the primitive smaller foramen is seen in more basal tyrannosauroids, such as
|
||
<taxonomicName id="4C2AAAD2994CFFDEB84F007BFF1BFC62" box="[113,250,833,857]" genus="Suskityrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95E0D43994CFFDEB84F007BFF1BFC62" box="[113,250,833,857]" italics="true" pageId="33" pageNumber="34">Suskityrannus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C2AAAD2994CFFDEB93B0078FE94FC62" authorityName="Hutt, Naish, Martill, Barker & Newbery" authorityYear="2001" box="[261,373,834,857]" class="Reptilia" genus="Eotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95E0D43994CFFDEB93B0078FE94FC62" box="[261,373,834,857]" italics="true" pageId="33" pageNumber="34">Eotyrannus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C2AAAD2994CFFDEB941007BFE20FC62" box="[383,449,833,857]" class="Reptilia" family="Tyrannosauridae" genus="Dilong" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95E0D43994CFFDEB941007BFE20FC62" box="[383,449,833,857]" italics="true" pageId="33" pageNumber="34">Dilong</emphasis>
|
||
</taxonomicName>
|
||
, and proceratosaurids.
|
||
</paragraph>
|
||
<paragraph id="8B95D151994CFFDEB8B3005BFD86FA55" blockId="33.[113,766,144,1985]" pageId="33" pageNumber="34">
|
||
Other authors, however, have considered the foramen differently. The size of the surangular foramen in tyrannosaurids was divided into moderate (
|
||
<taxonomicName id="4C2AAAD2994CFFDEB98C009AFDCCFC8C" box="[434,557,927,951]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis id="B95E0D43994CFFDEB98C009AFDCCFC8C" box="[434,557,927,951]" italics="true" pageId="33" pageNumber="34">Go. libratus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C2AAAD2994CFFDEBA7E00A5FE88FCED" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="sarcophagus">
|
||
<emphasis id="B95E0D43994CFFDEBA7E00A5FF2BFCEC" italics="true" pageId="33" pageNumber="34">Albertosaurus sarcophagus</emphasis>
|
||
<bibRefCitation id="EFBBACA0994CFFDEB8E80085FE88FCED" author="Osborn HF" box="[214,361,958,983]" pageId="33" pageNumber="34" pagination="259 - 65" refId="ref28311" refString="Osborn HF. Tyrannosaurus and other Cretaceous carnivorous dinosaurs. Bulletin of the American Museum of Natural History 1905; 21: 259 - 65." year="1905">Osborn, 1905</bibRefCitation>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C2AAAD2994CFFDEB9470085FE5EFCED" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[377,447,959,982]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B95E0D43994CFFDEB9470085FE5EFCED" box="[377,447,959,982]" italics="true" pageId="33" pageNumber="34">Ty. rex</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C2AAAD2994CFFDEBA3F0085FD8DFCEC" box="[513,620,959,983]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis id="B95E0D43994CFFDEBA3F0085FD8DFCEC" box="[513,620,959,983]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
|
||
</taxonomicName>
|
||
) or enlarged (
|
||
<taxonomicName id="4C2AAAD2994CFFDEB84200E5FF07FCCD" box="[124,230,990,1014]" class="Reptilia" genus="Bistahieversor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="sealeyi">
|
||
<emphasis id="B95E0D43994CFFDEB84200E5FF07FCCD" box="[124,230,990,1014]" italics="true" pageId="33" pageNumber="34">Bi. sealeyi</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C2AAAD2994CFFDEB8C400E4FE28FCCE" box="[250,457,990,1014]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="undetermined">
|
||
<emphasis id="B95E0D43994CFFDEB8C400E4FE6FFCCD" box="[250,398,990,1014]" italics="true" pageId="33" pageNumber="34">Daspletosaurus</emphasis>
|
||
spp.
