927 lines
141 KiB
XML
927 lines
141 KiB
XML
<document ID-DOI="http://doi.org/10.5281/zenodo.3679538" ID-GBIF-Dataset="75f859ce-4da6-4513-871e-e1fb3b41a55f" ID-GBIF-Taxon="162147819" ID-Zenodo-Dep="3679538" checkinTime="1581972654346" checkinUser="plazi" docAuthor="Voris, Jared T., Therrien, François, Zelenitsky, Darla K. & Brown, Caleb M." docDate="2020" docId="039E87F3FFE7FFDAB92BDBF56DCEFC17" docLanguage="en" docName="createceousResearch.110.104.388.pdf" docOrigin="Cretaceous Research 110 (104388)" docStyle="DocumentStyle{}" docTitle="Thanatotheristes Voris & Therrien & Zelenitsky & Brown 2020" docType="treatment" docVersion="4" lastPageId="8" lastPageNumber="9" masterDocId="FFA7FF8BFFE5FFD2B95ADE6C695CFF97" masterDocTitle="A new tyrannosaurine (Theropoda: Tyrannosauridae) from the Campanian Foremost Formation of Alberta, Canada, provides insight into the evolution and biogeography of tyrannosaurids" masterLastPageNumber="15" masterPageNumber="1" pageId="2" pageNumber="3" updateTime="1643450941510" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>A new tyrannosaurine (Theropoda: Tyrannosauridae) from the Campanian Foremost Formation of Alberta, Canada, provides insight into the evolution and biogeography of tyrannosaurids</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Voris, Jared T.</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Therrien, François</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Zelenitsky, Darla K.</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Brown, Caleb M.</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>Cretaceous Research</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2020</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2020-01-23</mods:number>
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</mods:detail>
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<mods:detail type="volume">
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<mods:number>110</mods:number>
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</mods:detail>
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<mods:detail type="issue">
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<mods:number>104388</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>1</mods:start>
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<mods:end>15</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1016/j.cretres.2020.104388</mods:identifier>
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<mods:identifier type="GBIF-Dataset">75f859ce-4da6-4513-871e-e1fb3b41a55f</mods:identifier>
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<mods:identifier type="Zenodo-Dep">3672003</mods:identifier>
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<mods:identifier type="ZooBank">26778F25-3135-412D-A203-752CB9C078F1</mods:identifier>
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</mods:mods>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.3679538" ID-GBIF-Taxon="162147819" ID-Zenodo-Dep="3679538" LSID="urn:lsid:plazi:treatment:039E87F3FFE7FFDAB92BDBF56DCEFC17" httpUri="http://treatment.plazi.org/id/039E87F3FFE7FFDAB92BDBF56DCEFC17" lastPageId="8" lastPageNumber="9" pageId="2" pageNumber="3">
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<subSubSection pageId="2" pageNumber="3" type="nomenclature">
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<paragraph blockId="2.[113,783,1433,1509]" box="[113,595,1433,1453]" pageId="2" pageNumber="3">
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<emphasis bold="true" box="[113,432,1433,1453]" italics="true" pageId="2" pageNumber="3">
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<taxonomicName authority="Voris & Therrien & Zelenitsky & Brown, 2020" authorityName="Voris & Therrien & Zelenitsky & Brown" authorityYear="2020" box="[113,289,1433,1453]" class="Reptilia" family="Tyrannosauridae" genus="Thanatotheristes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus" status="gen. nov.">Thanatotheristes</taxonomicName>
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degrootorum
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</emphasis>
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gen. et. sp. nov.
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|
</paragraph>
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<paragraph blockId="2.[113,783,1433,1509]" pageId="2" pageNumber="3">
|
|
ZOOBANK ID:
|
|
<uri pageId="2" pageNumber="3">
|
|
urn:lsid:zoobank.org:act:
|
|
<uuid pageId="2" pageNumber="3">581C0612-B697-4D01- 915C-E3E937D3667A</uuid>
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|
</uri>
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection pageId="2" pageNumber="3" type="etymology">
|
|
<paragraph blockId="2.[113,784,1530,1969]" pageId="2" pageNumber="3">
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|
<emphasis box="[113,223,1530,1550]" italics="true" pageId="2" pageNumber="3">Etymology.</emphasis>
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From
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<taxonomicName box="[290,380,1530,1550]" genus="Thanatos" kingdom="Animalia" pageId="2" pageNumber="3" rank="genus">
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<emphasis box="[290,380,1530,1550]" italics="true" pageId="2" pageNumber="3">Thanatos</emphasis>
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</taxonomicName>
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|
, the Greek god of and embodiment of death, and
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<emphasis box="[229,313,1558,1578]" italics="true" pageId="2" pageNumber="3">theristes</emphasis>
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(Greek), one who reaps or harvests, a reaper. The specific name,
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<emphasis box="[316,444,1586,1606]" italics="true" pageId="2" pageNumber="3">degrootorum</emphasis>
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, is in honor of John and Sandra De Groot of Hays,
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<collectingRegion box="[315,392,1615,1634]" country="Canada" name="Alberta" pageId="2" pageNumber="3">Alberta</collectingRegion>
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, who discovered the
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|
<typeStatus box="[627,718,1615,1634]" pageId="2" pageNumber="3">holotype</typeStatus>
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specimen and have been supportive of paleontological research in the area.
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</paragraph>
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<paragraph blockId="2.[113,784,1530,1969]" pageId="2" pageNumber="3">
|
|
<emphasis box="[113,206,1698,1718]" italics="true" pageId="2" pageNumber="3">
|
|
<typeStatus box="[113,202,1698,1718]" pageId="2" pageNumber="3">Holotype</typeStatus>
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.
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|
</emphasis>
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<collectionCode box="[210,255,1699,1718]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="2" pageNumber="3">TMP</collectionCode>
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<date box="[261,345,1699,1718]" pageId="2" pageNumber="3" value="2010-05-07">2010.5.7</date>
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(
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<figureCitation box="[358,411,1699,1718]" captionStart="Fig" captionStartId="2.[831,856,771,785]" captionTargetBox="[837,1494,181,740]" captionTargetId="figure@2.[836,1497,178,741]" captionTargetPageId="2" captionText="Fig. 2. Skull reconstruction of Thanatotheristes holotype TMP 2010.5.7. Known bones appear in white. Missing bone morphologies and proportions are based on the holotype of Daspletosaurus torosus (CMN 8506). Scale bar equals 10 cm." figureDoi="http://doi.org/10.5281/zenodo.3672009" httpUri="https://zenodo.org/record/3672009/files/figure.png" pageId="2" pageNumber="3">Fig. 2</figureCitation>
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), a partial skull from a large subadult individual (estimated skull length
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<emphasis bold="true" box="[472,489,1727,1746]" pageId="2" pageNumber="3">¼</emphasis>
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~
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<quantity box="[511,600,1727,1746]" metricMagnitude="-1" metricUnit="m" metricValue="8.0" pageId="2" pageNumber="3" unit="mm" value="800.0">800 mm</quantity>
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, based on frontal width, metrics from
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<bibRefCitation author="Currie, P. J." box="[321,457,1755,1774]" journalOrPublisher="Canadian Journal of Earth Sciences" pageId="2" pageNumber="3" pagination="665" part="40" refId="ref10471" refString="Currie, P. J., 2003 a. Allometric growth in tyrannosaurids (Dinosauria: Theropoda) from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences 40, 651 e 665." title="Allometric growth in tyrannosaurids (Dinosauria: Theropoda) from the Upper Cretaceous of North America and Asia" type="journal article" year="2003">Currie, 2003a</bibRefCitation>
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) that includes the right maxilla, right jugal, right postorbital, right surangular, right quadrate, right laterosphenoid, left frontal, and both dentaries. Immature bone grain (most apparent within the antorbital fossa of the maxilla) indicates the specimen was not osteologically mature at the time of death (
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<bibRefCitation author="Carr, T. D." box="[184,282,1894,1913]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="2" pageNumber="3" pagination="520" part="19" refId="ref10070" refString="Carr, T. D., 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology 19, 497 e 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">Carr, 1999</bibRefCitation>
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), but the well-developed nature of ornamentation on the maxilla suggests the individual was probably sexually mature (
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<bibRefCitation author="Knell, R. J. & Naish, D. & Tomkins, J. L. & Hone, D. W. E." box="[201,372,1950,1969]" journalOrPublisher="Trends in Ecology & Evolution" pageId="2" pageNumber="3" pagination="47" part="28" refId="ref11827" refString="Knell, R. J., Naish, D., Tomkins, J. L., Hone, D. W. E., 2013. Sexual selection in prehistoric animals: detection and implications. Trends in Ecology & Evolution 28, 38 e 47." title="Sexual selection in prehistoric animals: detection and implications" type="journal article" year="2013">Knell et al., 2013</bibRefCitation>
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;
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<bibRefCitation author="Hone, D. W. E. & Farke, A. A. & Wedel, M. J." box="[383,552,1950,1969]" journalOrPublisher="Biology Letters" pageId="2" pageNumber="3" pagination="20150947" part="12" refId="ref11597" refString="Hone, D. W. E., Farke, A. A., Wedel, M. J., 2016. Ontogeny and the fossil record: what, if anything, is an adult dinosaur? Biology Letters 12, 20150947. https: // doi. org / 10.1098 / rsbl. 2015.0947." title="Ontogeny and the fossil record: what, if anything, is an adult dinosaur?" type="journal article" year="2016">Hone et al., 2016</bibRefCitation>
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).