|
||
</taxonomicName>
|
||
,
|
||
<emphasis id="B95E0D43994CFFDEB9E500E5FE18FCCD" box="[475,505,991,1014]" italics="true" pageId="33" pageNumber="34">Te.</emphasis>
|
||
<taxonomicName id="4C2AAAD2994CFFDEBA3900E5FD1BFCCD" authority=", Nanatotheristes" authorityName="Nanatotheristes" box="[519,762,990,1014]" class="Reptilia" genus="Teratophoneus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Avetheropoda" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="curriei">
|
||
<emphasis id="B95E0D43994CFFDEBA3900E5FDA4FCCD" box="[519,581,991,1014]" italics="true" pageId="33" pageNumber="34">curriei</emphasis>
|
||
,
|
||
<emphasis id="B95E0D43994CFFDEBA6400E4FD1BFCCD" box="[602,762,990,1014]" italics="true" pageId="33" pageNumber="34">Ŋanatotheristes</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B95E0D43994CFFDEB84F00C7FF0CFB2E" box="[113,237,1021,1045]" italics="true" pageId="33" pageNumber="34">degrootorum</emphasis>
|
||
Voris
|
||
<emphasis id="B95E0D43994CFFDEB90D00C4FE84FB2E" box="[307,357,1021,1045]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
|
||
, 2020,
|
||
<taxonomicName id="4C2AAAD2994CFFDEB98D00C4FDFBFB2E" box="[435,538,1022,1045]" family="Tyrannosauridae" genus="Qianzhousaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="sinensis">
|
||
<emphasis id="B95E0D43994CFFDEB98D00C4FDFBFB2E" box="[435,538,1022,1045]" italics="true" pageId="33" pageNumber="34">Q. sinensis</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C2AAAD2994CFFDEBA6800C7FD0EFB2E" box="[598,751,1021,1045]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="altai">
|
||
<emphasis id="B95E0D43994CFFDEBA6800C7FD0EFB2E" box="[598,751,1021,1045]" italics="true" pageId="33" pageNumber="34">Alioramus altai</emphasis>
|
||
</taxonomicName>
|
||
) states by Voris
|
||
<emphasis id="B95E0D43994CFFDEB92E0727FEA0FB0E" box="[272,321,1053,1077]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
|
||
(2021). However, the surangular foramen in
|
||
<taxonomicName id="4C2AAAD2994CFFDEB8B50707FF2AFB6F" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[139,203,1085,1108]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B95E0D43994CFFDEB8B50707FF2AFB6F" box="[139,203,1085,1108]" italics="true" pageId="33" pageNumber="34">Ty.rex</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C2AAAD2994CFFDEB8C20706FE1FFB6F" authorityName="Osborn" authorityYear="1905" box="[252,510,1084,1108]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="sarcophagus">
|
||
<emphasis id="B95E0D43994CFFDEB8C20706FE1FFB6F" box="[252,510,1084,1108]" italics="true" pageId="33" pageNumber="34">Albertosaurus sarcophagus</emphasis>
|
||
</taxonomicName>
|
||
also used to be classified as smaller than in other tyrannosaurids (
|
||
<bibRefCitation id="EFBBACA0994CFFDEBA210761FF44FBA8" author="Carr TD & Williamson TE" pageId="33" pageNumber="34" pagination="479 - 523" refId="ref25757" refString="Carr TD, Williamson TE. Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America. Zoological Journal of the Linnean Society 2004; 142: 479 - 523. https: // doi. org / 10.1111 / j. 1096 - 3642.2004.00130. x" year="2004">Carr and Williamson 2004</bibRefCitation>
|
||
). Other authors reported that the surangular foramen in
|
||
<taxonomicName id="4C2AAAD2994CFFDEB84F07A1FF39FB89" box="[113,216,1178,1202]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis id="B95E0D43994CFFDEB84F07A1FF39FB89" box="[113,216,1178,1202]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
|
||
</taxonomicName>
|
||
is smaller than in other tyrannosaurids and invariant during ontogeny (Tsuihiji
|
||
<emphasis id="B95E0D43994CFFDEB9420780FE4BFBEA" box="[380,426,1209,1233]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
|
||
2011, Voris
|
||
<emphasis id="B95E0D43994CFFDEBA180780FDB5FBEA" box="[550,596,1209,1233]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
|
||
2021). Also, the surangular foramen in ‘
|
||
<taxonomicName id="4C2AAAD2994CFFDEB95307E2FD25FBCB" authorityName="' Dong" authorityYear="1977" box="[365,708,1240,1264]" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="huoyanshanensis">
|
||