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</paragraph>
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</subSubSection>
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<subSubSection pageId="2" pageNumber="3" type="materials_examined">
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<paragraph blockId="2.[831,1501,872,1590]" pageId="2" pageNumber="3">
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<emphasis box="[831,1081,872,892]" italics="true" pageId="2" pageNumber="3">Type locality and horizon.</emphasis>
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<materialsCitation ID-GBIF-Occurrence="2565937398" collectingDate="2010-05-07" collectionCode="TMP" country="Canada" county="Taber Coal Zone" elevation="11" location="Bow River" municipality="The" pageId="2" pageNumber="3" specimenCount="1" stateProvince="Alberta" typeStatus="holotype">
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<collectionCode box="[1088,1132,874,893]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="2" pageNumber="3">TMP</collectionCode>
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<date box="[1140,1222,873,892]" pageId="2" pageNumber="3" value="2010-05-07">
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<collectingDate box="[1140,1222,873,892]" pageId="2" pageNumber="3" value="2010-05-07">2010.5.7</collectingDate>
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</date>
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was found at the base of a steep cliff on the eastern shore of the
|
|
<location LSID="urn:lsid:plazi:treatment:039E87F3FFE7FFDAB92BDBF56DCEFC17:8EE8603EFFE7FFD0BDE6DDEA6C7EFC0F" box="[1212,1314,901,921]" country="Canada" county="Taber Coal Zone" municipality="The" name="Bow River" pageId="2" pageNumber="3" stateProvince="Alberta">Bow River</location>
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approximately
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<quantity metricMagnitude="2" metricUnit="m" metricValue="5.0" pageId="2" pageNumber="3" unit="km" value="0.5">0.5 km</quantity>
|
|
upstream of the confluence with the
|
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<location LSID="urn:lsid:plazi:treatment:039E87F3FFE7FFDAB92BDBF56DCEFC17:8EE8603EFFE7FFD0BDAADDCD6C20FC23" box="[1264,1404,929,948]" country="Canada" county="Taber Coal Zone" municipality="The" name="Oldman River" pageId="2" pageNumber="3" stateProvince="Alberta">Oldman River</location>
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(i.e., The Forks) in
|
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<collectingRegion box="[922,995,957,976]" country="Canada" name="Alberta" pageId="2" pageNumber="3">Alberta</collectingRegion>
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,
|
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<collectingCountry box="[1004,1076,957,976]" name="Canada" pageId="2" pageNumber="3">Canada</collectingCountry>
|
|
(
|
|
<figureCitation box="[1089,1159,957,976]" captionStart="Fig" captionStartId="3.[88,113,1453,1467]" captionTargetBox="[178,1387,179,1422]" captionTargetId="figure@3.[175,1388,178,1423]" captionTargetPageId="3" captionText="Fig. 3. Locality of Thanatotheristes degrootorum holotype (TMP 2010.5.7).A) Geographic location of the locality.The specimen was found 0.5 km upstream from the confluence of the Bow and Oldman rivers. B) Photograph of locality looking NNE. Specimen was found scattered over a lateral extent of 10 m on the shore of the river. The nearby cliff is composed of coal beds and interbedded sandstones and mudstones of the Taber Coal Zone of the Foremost Formation. C) Stratigraphic section of the exposures near the fossil locality. Bracket indicates likely stratigraphic provenance of specimen. Dotted line marks the extent of the exposures at the type locality; the Herronton Sandstone is exposed farther downstream." figureDoi="http://doi.org/10.5281/zenodo.3672011" httpUri="https://zenodo.org/record/3672011/files/figure.png" pageId="2" pageNumber="3">Fig. 3A</figureCitation>
|
|
).
|
|
<location LSID="urn:lsid:plazi:treatment:039E87F3FFE7FFDAB92BDBF56DCEFC17:8EE8603EFFE7FFD0BDCFDDD16DE5FC47" box="[1173,1209,957,976]" country="Canada" county="Taber Coal Zone" municipality="The" name="The" pageId="2" pageNumber="3" stateProvince="Alberta">The</location>
|
|
cliff is approximately
|
|
<quantity box="[1427,1475,957,976]" metricMagnitude="1" metricUnit="m" metricValue="1.1" pageId="2" pageNumber="3" unit="m" value="11.0">
|
|
<elevation box="[1427,1475,957,976]" metricMagnitude="1" metricUnit="m" metricValue="1.1" pageId="2" pageNumber="3" unit="m" value="11.0">11 m</elevation>
|
|
</quantity>
|
|
in height and consists of exposures of the
|
|
<location LSID="urn:lsid:plazi:treatment:039E87F3FFE7FFDAB92BDBF56DCEFC17:8EE8603EFFE7FFD0BD97DDB56C31FC7B" box="[1229,1389,985,1004]" country="Canada" county="Taber Coal Zone" municipality="The" name="Taber Coal Zone" pageId="2" pageNumber="3" stateProvince="Alberta">Taber Coal Zone</location>
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|
, below the
|
|
<location LSID="urn:lsid:plazi:treatment:039E87F3FFE7FFDAB92BDBF56DCEFC17:8EE8603EFFE7FFD0BA65DD996D4EFB9F" box="[831,1042,1013,1032]" country="Canada" county="Taber Coal Zone" municipality="The" name="Herronton Sandstone" pageId="2" pageNumber="3" stateProvince="Alberta">Herronton Sandstone</location>
|
|
, of the
|
|
<location LSID="urn:lsid:plazi:treatment:039E87F3FFE7FFDAB92BDBF56DCEFC17:8EE8603EFFE7FFD0BD03DD996C7CFB9F" box="[1113,1312,1013,1032]" country="Canada" county="Taber Coal Zone" municipality="The" name="Foremost Formation" pageId="2" pageNumber="3" stateProvince="Alberta">Foremost Formation</location>
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|
(
|
|
<figureCitation box="[1324,1392,1013,1032]" captionStart="Fig" captionStartId="3.[88,113,1453,1467]" captionTargetBox="[178,1387,179,1422]" captionTargetId="figure@3.[175,1388,178,1423]" captionTargetPageId="3" captionText="Fig. 3. Locality of Thanatotheristes degrootorum holotype (TMP 2010.5.7).A) Geographic location of the locality.The specimen was found 0.5 km upstream from the confluence of the Bow and Oldman rivers. B) Photograph of locality looking NNE. Specimen was found scattered over a lateral extent of 10 m on the shore of the river. The nearby cliff is composed of coal beds and interbedded sandstones and mudstones of the Taber Coal Zone of the Foremost Formation. C) Stratigraphic section of the exposures near the fossil locality. Bracket indicates likely stratigraphic provenance of specimen. Dotted line marks the extent of the exposures at the type locality; the Herronton Sandstone is exposed farther downstream." figureDoi="http://doi.org/10.5281/zenodo.3672011" httpUri="https://zenodo.org/record/3672011/files/figure.png" pageId="2" pageNumber="3">Fig. 3B</figureCitation>
|
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).