<emphasis id="B95E0D43994CFFDEB95307E2FD25FBCB" box="[365,708,1240,1264]" italics="true" pageId="33" pageNumber="34">Shanshanosaurus huoyanshanensis</emphasis>
|
||
</taxonomicName>
|
||
’ was described as large (
|
||
<bibRefCitation id="EFBBACA0994CFFDEB90007C2FDCFFA2B" author="Currie PJ & Dong ZM" box="[318,558,1272,1296]" pageId="33" pageNumber="34" pagination="1729 - 37" refId="ref26336" refString="Currie PJ, Dong ZM. New information on Shanshanosaurus huoyanshanensis, a juvenile tyronnsaurid (Theropoda, Dinosauria) from the Late Cretaceous of China. Canadian Journal of Earth Sciences 2001; 38: 1729 - 37." year="2001">Currie and Dong 2001</bibRefCitation>
|
||
), but later as small (Tsuihiji
|
||
<emphasis id="B95E0D43994CFFDEB8EE0622FEE0FA14" box="[208,257,1303,1327]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
|
||
2011). For
|
||
<taxonomicName id="4C2AAAD2994CFFDEB9430622FE5EFA14" baseAuthorityName="Carr" baseAuthorityYear="2020" box="[381,447,1304,1327]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B95E0D43994CFFDEB9430622FE5EFA14" box="[381,447,1304,1327]" italics="true" pageId="33" pageNumber="34">Ty. rex</emphasis>
|
||
</taxonomicName>
|
||
, the size of the surangular foramen was first reported as increasing (
|
||
<bibRefCitation id="EFBBACA0994CFFDEBA39060DFD93FA74" author="Carr TD" box="[519,626,1335,1359]" pageId="33" pageNumber="34" pagination="497 - 520" refId="ref25647" refString="Carr TD. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology 1999; 19: 497 - 520. https: // doi. org / 10.1080 / 02724634.1999.10011161" year="1999">Carr 1999</bibRefCitation>
|
||
) but later as decreasing in size through ontogeny (
|
||
<bibRefCitation id="EFBBACA0994CFFDEB9D0066CFDB7FA55" author="Carr TD" box="[494,598,1366,1390]" pageId="33" pageNumber="34" refId="ref25682" refString="Carr TD. A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence. PeerJ 2020; 8: e 9192. https: // doi. org / 10.7717 / peerj. 9192" year="2020">Carr 2020</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8B95D151994CFFDEB8B3064CFC8EFC62" blockId="33.[113,766,144,1985]" lastBlockId="33.[810,1460,144,1986]" pageId="33" pageNumber="34">
|
||
Because of this confusion, we built a dataset to examine the size of foramina in a quantitative context. In tyrannosaurids, growth of the mandible, skull, and femur is isometric and related to the body size of the individual (
|
||
<bibRefCitation id="EFBBACA0994CFFDEBA2006E9FD4BFAD0" author="Currie PJ" box="[542,682,1491,1515]" pageId="33" pageNumber="34" pagination="651 - 65" refId="ref26287" refString="Currie PJ. Allometric growth in tyrannosaurids (Dinosauria: Theropoda) from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences 2003 b; 40: 651 - 65. https: // doi. org / 10.1139 / e 02 - 083" year="2003">Currie 2003b</bibRefCitation>
|
||
). Thus, we assessed the relationship between the size of the surangular foramen and skull length (as a proxy for body size). Our results (
|
||
<figureCitation id="1311CDD4994CFFDEB842050BFF20F972" box="[124,193,1585,1609]" captionStart="Figure 22" captionStartId="29.[113,178,818,842]" captionTargetBox="[117,1455,147,788]" captionTargetId="figure-606@29.[114,1458,144,791]" captionTargetPageId="29" captionText="Figure 22. Plots showing the relationship between the anteroposterior length of the surangular foramen and the skull length (as a proxy of body size) in tyrannosaurids.Bagaraatan ostromi was measured for the single (posterior only) or double (posterior + anterior) foramina. A, regression analysis including all specimens shows a negative slope and a high correlation coefficient. B, regression analysis for specific taxa shows the same trend. No distinction between small, medium, or large size of the surangular foramen can be noticed in the tyrannosaurids." figureDoi="http://doi.org/10.5281/zenodo.14284277" httpUri="https://zenodo.org/record/14284277/files/figure.png" pageId="33" pageNumber="34">Fig. 22</figureCitation>
|
||
) show that in all taxa the size of the surangular foramen decreases during ontogeny (negative allometry) and is strongly correlated with the length of the skull. Thus, e.g.