|
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<collectingMunicipality box="[1407,1443,1013,1032]" pageId="2" pageNumber="3">The</collectingMunicipality>
|
|
exact stratigraphic provenance of the specimen within the
|
|
<collectingCounty box="[1345,1500,1041,1060]" pageId="2" pageNumber="3">Taber Coal Zone</collectingCounty>
|
|
is unknown, as the skull was found scattered along the shoreline over a lateral extent of ~
|
|
<quantity box="[1027,1078,1097,1116]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="2" pageNumber="3" unit="m" value="10.0">
|
|
<elevation box="[1027,1078,1097,1116]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="2" pageNumber="3" unit="m" value="10.0">10 m</elevation>
|
|
</quantity>
|
|
, but the specimen likely derived from the lower half of the cliff due to a small-scale landslide (
|
|
<figureCitation box="[1351,1420,1125,1144]" captionStart="Fig" captionStartId="3.[88,113,1453,1467]" captionTargetBox="[178,1387,179,1422]" captionTargetId="figure@3.[175,1388,178,1423]" captionTargetPageId="3" captionText="Fig. 3. Locality of Thanatotheristes degrootorum holotype (TMP 2010.5.7).A) Geographic location of the locality.The specimen was found 0.5 km upstream from the confluence of the Bow and Oldman rivers. B) Photograph of locality looking NNE. Specimen was found scattered over a lateral extent of 10 m on the shore of the river. The nearby cliff is composed of coal beds and interbedded sandstones and mudstones of the Taber Coal Zone of the Foremost Formation. C) Stratigraphic section of the exposures near the fossil locality. Bracket indicates likely stratigraphic provenance of specimen. Dotted line marks the extent of the exposures at the type locality; the Herronton Sandstone is exposed farther downstream." figureDoi="http://doi.org/10.5281/zenodo.3672011" httpUri="https://zenodo.org/record/3672011/files/figure.png" pageId="2" pageNumber="3">Fig. 3C</figureCitation>
|
|
)
|
|
</materialsCitation>
|
|
.
|
|
</paragraph>
|
|
<paragraph blockId="2.[831,1501,872,1590]" pageId="2" pageNumber="3">
|
|
<emphasis box="[831,1017,1151,1171]" italics="true" pageId="2" pageNumber="3">Referred specimen.</emphasis>
|
|
<collectionCode box="[1027,1072,1153,1172]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="2" pageNumber="3">TMP</collectionCode>
|
|
2018.16.1, a partial right maxilla from a subadult individual.
|
|
<collectionCode box="[1042,1087,1181,1200]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="2" pageNumber="3">TMP</collectionCode>
|
|
2018.16.1 was found along Twelve Mile Coulee, approximately
|
|
<quantity box="[1071,1137,1208,1227]" metricMagnitude="4" metricUnit="m" metricValue="1.0" pageId="2" pageNumber="3" unit="km" value="10.0">10 km</quantity>
|
|
northeast of Hays,
|
|
<collectingRegion box="[1350,1425,1208,1227]" country="Canada" name="Alberta" pageId="2" pageNumber="3">Alberta</collectingRegion>
|
|
, stratigraphically high within the Herronton Sandstone of the Foremost Formation.
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection pageId="2" pageNumber="3" type="diagnosis">
|
|
<paragraph blockId="2.[831,1501,872,1590]" pageId="2" pageNumber="3">
|
|
<emphasis box="[831,930,1291,1311]" italics="true" pageId="2" pageNumber="3">Diagnosis.</emphasis>
|
|
Tyrannosaurine tyrannosaurid diagnosed by the following autapomorphies: single row of evenly-spaced dorsoventrally-oriented ridges on the subcutaneous surface of the maxilla ventral and anteroventral to the antorbital fossa; rounded and inflated orbital margin of the jugal; sagittal crest on the frontal extends anterior to the supratemporal ridge as a broad and rounded ridge; lacrimal contact surface on the frontal extends anteromedially at ~60 relative to interfrontal suture; prefrontal with two posteriorly-projecting prongs articulating with the frontal on ventral surface of skull roof (long, medial, primary prong and shorter secondary, lateral prong).
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection lastPageId="8" lastPageNumber="9" pageId="2" pageNumber="3" type="description">
|
|
<paragraph blockId="2.[831,984,1657,1677]" box="[831,984,1657,1677]" pageId="2" pageNumber="3">
|
|
<heading bold="true" box="[831,984,1657,1677]" fontSize="8" level="1" pageId="2" pageNumber="3" reason="0">
|
|
<emphasis bold="true" box="[831,984,1657,1677]" pageId="2" pageNumber="3">5. Description</emphasis>
|
|
</heading>
|
|
</paragraph>
|
|
<paragraph blockId="2.[831,1501,1713,1984]" lastBlockId="3.[88,758,1594,1976]" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3">
|
|
<emphasis bold="true" box="[863,943,1713,1732]" pageId="2" pageNumber="3">Maxilla</emphasis>
|
|
-The partial right maxilla of the
|
|
<typeStatus box="[1298,1389,1714,1733]" pageId="2" pageNumber="3">holotype</typeStatus>
|
|
specimen,
|
|
<collectionCode box="[831,877,1742,1761]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="2" pageNumber="3">TMP</collectionCode>
|
|
<date box="[889,975,1741,1760]" pageId="2" pageNumber="3" value="2010-05-07">2010.5.7</date>
|
|
, consists of a complete anterior body and two fragments of the jugal ramus, one of which preserves the three posterior-most alveoli and the groove-like primary posterior foramen of the maxillary nerve and the second preserves a single alveolus (
|
|
<figureCitation box="[1008,1068,1853,1872]" captionStart="Fig" captionStartId="4.[113,138,1672,1686]" captionTargetBox="[202,1409,184,1640]" captionTargetId="figure@4.[200,1413,178,1642]" captionTargetPageId="4" captionText="Fig. 4. Right maxilla of Thanatotheristes holotype (TMP 2010.5.7) in lateral view with A) known fragments overlain on hypothesized reconstruction and B) subcutaneous surface magnified. The subcutaneous surface is strongly ornamented and possesses autapomorphic dorsoventral ridges (arrows). Extent of antorbital fossa indicated by black dash line. Large healed bite mark is indicated by white dash line. Abbreviations: aofo, antorbital fossa; mf, maxillary fenestra; pmf, promaxillary fenestra; pnf, posterior-most neurovascular foramen; scs, subcutaneous surface; sppm, subnarial process of premaxilla. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672013" httpUri="https://zenodo.org/record/3672013/files/figure.png" pageId="2" pageNumber="3">Fig. 4</figureCitation>
|
|
). The anterior body is deep, consistent with the proportions of other medium to large-sized tyrannosaurines. The anteroventral corner of the bone is slightly downturned and tapers into a shallow angle, as in
|
|
<taxonomicName authorityName="Russell" authorityYear="1970" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
|
|
<emphasis italics="true" pageId="2" pageNumber="3">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
. The referred maxilla (
|
|
<collectionCode box="[1159,1207,1965,1984]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="2" pageNumber="3">TMP</collectionCode>
|
|
2018.16.1) consists of the subcutaneous surface ventral to the anterior half of the antorbital fossa (
|
|
<figureCitation box="[211,268,1622,1641]" captionStart="Fig" captionStartId="5.[88,113,874,888]" captionTargetBox="[329,1235,181,841]" captionTargetId="figure@5.[326,1236,178,844]" captionTargetPageId="5" captionText="Fig. 5. Left maxilla of referred specimen of Thanatotheristes (TMP 2018.16.1) in lateral view. Note the presence of pronounced vertical ridges (arrows) on subcutaneous surface, an autapomorphy of the taxon. Abbreviations: dvr, dorsoventral ridges; sc-afos, subcutaneous-antorbital fossa boundary. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672015" httpUri="https://zenodo.org/record/3672015/files/figure.png" pageId="3" pageNumber="4">Fig. 5</figureCitation>
|
|
).