|
||
<taxonomicName id="4C2AAAD2994CFFDEBAA8054AFF7CF99C" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="altai">
|
||
<emphasis id="B95E0D43994CFFDEBAA8054AFF7CF99C" italics="true" pageId="33" pageNumber="34">Alioramus altai</emphasis>
|
||
</taxonomicName>
|
||
(IGM 100/1844) and the similar-sized
|
||
<taxonomicName id="4C2AAAD2994CFFDEBA0005B5FD50F99C" box="[574,689,1679,1703]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis id="B95E0D43994CFFDEBA0005B5FD50F99C" box="[574,689,1679,1703]" italics="true" pageId="33" pageNumber="34">Go. libratus</emphasis>
|
||
</taxonomicName>
|
||
(TMP 1991.36.500) have surangular foramina of proportionally the same size. Although the surangular foramen–skull length correlation is statistically significant, variability in surangular foramen size is also apparent, especially in
|
||
<taxonomicName id="4C2AAAD2994CFFDEB9F70437FDDAF81F" box="[457,571,1804,1828]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis id="B95E0D43994CFFDEB9F70437FDDAF81F" box="[457,571,1804,1828]" italics="true" pageId="33" pageNumber="34">Go. libratus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C2AAAD2994CFFDEBA520437FD33F81F" box="[620,722,1804,1828]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis id="B95E0D43994CFFDEBA520437FD33F81F" box="[620,722,1804,1828]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
|
||
</taxonomicName>
|
||
, for which the data are less fitted to the trend than for the other species (
|
||
<emphasis id="B95E0D43994CFFDEB8990476FF56F858" box="[167,183,1868,1891]" italics="true" pageId="33" pageNumber="34">
|
||
<collectionCode id="ED3B4994994CFFDEB8990476FF56F858" box="[167,183,1868,1891]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="33" pageNumber="34">R</collectionCode>
|
||
</emphasis>
|
||
<superScript id="7C5F7C19994CFFDEB8890470FF5EF863" attach="left" box="[183,191,1866,1880]" fontSize="6" pageId="33" pageNumber="34">2</superScript>
|
||
=.76–.78, vs.
|
||
<emphasis id="B95E0D43994CFFDEB96C0476FE81F858" box="[338,352,1868,1891]" italics="true" pageId="33" pageNumber="34">
|
||
<collectionCode id="ED3B4994994CFFDEB96C0476FE81F858" box="[338,352,1868,1891]" country="Chile" name="Departamento de Geologia, Universidad de Chile" pageId="33" pageNumber="34">R</collectionCode>
|
||
</emphasis>
|
||
<superScript id="7C5F7C19994CFFDEB95E0470FE89F863" attach="left" box="[352,360,1866,1880]" fontSize="6" pageId="33" pageNumber="34">2</superScript>
|
||
>.88–.
|
||
<quantity id="4CD27CB4994CFFDEB9880471FE0AF858" box="[438,491,1867,1891]" metricMagnitude="0" metricUnit="m" metricValue="2.3876" pageId="33" pageNumber="34" unit="in" value="94.0">94 in</quantity>
|
||
<taxonomicName id="4C2AAAD2994CFFDEB9D10471FD50F858" box="[495,689,1867,1891]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="undetermined">
|
||
<emphasis id="B95E0D43994CFFDEB9D10471FD62F858" box="[495,643,1867,1891]" italics="true" pageId="33" pageNumber="34">Daspletosaurus</emphasis>
|
||
spp.
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C2AAAD2994CFFDEBAE10476FF6EF8B9" baseAuthorityName="Carr" baseAuthorityYear="2020" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B95E0D43994CFFDEBAE10476FF6EF8B9" italics="true" pageId="33" pageNumber="34">Ty. rex</emphasis>
|
||
</taxonomicName>
|
||
;
|
||
<figureCitation id="1311CDD4994CFFDEB8A00451FF05F8B8" box="[158,228,1899,1923]" captionStart="Figure 22" captionStartId="29.[113,178,818,842]" captionTargetBox="[117,1455,147,788]" captionTargetId="figure-606@29.[114,1458,144,791]" captionTargetPageId="29" captionText="Figure 22. Plots showing the relationship between the anteroposterior length of the surangular foramen and the skull length (as a proxy of body size) in tyrannosaurids.Bagaraatan ostromi was measured for the single (posterior only) or double (posterior + anterior) foramina. A, regression analysis including all specimens shows a negative slope and a high correlation coefficient. B, regression analysis for specific taxa shows the same trend. No distinction between small, medium, or large size of the surangular foramen can be noticed in the tyrannosaurids." figureDoi="http://doi.org/10.5281/zenodo.14284277" httpUri="https://zenodo.org/record/14284277/files/figure.png" pageId="33" pageNumber="34">Fig. 22</figureCitation>