|
|
</paragraph>
|
|
<caption ID-DOI="http://doi.org/10.5281/zenodo.3672011" ID-Zenodo-Dep="3672011" httpUri="https://zenodo.org/record/3672011/files/figure.png" pageId="3" pageNumber="4" startId="3.[88,113,1453,1467]" targetBox="[178,1387,179,1422]" targetPageId="3">
|
|
<paragraph blockId="3.[88,1476,1452,1536]" pageId="3" pageNumber="4">
|
|
<emphasis bold="true" box="[88,134,1453,1467]" pageId="3" pageNumber="4">Fig. 3.</emphasis>
|
|
Locality of
|
|
<emphasis box="[225,450,1452,1467]" italics="true" pageId="3" pageNumber="4">
|
|
<taxonomicName authority="Voris & Therrien & Zelenitsky & Brown, 2020" authorityName="Voris & Therrien & Zelenitsky & Brown" authorityYear="2020" box="[225,349,1452,1467]" class="Reptilia" family="Tyrannosauridae" genus="Thanatotheristes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus" status="gen. nov.">Thanatotheristes</taxonomicName>
|
|
degrootorum
|
|
</emphasis>
|
|
holotype (TMP 2010.5.7). A) Geographic location of the locality. The specimen was found 0.5 km upstream from the confluence of the Bow and Oldman rivers. B) Photograph of locality looking NNE. Specimen was found scattered over a lateral extent of 10 m on the shore of the river. The nearby cliff is composed of coal beds and interbedded sandstones and mudstones of the Taber Coal Zone of the Foremost Formation. C) Stratigraphic section of the exposures near the fossil locality. Bracket indicates likely stratigraphic provenance of specimen. Dotted line marks the extent of the exposures at the type locality; the Herronton Sandstone is exposed farther downstream.
|
|
</paragraph>
|
|
</caption>
|
|
<paragraph blockId="3.[88,758,1594,1976]" pageId="3" pageNumber="4">
|
|
The subcutaneous surface of both specimens is extremely rugose, particularly anteroventral to the antorbital fossa where it is incised by deep anastomosing sulci, as in
|
|
<taxonomicName authority="(Carr et al., 2017)" baseAuthorityName="Carr" baseAuthorityYear="2017" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|
<emphasis box="[498,645,1704,1724]" italics="true" pageId="3" pageNumber="4">Daspletosaurus</emphasis>
|
|
(
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." journalOrPublisher="Scientific Reports" pageId="3" pageNumber="4" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
)
|
|
</taxonomicName>
|
|
. The subcutaneous texturing differs from that of
|
|
<taxonomicName class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|
<emphasis italics="true" pageId="3" pageNumber="4">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
and all other tyrannosaurids in the presence of a single row of prominent and evenly spaced dorsoventrally-oriented ridges located ventral and anteroventral to the antorbital fossa (
|
|
<figureCitation box="[653,715,1817,1836]" captionStart="Fig" captionStartId="4.[113,138,1672,1686]" captionTargetBox="[202,1409,184,1640]" captionTargetId="figure@4.[200,1413,178,1642]" captionTargetPageId="4" captionText="Fig. 4. Right maxilla of Thanatotheristes holotype (TMP 2010.5.7) in lateral view with A) known fragments overlain on hypothesized reconstruction and B) subcutaneous surface magnified. The subcutaneous surface is strongly ornamented and possesses autapomorphic dorsoventral ridges (arrows). Extent of antorbital fossa indicated by black dash line. Large healed bite mark is indicated by white dash line. Abbreviations: aofo, antorbital fossa; mf, maxillary fenestra; pmf, promaxillary fenestra; pnf, posterior-most neurovascular foramen; scs, subcutaneous surface; sppm, subnarial process of premaxilla. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672013" httpUri="https://zenodo.org/record/3672013/files/figure.png" pageId="3" pageNumber="4">Figs. 4</figureCitation>
|
|
and
|
|
<figureCitation box="[88,101,1845,1864]" captionStart="Fig" captionStartId="5.[88,113,874,888]" captionTargetBox="[329,1235,181,841]" captionTargetId="figure@5.[326,1236,178,844]" captionTargetPageId="5" captionText="Fig. 5. Left maxilla of referred specimen of Thanatotheristes (TMP 2018.16.1) in lateral view. Note the presence of pronounced vertical ridges (arrows) on subcutaneous surface, an autapomorphy of the taxon. Abbreviations: dvr, dorsoventral ridges; sc-afos, subcutaneous-antorbital fossa boundary. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672015" httpUri="https://zenodo.org/record/3672015/files/figure.png" pageId="3" pageNumber="4">5</figureCitation>
|
|
). In
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[148,291,1844,1864]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|
<emphasis box="[148,291,1844,1864]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[304,422,1844,1864]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|
<emphasis box="[304,422,1844,1864]" italics="true" pageId="3" pageNumber="4">Tarbosaurus</emphasis>
|
|
</taxonomicName>
|
|
, and
|
|
<taxonomicName box="[480,659,1844,1864]" class="Reptilia" family="Tyrannosauridae" genus="Zhuchengtyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|
<emphasis box="[480,659,1844,1864]" italics="true" pageId="3" pageNumber="4">Zhuchengtyrannus</emphasis>
|
|
</taxonomicName>
|
|
, the subcutaneous surface is ornamented with at least two rows of unevenly spaced and anastomosing ridges that circumferent shallow oval fossae (
|
|
<bibRefCitation author="Hone, D. W. E. & Wang, K. & Sullivan, C. & Zhao, X. & Chen, S. & Li, D. & Ji, S. & Ji, Q. & Xu, X." box="[217,396,1929,1948]" journalOrPublisher="Cretaceous Research" pageId="3" pageNumber="4" pagination="503" part="32" refId="ref11657" refString="Hone, D. W. E., Wang, K., Sullivan, C., Zhao, X., Chen, S., Li, D., Ji, S., Ji, Q., Xu, X., 2011. A new, large tyrannosaurine theropod from the Upper Cretaceous of China. Cretaceous Research 32, 495 e 503." title="A new, large tyrannosaurine theropod from the Upper Cretaceous of China" type="journal article" year="2011">Hone et al., 2011</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[410,575,1929,1948]" journalOrPublisher="Scientific Reports" pageId="3" pageNumber="4" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
); such fossae are absent in
|
|
<taxonomicName authority="Voris & Therrien & Zelenitsky & Brown, 2020" authorityName="Voris & Therrien & Zelenitsky & Brown" authorityYear="2020" box="[186,347,1955,1975]" class="Reptilia" family="Tyrannosauridae" genus="Thanatotheristes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus" status="gen. nov.">
|
|
<emphasis box="[186,347,1955,1975]" italics="true" pageId="3" pageNumber="4">Thanatotheristes</emphasis>
|
|
</taxonomicName>
|
|
.