|
||
). Indeed, although the surangular foramen is rather enlarged in
|
||
<taxonomicName id="4C2AAAD2994CFFDEB8D704B0FE83F899" authorityName="Maleev" authorityYear="1955" box="[233,354,1930,1954]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95E0D43994CFFDEB8D704B0FE83F899" box="[233,354,1930,1954]" italics="true" pageId="33" pageNumber="34">Tarbosaurus</emphasis>
|
||
</taxonomicName>
|
||
individuals (as in other tyrannosaurids; Fig. 22) bigger than MPC-D 100/66 (skull length:
|
||
<quantity id="4CD27CB4994CFFDEBABF0493FD25F8FA" box="[641,708,1961,1985]" metricMagnitude="-1" metricUnit="m" metricValue="4.5" pageId="33" pageNumber="34" unit="cm" value="45.0">45 cm</quantity>
|
||
), few exceptions were found within the hypodigm. The surangular foramen of the medium-sized specimen MPC-D 107/14 is exceptionally small in comparison to other
|
||
<taxonomicName id="4C2AAAD2994CFFDEBCEA03F5FADAFFDC" box="[1236,1339,207,231]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis id="B95E0D43994CFFDEBCEA03F5FADAFFDC" box="[1236,1339,207,231]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
|
||
</taxonomicName>
|
||
individuals of similar size (e.g. ZPAL MgD-I/3 and MPC-D 107/5;
|
||
<figureCitation id="1311CDD4994CFFDEBD5403D4FA43FE3D" box="[1386,1442,238,262]" captionStart="Figure 5" captionStartId="8.[129,194,1539,1563]" captionTargetBox="[131,1471,146,1509]" captionTargetId="figure-205@8.[129,1473,144,1511]" captionTargetPageId="8" captionText="Figure 5. Comparison of Bagaraatan ostromi (A) and Tarbosaurus bataar (B–F) distal end of the mandible in lateral view.A, ZPAL MgD-I/108. B, ZPAL MgD-I/31. C, ZPAL MgD-I/3. D, MPC-D 107/14. E, ZPAL MgD-I/4. F, ZPAL MgD-I/5. Note the exceptionally small surangular foramen in the medium-sized individual MPC-D 107/14 (D) in comparison to similar-sized ZPAL MgD-I/3 (C). Asterisks indicate the posterior surangular foramen.Note that the placement of the double surangular foramina in B.ostromi is similar to the enlarged posterior surangular foramen in Ta.bataar. During ontogeny, the surangular shelf faces more laterally and the glenoid becomes longer anteroposteriorly." figureDoi="http://doi.org/10.5281/zenodo.14284239" httpUri="https://zenodo.org/record/14284239/files/figure.png" pageId="33" pageNumber="34">Fig. 5</figureCitation>
|
||
). Moreover, a specimen larger than those listed above, MPC-D 100/60, shows asymmetrical surangular foramina: the left one is smaller (anteroposterior length:
|
||
<quantity id="4CD27CB4994CFFDEBCA40276FB09FE5F" box="[1178,1256,332,356]" metricMagnitude="-2" metricUnit="m" metricValue="2.3" pageId="33" pageNumber="34" unit="mm" value="23.0">23 mm</quantity>
|
||
) and the right one larger (anteroposterior length:
|
||
<quantity id="4CD27CB4994CFFDEBC570251FB56FEB8" box="[1129,1207,363,387]" metricMagnitude="-2" metricUnit="m" metricValue="4.0" pageId="33" pageNumber="34" unit="mm" value="40.0">40 mm</quantity>
|
||
). The smaller surangular foramen of the left mandible can be noticed as an outlier in the
|
||
<figureCitation id="1311CDD4994CFFDEBB6D0290FC56FEF9" box="[851,951,426,450]" captionStart="Figure 22" captionStartId="29.[113,178,818,842]" captionTargetBox="[117,1455,147,788]" captionTargetId="figure-606@29.[114,1458,144,791]" captionTargetPageId="29" captionText="Figure 22. Plots showing the relationship between the anteroposterior length of the surangular foramen and the skull length (as a proxy of body size) in tyrannosaurids.Bagaraatan ostromi was measured for the single (posterior only) or double (posterior + anterior) foramina. A, regression analysis including all specimens shows a negative slope and a high correlation coefficient. B, regression analysis for specific taxa shows the same trend. No distinction between small, medium, or large size of the surangular foramen can be noticed in the tyrannosaurids." figureDoi="http://doi.org/10.5281/zenodo.14284277" httpUri="https://zenodo.org/record/14284277/files/figure.png" pageId="33" pageNumber="34">Figure 22</figureCitation>
|
||
. It would appear that there was some variability in the timing of the surangular foramen enlargement, at least in
|
||