|
|
</paragraph>
|
|
<paragraph blockId="3.[805,1475,1594,1976]" pageId="3" pageNumber="4">
|
|
The ventral margin of the promaxillary fenestra in
|
|
<collectionCode box="[1340,1385,1595,1614]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="3" pageNumber="4">TMP</collectionCode>
|
|
<date box="[1390,1474,1594,1613]" pageId="3" pageNumber="4" value="2010-05-07">2010.5.7</date>
|
|
appears as a wide concavity in the far anterior corner of the antorbital fossa in lateral view (
|
|
<figureCitation box="[1121,1174,1650,1669]" captionStart="Fig" captionStartId="4.[113,138,1672,1686]" captionTargetBox="[202,1409,184,1640]" captionTargetId="figure@4.[200,1413,178,1642]" captionTargetPageId="4" captionText="Fig. 4. Right maxilla of Thanatotheristes holotype (TMP 2010.5.7) in lateral view with A) known fragments overlain on hypothesized reconstruction and B) subcutaneous surface magnified. The subcutaneous surface is strongly ornamented and possesses autapomorphic dorsoventral ridges (arrows). Extent of antorbital fossa indicated by black dash line. Large healed bite mark is indicated by white dash line. Abbreviations: aofo, antorbital fossa; mf, maxillary fenestra; pmf, promaxillary fenestra; pnf, posterior-most neurovascular foramen; scs, subcutaneous surface; sppm, subnarial process of premaxilla. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672013" httpUri="https://zenodo.org/record/3672013/files/figure.png" pageId="3" pageNumber="4">Fig. 4</figureCitation>
|
|
). The presence of a short base of the preantral strut, however, suggests the exposure of the promaxillary fenestra in lateral view is the result of taphonomic postburial deformation, and that the promaxillary fenestra would have been concealed in lateral view, as in derived tyrannosaurines (e.g.,
|
|
<taxonomicName box="[856,1003,1788,1808]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|
<emphasis box="[856,1003,1788,1808]" italics="true" pageId="3" pageNumber="4">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[1014,1132,1788,1808]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1014,1132,1788,1808]" italics="true" pageId="3" pageNumber="4">Tarbosaurus</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1141,1284,1788,1808]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1141,1284,1788,1808]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
|
|
</taxonomicName>
|
|
;
|
|
<bibRefCitation author="Brusatte, S. L. & Carr, T. D." journalOrPublisher="Scientific Reports" pageId="3" pageNumber="4" pagination="20252" part="6" refId="ref9758" refString="Brusatte, S. L., Carr, T. D., 2016. The phylogeny and evolutionary history of tyrannosauroid dinosaurs. Scientific Reports 6, 20252. https: // doi. org / 10.1038 / srep 20252." title="The phylogeny and evolutionary history of tyrannosauroid dinosaurs" type="journal article" year="2016">Brusatte and Carr, 2016</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[868,1025,1817,1837]" journalOrPublisher="Scientific Reports" pageId="3" pageNumber="4" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
).
|
|
</paragraph>
|
|
<paragraph blockId="3.[805,1475,1594,1976]" lastBlockId="4.[831,1501,1813,1972]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
|
|
The ventral margin of the maxillary fenestra is broken in
|
|
<collectionCode box="[1429,1474,1846,1865]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="3" pageNumber="4">TMP</collectionCode>
|
|
<date box="[805,889,1873,1892]" pageId="3" pageNumber="4" value="2010-05-07">2010.5.7</date>
|
|
but its location can be determined based on the thinness of the bone between two thickened areas representing the base of the interfenestral and preantral struts. The ventral margin of the maxillary fenestra is widely separated from the ventral margin of the antorbital fossa, as in albertosaurines, alioramins,
|
|
<emphasis italics="true" pageId="4" pageNumber="5">Teratophoneus</emphasis>
|
|
, and juvenile individuals of the genera
|
|
<taxonomicName box="[636,783,1840,1860]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|
<emphasis box="[636,783,1840,1860]" italics="true" pageId="4" pageNumber="5">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[156,299,1868,1888]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|
<emphasis box="[156,299,1868,1888]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
|
|
</taxonomicName>
|
|
(
|
|
<bibRefCitation author="Carr, T. D." box="[312,415,1869,1888]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="4" pageNumber="5" pagination="520" part="19" refId="ref10070" refString="Carr, T. D., 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology 19, 497 e 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">Carr, 1999</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[423,575,1869,1889]" journalOrPublisher="Scientific Reports" pageId="4" pageNumber="5" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
). This contrasts with the condition seen in subadult and adult
|
|
<taxonomicName box="[592,739,1896,1916]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|
<emphasis box="[592,739,1896,1916]" italics="true" pageId="4" pageNumber="5">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
individuals, where the ventral margin of the maxillary fenestra is closer to the ventral margin of the antorbital fossa (
|
|
<bibRefCitation author="Brusatte, S. L. & Carr, T. D. & Norell, M. A." box="[639,880,1813,1972]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="4" pageNumber="5" pagination="197" part="366" refId="ref9853" refString="Brusatte, S. L., Carr, T. D., Norell, M. A., 2012. The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bulletin of the American Museum of Natural History 366, 1 e 197." title="The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia" type="journal article" year="2012">Brusatte et al., 2012</bibRefCitation>
|
|
), and in subadult and adult individuals of more derived tyrannosaurines (
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1013,1156,1840,1860]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1013,1156,1840,1860]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[1173,1291,1840,1860]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1173,1291,1840,1860]" italics="true" pageId="4" pageNumber="5">Tarbosaurus</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<taxonomicName box="[1308,1487,1840,1860]" class="Reptilia" family="Tyrannosauridae" genus="Zhuchengtyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1308,1487,1840,1860]" italics="true" pageId="4" pageNumber="5">Zhuchengtyrannus</emphasis>
|
|
</taxonomicName>
|
|
), where the maxillary fenestra is in contact with the margin of the antorbital fossa (
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[1001,1158,1897,1917]" journalOrPublisher="Scientific Reports" pageId="4" pageNumber="5" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
).
|
|
</paragraph>
|
|
<caption ID-DOI="http://doi.org/10.5281/zenodo.3672013" ID-Zenodo-Dep="3672013" httpUri="https://zenodo.org/record/3672013/files/figure.png" pageId="4" pageNumber="5" startId="4.[113,138,1672,1686]" targetBox="[202,1409,184,1640]" targetPageId="4">
|
|
<paragraph blockId="4.[113,1502,1671,1755]" pageId="4" pageNumber="5">
|
|
<emphasis bold="true" box="[113,162,1672,1686]" pageId="4" pageNumber="5">Fig. 4.</emphasis>
|
|
Right maxilla of
|
|
<taxonomicName authority="Voris & Therrien & Zelenitsky & Brown, 2020" authorityName="Voris & Therrien & Zelenitsky & Brown" authorityYear="2020" box="[301,425,1671,1686]" class="Reptilia" family="Tyrannosauridae" genus="Thanatotheristes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="3" phylum="Chordata" rank="genus" status="gen. nov.">
|
|
<emphasis box="[301,425,1671,1686]" italics="true" pageId="4" pageNumber="5">Thanatotheristes</emphasis>
|
|
</taxonomicName>
|
|
holotype (TMP 2010.5.7) in lateral view with A) known fragments overlain on hypothesized reconstruction and B) subcutaneous surface magnified. The subcutaneous surface is strongly ornamented and possesses autapomorphic dorsoventral ridges (arrows). Extent of antorbital fossa indicated by black dash line. Large healed bite mark is indicated by white dash line. Abbreviations: aofo, antorbital fossa; mf, maxillary fenestra; pmf, promaxillary fenestra; pnf, posterior-most neurovascular foramen; scs, subcutaneous surface; sppm, subnarial process of premaxilla. Scale bar equals 5 cm.
|
|
</paragraph>
|
|
</caption>
|
|
<paragraph blockId="4.[831,1501,1813,1972]" lastBlockId="5.[88,758,969,1965]" lastPageId="5" lastPageNumber="6" pageId="4" pageNumber="5">
|
|
The neurovascular foramina of the maxilla are large in
|
|
<collectionCode box="[1455,1500,1925,1944]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="4" pageNumber="5">TMP</collectionCode>
|
|
<date box="[831,914,1953,1972]" pageId="4" pageNumber="5" value="2010-05-07">2010.5.7</date>
|
|
and occur in two distinct rows: a dorsal circumfenestral row inset into an elongate fossa and a ventral alveolar row. Anteriorly, the circumfenestral foramina diverge from the linear arrangement with individual foramina becoming randomly dispersed. Several deep and anteroventrally-oriented neurovascular sulci originate from these anterior circumfenestral foramina. The posterior-most foramen of the maxilla exits the subcutaneous surface between the two posterior-most alveoli and continues posteriorly as an elongate neurovascular sulcus, as seen in
|
|
<taxonomicName box="[117,213,1191,1211]" class="Reptilia" family="Tyrannosauridae" genus="Lythronax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis box="[117,213,1191,1211]" italics="true" pageId="5" pageNumber="6">Lythronax</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<taxonomicName box="[229,376,1191,1211]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis box="[229,376,1191,1211]" italics="true" pageId="5" pageNumber="6">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
, and
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[439,582,1191,1211]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis box="[439,582,1191,1211]" italics="true" pageId="5" pageNumber="6">Tyrannosaurus</emphasis>
|
|
</taxonomicName>
|
|
. In contrast, the foramen is located posterior to the posterior-most alveolus in albertosaurines.
|
|
</paragraph>
|
|
<caption ID-DOI="http://doi.org/10.5281/zenodo.3672015" ID-Zenodo-Dep="3672015" httpUri="https://zenodo.org/record/3672015/files/figure.png" pageId="5" pageNumber="6" startId="5.[88,113,874,888]" targetBox="[329,1235,181,841]" targetPageId="5">
|
|
<paragraph blockId="5.[88,1474,873,911]" pageId="5" pageNumber="6">
|
|
<emphasis bold="true" box="[88,135,874,888]" pageId="5" pageNumber="6">Fig. 5.</emphasis>
|
|
Left maxilla of referred specimen of
|
|
<taxonomicName authority="Voris & Therrien & Zelenitsky & Brown, 2020" authorityName="Voris & Therrien & Zelenitsky & Brown" authorityYear="2020" box="[427,551,873,888]" class="Reptilia" family="Tyrannosauridae" genus="Thanatotheristes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="3" phylum="Chordata" rank="genus" status="gen. nov.">
|
|
<emphasis box="[427,551,873,888]" italics="true" pageId="5" pageNumber="6">Thanatotheristes</emphasis>
|
|
</taxonomicName>
|
|
(TMP 2018.16.1) in lateral view. Note the presence of pronounced vertical ridges (arrows) on subcutaneous surface, an autapomorphy of the taxon. Abbreviations: dvr, dorsoventral ridges; sc-afos, subcutaneous-antorbital fossa boundary. Scale bar equals 5 cm.
|
|
</paragraph>
|
|
</caption>
|
|
<paragraph blockId="5.[88,758,969,1965]" pageId="5" pageNumber="6">
|
|
The jugal ramus of the maxilla in
|
|
<collectionCode box="[451,496,1277,1296]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="5" pageNumber="6">TMP</collectionCode>
|
|
<date box="[501,585,1276,1295]" pageId="5" pageNumber="6" value="2010-05-07">2010.5.7</date>
|
|
is dorsoventrally constricted at the level of the two posterior alveoli, such that the posterior-most maxillary foramen lies near the dorsal margin of the subcutaneous surface (
|
|
<figureCitation box="[332,393,1360,1379]" captionStart="Fig" captionStartId="4.[113,138,1672,1686]" captionTargetBox="[202,1409,184,1640]" captionTargetId="figure@4.[200,1413,178,1642]" captionTargetPageId="4" captionText="Fig. 4. Right maxilla of Thanatotheristes holotype (TMP 2010.5.7) in lateral view with A) known fragments overlain on hypothesized reconstruction and B) subcutaneous surface magnified. The subcutaneous surface is strongly ornamented and possesses autapomorphic dorsoventral ridges (arrows). Extent of antorbital fossa indicated by black dash line. Large healed bite mark is indicated by white dash line. Abbreviations: aofo, antorbital fossa; mf, maxillary fenestra; pmf, promaxillary fenestra; pnf, posterior-most neurovascular foramen; scs, subcutaneous surface; sppm, subnarial process of premaxilla. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672013" httpUri="https://zenodo.org/record/3672013/files/figure.png" pageId="5" pageNumber="6">Fig. 4</figureCitation>
|
|
), as in
|
|
<taxonomicName box="[485,632,1359,1379]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis box="[485,632,1359,1379]" italics="true" pageId="5" pageNumber="6">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
. In mature specimens of all other tyrannosaurids, the jugal ramus gradually tapers posteriorly and the posterior-most maxillary foramen is widely separated from the dorsal margin of the subcutaneous surface.
|
|
</paragraph>
|
|
<paragraph blockId="5.[88,758,969,1965]" pageId="5" pageNumber="6">
|
|
In total, the anterior body of the maxilla and posterior jugal ramus fragments of
|
|
<collectionCode box="[296,341,1528,1547]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="5" pageNumber="6">TMP</collectionCode>
|
|
<date box="[349,433,1527,1546]" pageId="5" pageNumber="6" value="2010-05-07">2010.5.7</date>
|
|
preserve 12 teeth. However, as these fragments do not articulate with each other and are broadly separated, it is likely that the individual possessed at least 15 maxillary teeth, indicating the animal possessed a long snout (
|
|
<figureCitation box="[96,153,1639,1658]" captionStart="Fig" captionStartId="4.[113,138,1672,1686]" captionTargetBox="[202,1409,184,1640]" captionTargetId="figure@4.[200,1413,178,1642]" captionTargetPageId="4" captionText="Fig. 4. Right maxilla of Thanatotheristes holotype (TMP 2010.5.7) in lateral view with A) known fragments overlain on hypothesized reconstruction and B) subcutaneous surface magnified. The subcutaneous surface is strongly ornamented and possesses autapomorphic dorsoventral ridges (arrows). Extent of antorbital fossa indicated by black dash line. Large healed bite mark is indicated by white dash line. Abbreviations: aofo, antorbital fossa; mf, maxillary fenestra; pmf, promaxillary fenestra; pnf, posterior-most neurovascular foramen; scs, subcutaneous surface; sppm, subnarial process of premaxilla. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672013" httpUri="https://zenodo.org/record/3672013/files/figure.png" pageId="5" pageNumber="6">Fig. 4</figureCitation>
|
|
), as in
|
|
<taxonomicName box="[238,385,1638,1658]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis box="[238,385,1638,1658]" italics="true" pageId="5" pageNumber="6">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
. The base of the tooth crowns are marginally narrower than long (fore-aft base length
|
|
<emphasis box="[665,680,1666,1685]" italics="true" pageId="5" pageNumber="6">></emphasis>
|
|
crown width). The mesial carinae terminate distal to the base of the crown, which differs from that reported for specimens of
|
|
<taxonomicName class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis italics="true" pageId="5" pageNumber="6">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
in which the carinae extend to the base of the crown (
|
|
<bibRefCitation author="Carr, T. D. & Williamson, T. E." journalOrPublisher="New Mexico Museum of Natural History and Science Bulletin" pageId="5" pageNumber="6" pagination="145" part="17" refId="ref10099" refString="Carr, T. D., Williamson, T. E., 2000. A review of Tyrannosauridae (Dinosauria, Coelurosauria) from New Mexico. New Mexico Museum of Natural History and Science Bulletin 17, 113 e 145." title="A review of Tyrannosauridae (Dinosauria, Coelurosauria) from New Mexico" type="journal article" year="2000">Carr and Williamson, 2000</bibRefCitation>
|
|
).