<taxonomicName id="4C2AAAD2994CFFDEBB1402D3FC70FD3B" box="[810,913,488,512]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis id="B95E0D43994CFFDEBB1402D3FC70FD3B" box="[810,913,488,512]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
|
||
</taxonomicName>
|
||
. The size of the surangular foramen in
|
||
<taxonomicName id="4C2AAAD2994CFFDEBD1902D3FA6BFD3B" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1319,1418,489,512]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
|
||
<emphasis id="B95E0D43994CFFDEBD1902D3FA6BFD3B" box="[1319,1418,489,512]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
|
||
</taxonomicName>
|
||
, regardless of whether it is measured for the single (posterior only) or double (posterior + anterior) foramina, falls into the overall variability of surangular size in the tyrannosaurids generally and
|
||
<taxonomicName id="4C2AAAD2994CFFDEBB14015DFC6EFD45" box="[810,911,614,638]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis id="B95E0D43994CFFDEBB14015DFC6EFD45" box="[810,911,614,638]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
|
||
</taxonomicName>
|
||
specifically. The position of the surangular foramen in ‘
|
||
<taxonomicName id="4C2AAAD2994CFFDEBB0E01BFFC3BFDA6" authorityName="Dong" authorityYear="1977" box="[816,986,645,669]" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95E0D43994CFFDEBB0E01BFFC3BFDA6" box="[816,986,645,669]" italics="true" pageId="33" pageNumber="34">Shanshanosaurus</emphasis>
|
||
</taxonomicName>
|
||
’ and
|
||
<taxonomicName id="4C2AAAD2994CFFDEBC3101BCFB94FDA6" box="[1039,1141,645,669]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis id="B95E0D43994CFFDEBC3101BCFB94FDA6" box="[1039,1141,645,669]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
|
||
</taxonomicName>
|
||
MPC-D 107/7 is similar to the position of the posterior opening in the surangular of
|
||
<taxonomicName id="4C2AAAD2994CFFDEBD05019FFA4FFD87" authorityName="Osmolska" authorityYear="1996" box="[1339,1454,677,700]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95E0D43994CFFDEBD05019FFA4FFD87" box="[1339,1454,677,700]" italics="true" pageId="33" pageNumber="34">Bagaraatan</emphasis>
|
||
</taxonomicName>
|
||
, and those individuals cluster together on the plot. In turn, if the length of the area of both surangular foramina is measured for
|
||
<taxonomicName id="4C2AAAD2994CFFDEBB140039FC6CFC21" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[810,909,771,794]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
|
||
<emphasis id="B95E0D43994CFFDEBB140039FC6CFC21" box="[810,909,771,794]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
|
||
</taxonomicName>
|
||
, it clusters with
|
||
<taxonomicName id="4C2AAAD2994CFFDEBC060039FB56FC20" box="[1080,1207,771,795]" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="kriegsteini">
|
||
<emphasis id="B95E0D43994CFFDEBC060039FB56FC20" box="[1080,1207,771,795]" italics="true" pageId="33" pageNumber="34">R. kriegsteini</emphasis>
|
||
</taxonomicName>
|
||
, the surangular foramen of which was previously reported to be ‘enlarged’ (
|
||
<bibRefCitation id="EFBBACA0994CFFDEBD080018FCBFFC62" author="Fowler DW & Woodward HN & Freedman EA" pageId="33" pageNumber="34" refId="ref26728" refString="Fowler DW, Woodward HN, Freedman EA et al. Reanalysis of " Raptorex kriegsteini ": a juvenile tyrannosaurid dinosaur from Mongolia. PLoS One 2011; 6: e 21376. https: // doi. org / 10.1371 / journal. pone. 0021376" year="2011">
|
||
Fowler
|
||
<emphasis id="B95E0D43994CFFDEBDBA0019FA55FC01" box="[1412,1460,802,826]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
|
||
2011
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8B95D151994CFFDDBB7B005BFED1FFFC" blockId="33.[810,1460,144,1986]" lastBlockId="34.[129,777,144,199]" lastPageId="34" lastPageNumber="35" pageId="33" pageNumber="34">
|
||
What might explain the strange double set of foramina in
|
||
<taxonomicName id="4C2AAAD2994CFFDEBB1400BBFC7CFCA3" authority="Ne" authorityName="Osmolska" authorityYear="1996" box="[810,925,897,920]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95E0D43994CFFDEBB1400BBFC7CFCA3" box="[810,925,897,920]" italics="true" pageId="33" pageNumber="34">Bagaraatan</emphasis>
|
||
</taxonomicName>
|
||