|
|
</paragraph>
|
|
<paragraph blockId="5.[88,758,969,1965]" lastBlockId="5.[805,1475,969,1965]" pageId="5" pageNumber="6">
|
|
The subcutaneous surface of
|
|
<collectionCode box="[439,484,1807,1826]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="5" pageNumber="6">TMP</collectionCode>
|
|
<date box="[498,581,1806,1825]" pageId="5" pageNumber="6" value="2010-05-07">2010.5.7</date>
|
|
is incised by a prominent anteroventral bite mark located immediately ventral to the antorbital fossa (
|
|
<typeStatus box="[319,369,1862,1881]" pageId="5" pageNumber="6">Type</typeStatus>
|
|
2: sensu
|
|
<bibRefCitation author="Tanke, D. H. & Currie, P. J." box="[489,744,1862,1881]" journalOrPublisher="Gaia" pageId="5" pageNumber="6" pagination="184" part="15" refId="ref13453" refString="Tanke, D. H., Currie, P. J., 1998. Head-biting behavior in theropod dinosaurs: paleopathological evidence. Gaia 15, 167 e 184." title="Head-biting behavior in theropod dinosaurs: paleopathological evidence" type="journal article" year="1998">Tanke and Currie, 1998</bibRefCitation>
|
|
). Orientation of the healed bone grain indicates the bite originated at the dorsal extent of the scar and was dragged downward to the alveolar margin at the anterior edge of the sixth alveolus (
|
|
<figureCitation box="[689,744,1946,1965]" captionStart="Fig" captionStartId="4.[113,138,1672,1686]" captionTargetBox="[202,1409,184,1640]" captionTargetId="figure@4.[200,1413,178,1642]" captionTargetPageId="4" captionText="Fig. 4. Right maxilla of Thanatotheristes holotype (TMP 2010.5.7) in lateral view with A) known fragments overlain on hypothesized reconstruction and B) subcutaneous surface magnified. The subcutaneous surface is strongly ornamented and possesses autapomorphic dorsoventral ridges (arrows). Extent of antorbital fossa indicated by black dash line. Large healed bite mark is indicated by white dash line. Abbreviations: aofo, antorbital fossa; mf, maxillary fenestra; pmf, promaxillary fenestra; pnf, posterior-most neurovascular foramen; scs, subcutaneous surface; sppm, subnarial process of premaxilla. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672013" httpUri="https://zenodo.org/record/3672013/files/figure.png" pageId="5" pageNumber="6">Fig. 4</figureCitation>
|
|
). Several possible long-healed
|
|
<typeStatus box="[1103,1152,970,989]" pageId="5" pageNumber="6">Type</typeStatus>
|
|
1 bite marks (sensu
|
|
<bibRefCitation author="Tanke, D. H. & Currie, P. J." journalOrPublisher="Gaia" pageId="5" pageNumber="6" pagination="184" part="15" refId="ref13453" refString="Tanke, D. H., Currie, P. J., 1998. Head-biting behavior in theropod dinosaurs: paleopathological evidence. Gaia 15, 167 e 184." title="Head-biting behavior in theropod dinosaurs: paleopathological evidence" type="journal article" year="1998">Tanke and Currie, 1998</bibRefCitation>
|
|
) adorn the subcutaneous surface more anteriorly.
|
|
</paragraph>
|
|
<paragraph blockId="5.[805,1475,969,1965]" pageId="5" pageNumber="6">
|
|
<emphasis bold="true" box="[837,890,1025,1044]" pageId="5" pageNumber="6">Jugal</emphasis>
|
|
-The jugal of
|
|
<collectionCode box="[1031,1076,1025,1044]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="5" pageNumber="6">TMP</collectionCode>
|
|
<date box="[1080,1164,1025,1044]" pageId="5" pageNumber="6" value="2010-05-07">2010.5.7</date>
|
|
is represented by a fragment of the suborbital region of the bone and a natural endocast of the jugal sinus (
|
|
<figureCitation box="[872,941,1081,1100]" captionStart="Fig" captionStartId="6.[113,138,1638,1652]" captionTargetBox="[280,1335,183,1607]" captionTargetId="figure@6.[276,1338,178,1608]" captionTargetPageId="6" captionText="Fig. 6. Comparison of right jugal (lateral view) and jugal sinus in tyrannosaurids. A) Thanatotheristes holotype (TMP 2010.5.7). B) Daspletosaurus torosus (TMP 2001.36.1).Black line represents position of transverse CT slice in the top right.Note the elliptical shape and extensive sinus inflation in the suborbital region cross section of Thanatotheristes, constrasting with the tear-drop shaped and constricted sinus of Daspletosaurus. Abbreviation: jse, jugal sinus endocast. Scale bars equal 3 cm." figureDoi="http://doi.org/10.5281/zenodo.3672017" httpUri="https://zenodo.org/record/3672017/files/figure.png" pageId="5" pageNumber="6">Fig. 6A</figureCitation>
|
|
and
|
|
<figureCitation box="[994,1060,1081,1100]" captionStart="Fig" captionStartId="3.[88,113,1453,1467]" captionTargetBox="[178,1387,179,1422]" captionTargetId="figure@3.[175,1388,178,1423]" captionTargetPageId="3" captionText="Fig. 3. Locality of Thanatotheristes degrootorum holotype (TMP 2010.5.7).A) Geographic location of the locality.The specimen was found 0.5 km upstream from the confluence of the Bow and Oldman rivers. B) Photograph of locality looking NNE. Specimen was found scattered over a lateral extent of 10 m on the shore of the river. The nearby cliff is composed of coal beds and interbedded sandstones and mudstones of the Taber Coal Zone of the Foremost Formation. C) Stratigraphic section of the exposures near the fossil locality. Bracket indicates likely stratigraphic provenance of specimen. Dotted line marks the extent of the exposures at the type locality; the Herronton Sandstone is exposed farther downstream." figureDoi="http://doi.org/10.5281/zenodo.3672011" httpUri="https://zenodo.org/record/3672011/files/figure.png" pageId="5" pageNumber="6">Fig. S3</figureCitation>
|
|
). Despite the fragmentary nature of this bone, the jugal exhibits several features that permit comparison with other tyrannosaurids.
|
|
</paragraph>
|
|
<paragraph blockId="5.[805,1475,969,1965]" pageId="5" pageNumber="6">
|
|
The orbital margin of the jugal is deeply concave and is inclined steeply anterodorsally towards the dorsoventrally-expanded maxillary ramus. The orbital margin is wide and has a rounded edge, resulting in an oval cross-section of the suborbital region (
|
|
<figureCitation box="[813,886,1276,1295]" captionStart="Fig" captionStartId="6.[113,138,1638,1652]" captionTargetBox="[280,1335,183,1607]" captionTargetId="figure@6.[276,1338,178,1608]" captionTargetPageId="6" captionText="Fig. 6. Comparison of right jugal (lateral view) and jugal sinus in tyrannosaurids. A) Thanatotheristes holotype (TMP 2010.5.7). B) Daspletosaurus torosus (TMP 2001.36.1).Black line represents position of transverse CT slice in the top right.Note the elliptical shape and extensive sinus inflation in the suborbital region cross section of Thanatotheristes, constrasting with the tear-drop shaped and constricted sinus of Daspletosaurus. Abbreviation: jse, jugal sinus endocast. Scale bars equal 3 cm." figureDoi="http://doi.org/10.5281/zenodo.3672017" httpUri="https://zenodo.org/record/3672017/files/figure.png" pageId="5" pageNumber="6">Fig. 6A</figureCitation>
|
|
). This morphology differs from that of most other tyrannosaurids (including
|
|
<taxonomicName box="[1017,1164,1303,1323]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1017,1164,1303,1323]" italics="true" pageId="5" pageNumber="6">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
), where the orbital margin has a tapered blade-like edge, resulting in a tear drop-shaped crosssection (
|
|
<figureCitation box="[892,962,1360,1379]" captionStart="Fig" captionStartId="6.[113,138,1638,1652]" captionTargetBox="[280,1335,183,1607]" captionTargetId="figure@6.[276,1338,178,1608]" captionTargetPageId="6" captionText="Fig. 6. Comparison of right jugal (lateral view) and jugal sinus in tyrannosaurids. A) Thanatotheristes holotype (TMP 2010.5.7). B) Daspletosaurus torosus (TMP 2001.36.1).Black line represents position of transverse CT slice in the top right.Note the elliptical shape and extensive sinus inflation in the suborbital region cross section of Thanatotheristes, constrasting with the tear-drop shaped and constricted sinus of Daspletosaurus. Abbreviation: jse, jugal sinus endocast. Scale bars equal 3 cm." figureDoi="http://doi.org/10.5281/zenodo.3672017" httpUri="https://zenodo.org/record/3672017/files/figure.png" pageId="5" pageNumber="6">Fig. 6B</figureCitation>
|
|
). In
|
|
<taxonomicName box="[1007,1104,1359,1379]" class="Reptilia" family="Tyrannosauridae" genus="Lythronax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1007,1104,1359,1379]" italics="true" pageId="5" pageNumber="6">Lythronax</emphasis>
|
|
</taxonomicName>
|
|
(
|
|
<collectionCode box="[1117,1188,1360,1379]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34862" lsid="urn:lsid:biocol.org:col:34862" name="Utah Museum of Natural History" pageId="5" pageNumber="6" type="Museum">UMNH</collectionCode>
|
|
<collectionCode box="[1195,1222,1360,1379]" pageId="5" pageNumber="6">VP</collectionCode>
|
|
20200) and some specimens of
|
|
<emphasis box="[902,1086,1386,1406]" italics="true" pageId="5" pageNumber="6">
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[902,1045,1386,1406]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
|
rex
|
|
</emphasis>
|
|
(
|
|
<collectionCode box="[1104,1151,1388,1407]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="5" pageNumber="6">TMP</collectionCode>
|
|
<date box="[1162,1246,1388,1407]" pageId="5" pageNumber="6" value="1981-06-01">1981.6.1</date>
|
|
), the jugal suborbital margin is rounded but narrow, differing from the wider margin of
|
|
<taxonomicName authority="Voris & Therrien & Zelenitsky & Brown, 2020" authorityName="Voris & Therrien & Zelenitsky & Brown" authorityYear="2020" box="[805,965,1442,1462]" class="Reptilia" family="Tyrannosauridae" genus="Thanatotheristes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="3" phylum="Chordata" rank="genus" status="gen. nov.">
|
|
<emphasis box="[805,965,1442,1462]" italics="true" pageId="5" pageNumber="6">Thanatotheristes</emphasis>
|
|
</taxonomicName>
|
|
.
|
|
</paragraph>
|
|
<paragraph blockId="5.[805,1475,969,1965]" pageId="5" pageNumber="6">
|
|
The jugal sinus is greatly inflated in
|
|
<taxonomicName authority="Voris & Therrien & Zelenitsky & Brown, 2020" authorityName="Voris & Therrien & Zelenitsky & Brown" authorityYear="2020" box="[1191,1347,1470,1490]" class="Reptilia" family="Tyrannosauridae" genus="Thanatotheristes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="3" phylum="Chordata" rank="genus" status="gen. nov.">
|
|
<emphasis box="[1191,1347,1470,1490]" italics="true" pageId="5" pageNumber="6">Thanatotheristes</emphasis>
|
|
</taxonomicName>
|
|
, presumably related to the rounder cross-section of the element. The sinus is of consistent depth within the suborbital region, possessing roughly parallel dorsal and ventral margins (
|
|
<figureCitation box="[1160,1228,1555,1574]" captionStart="Fig" captionStartId="6.[113,138,1638,1652]" captionTargetBox="[280,1335,183,1607]" captionTargetId="figure@6.[276,1338,178,1608]" captionTargetPageId="6" captionText="Fig. 6. Comparison of right jugal (lateral view) and jugal sinus in tyrannosaurids. A) Thanatotheristes holotype (TMP 2010.5.7). B) Daspletosaurus torosus (TMP 2001.36.1).Black line represents position of transverse CT slice in the top right.Note the elliptical shape and extensive sinus inflation in the suborbital region cross section of Thanatotheristes, constrasting with the tear-drop shaped and constricted sinus of Daspletosaurus. Abbreviation: jse, jugal sinus endocast. Scale bars equal 3 cm." figureDoi="http://doi.org/10.5281/zenodo.3672017" httpUri="https://zenodo.org/record/3672017/files/figure.png" pageId="5" pageNumber="6">Fig. 6A</figureCitation>
|
|
), in contrast to
|
|
<emphasis box="[1379,1474,1554,1574]" italics="true" pageId="5" pageNumber="6">Alioramus</emphasis>
|
|
(
|
|
<bibRefCitation author="Gold, M. E. L. & Brusatte, S. L. & Norell, M. A." box="[813,973,1583,1603]" journalOrPublisher="American Museum Novitates" pageId="5" pageNumber="6" pagination="46" part="3790" refId="ref11395" refString="Gold, M. E. L., Brusatte, S. L., Norell, M. A., 2013. The cranial pneumatic sinuses of the tyrannosaurid Alioramus (Dinosauria: Theropoda) and the evolution of cranial pneumaticity in theropod dinosaurs. American Museum Novitates 3790, 1 e 46." title="The cranial pneumatic sinuses of the tyrannosaurid Alioramus (Dinosauria: Theropoda) and the evolution of cranial pneumaticity in theropod dinosaurs" type="journal article" year="2013">Gold et al., 2013</bibRefCitation>
|
|
:fig. 5),
|
|
<taxonomicName box="[1050,1169,1582,1602]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1050,1169,1582,1602]" italics="true" pageId="5" pageNumber="6">Gorgosaurus</emphasis>
|
|
</taxonomicName>
|
|
(
|
|
<collectionCode box="[1182,1228,1583,1602]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="5" pageNumber="6">TMP</collectionCode>
|
|
<date box="[1235,1342,1583,1602]" pageId="5" pageNumber="6" value="2000-12-11">2000.12.11</date>
|
|
), and
|
|
<taxonomicName class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis italics="true" pageId="5" pageNumber="6">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
(
|
|
<figureCitation box="[901,970,1611,1630]" captionStart="Fig" captionStartId="6.[113,138,1638,1652]" captionTargetBox="[280,1335,183,1607]