? The bone between the anterior and posterior surangular foramina in
|
||
<taxonomicName id="4C2AAAD2994CFFDEBC20009AFB62FC8C" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1054,1155,928,951]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
|
||
<emphasis id="B95E0D43994CFFDEBC20009AFB62FC8C" box="[1054,1155,928,951]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
|
||
</taxonomicName>
|
||
is very thin, and the relative position of this area and both foramina matches the surangular foramen of
|
||
<taxonomicName id="4C2AAAD2994CFFDEBB9E00E5FBFFFCCD" box="[928,1054,990,1014]" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="kriegsteini">
|
||
<emphasis id="B95E0D43994CFFDEBB9E00E5FBFFFCCD" box="[928,1054,990,1014]" italics="true" pageId="33" pageNumber="34">R. kriegsteini</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C2AAAD2994CFFDEBC7100E5FB55FCCD" box="[1103,1204,990,1014]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis id="B95E0D43994CFFDEBC7100E5FB55FCCD" box="[1103,1204,990,1014]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
|
||
</taxonomicName>
|
||
(ZPAL MgD-I/31). The surangular in tyrannosaurids during the early years of life was invaded by a pneumatic diverticulum (
|
||
<bibRefCitation id="EFBBACA0994CFFDEBC870726FABCFB0E" author="Gold MEL & Brusatte SL & Norell MA" box="[1209,1373,1052,1077]" pageId="33" pageNumber="34" pagination="1 - 46" refId="ref27220" refString="Gold MEL, Brusatte SL, Norell MA. The cranial pneumatic sinuses of the tyrannosaurid Alioramus (Dinosauria: Theropoda) and the evolution of cranial pneumaticity in theropod dinosaurs. American Museum Novitates 2013; 3790: 1 - 46. https: // doi. org / 10.1206 / 3790.1" year="2013">
|
||
Gold
|
||
<emphasis id="B95E0D43994CFFDEBCCA0727FAC5FB0F" box="[1268,1316,1052,1076]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
|
||
2013
|
||
</bibRefCitation>
|
||
), which pneumatized the bone and formed the enlarged surangular foramen, bordered by a pneumatic pocket posterodorsal to it. Given that more basal tyrannosauroids have a small foramen without a pneumatic pocket, it is not clear whether there was any pneumatic diverticulum in this region in these species. Owing to the fact that pneumatic diverticula induce bone resorption when they contact bone (
|
||
<bibRefCitation id="EFBBACA0994CFFDEBBC607C2FB61FA2B" author="Bremer JL" box="[1016,1152,1272,1296]" pageId="33" pageNumber="34" pagination="197 - 211" refId="ref25223" refString="Bremer JL. The pneumatization of the humerus in common fowl and the associated activity of theelin. Anatomical Record 1940; 77: 197 - 211. https: // doi. org / 10.1002 / ar. 1090770209" year="1940">Bremer 1940</bibRefCitation>
|
||
, Witmer 1997,
|
||
<bibRefCitation id="EFBBACA0994CFFDEBD1807C2FA43FA2B" author="Wedel M" box="[1318,1442,1272,1296]" pageId="33" pageNumber="34" refId="ref29166" refString="Wedel M. Postcranial pneumaticity in dinosaurs and the origin of the avian lung. Unpublished PhD Thesis, University of California, 2007." type="book" year="2007">Wedel 2007</bibRefCitation>
|
||
), we propose that the mandible of
|
||
<taxonomicName id="4C2AAAD2994CFFDEBCBA0622FB09FA14" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1156,1256,1304,1327]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
|
||
<emphasis id="B95E0D43994CFFDEBCBA0622FB09FA14" box="[1156,1256,1304,1327]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
|
||
</taxonomicName>
|
||
exhibits local bone resorption, induced by the pneumatic diverticula, that would explain the extremely thin bone between the anterior and posterior foramen. We hypothesize that if the pneumatization process continued slightly longer, the two foramina might have merged into a single large foramen, which is the common condition in
|
||
<taxonomicName id="4C2AAAD2994CFFDEBBAF06EEFBF7FAD0" authorityName="Marsh" authorityYear="1877" box="[913,1046,1492,1515]" class="Reptilia" family="Dryptosauridae" genus="Dryptosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95E0D43994CFFDEBBAF06EEFBF7FAD0" box="[913,1046,1492,1515]" italics="true" pageId="33" pageNumber="34">Dryptosaurus</emphasis>
|
||
</taxonomicName>
|
||
+ Tyrannosauridae (
|
||
<bibRefCitation id="EFBBACA0994CFFDEBCCB06EEFCBFF930" author="Brusatte SL & Carr TD" pageId="33" pageNumber="34" refId="ref25346" refString="Brusatte SL, Carr TD. The phylogeny and evolutionary history of tyrannosauroid dinosaurs. Scientific Reports 2016; 6: 20252." year="2016">Brusatte and Carr 2016</bibRefCitation>
|
||
). This proposal is supported by the fact that the posterior surangular foramen in
|
||
<taxonomicName id="4C2AAAD2994CFFDEBC260529FB9CF911" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1048,1149,1555,1578]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
|
||
<emphasis id="B95E0D43994CFFDEBC260529FB9CF911" box="[1048,1149,1555,1578]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
|
||
</taxonomicName>
|
||
is similar in length and positioned in a similar place as in the smaller ‘
|
||
<taxonomicName id="4C2AAAD2994CFFDEBCE9050BFA60F972" authorityName="Dong" authorityYear="1977" box="[1239,1409,1585,1609]" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95E0D43994CFFDEBCE9050BFA60F972" box="[1239,1409,1585,1609]" italics="true" pageId="33" pageNumber="34">Shanshanosaurus</emphasis>
|
||
</taxonomicName>
|
||
’ and
|
||
<taxonomicName id="4C2AAAD2994CFFDEBB14056BFC74F952" box="[810,917,1617,1641]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis id="B95E0D43994CFFDEBB14056BFC74F952" box="[810,917,1617,1641]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
|
||
</taxonomicName>
|
||
MPC-D 107/7 (skull length ~
|
||
<quantity id="4CD27CB4994CFFDEBCD6056BFAD0F952" box="[1256,1329,1617,1641]" metricMagnitude="-1" metricUnit="m" metricValue="2.9" pageId="33" pageNumber="34" unit="cm" value="29.0">29 cm</quantity>
|
||
) specimens. Furthermore, the area of the surangular containing the posterior and anterior surangular foramina and the thinned bone between them matches the length and position of the surangular foramen in
|
||
<taxonomicName id="4C2AAAD2994CFFDEBBB905F5FC00F9DD" box="[903,993,1743,1766]" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
|
||
<emphasis id="B95E0D43994CFFDEBBB905F5FC00F9DD" box="[903,993,1743,1766]" italics="true" pageId="33" pageNumber="34">Raptorex</emphasis>
|
||
</taxonomicName>
|
||
(skull length ~
|
||
<quantity id="4CD27CB4994CFFDEBCBD05F4FB2BF9DD" box="[1155,1226,1742,1766]" metricMagnitude="-1" metricUnit="m" metricValue="3.0" pageId="33" pageNumber="34" unit="cm" value="30.0">30 cm</quantity>
|
||
). Therefore,
|
||
<taxonomicName id="4C2AAAD2994CFFDEBD7005F5FA52F9DD" authority="Osmolska, 1996" authorityName="Osmolska" authorityYear="1996" box="[1358,1459,1743,1766]" genus="Bagaraatan" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="ostromi">
|
||
<emphasis id="B95E0D43994CFFDEBD7005F5FA52F9DD" box="[1358,1459,1743,1766]" italics="true" pageId="33" pageNumber="34">B. ostromi</emphasis>
|
||
</taxonomicName>
|
||
(skull also ~
|
||
<quantity id="4CD27CB4994CFFDEBB9905D7FC08F83E" box="[935,1001,1773,1797]" metricMagnitude="-1" metricUnit="m" metricValue="3.0" pageId="33" pageNumber="34" unit="cm" value="30.0">30 cm</quantity>
|
||
long) possibly captures the precise moment of ongoing bone resorption and perforation attributable to the pneumatic diverticulum. This process probably occurred early in ontogeny, in specimens
|
||
<quantity id="4CD27CB4994CFFDEBC1A0471FB89F858" box="[1060,1128,1867,1891]" metricMagnitude="0" metricUnit="m" metricValue="2.5" metricValueMax="3.0" metricValueMin="2.0" pageId="33" pageNumber="34" unit="m" value="2.5" valueMax="3.0" valueMin="2.0">2–3 m</quantity>
|
||
long, which were probably 2–3 years old at the time of death (as indicated for
|
||
<taxonomicName id="4C2AAAD2994CFFDEBCC80451FABAF8B9" box="[1270,1371,1898,1922]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis id="B95E0D43994CFFDEBCC80451FABAF8B9" box="[1270,1371,1898,1922]" italics="true" pageId="33" pageNumber="34">Ta. bataar</emphasis>
|
||
</taxonomicName>
|
||
MPC-D 107/7 by Tsuihiji
|
||
<emphasis id="B95E0D43994CFFDEBBD604B1FBFBF899" box="[1000,1050,1930,1954]" italics="true" pageId="33" pageNumber="34">et al.</emphasis>
|
||
2011). Apparently, around this growth stage the pneumatic diverticulum invaded the bone, and thus variability in the size, shape, and even the number of foramina is to be expected.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |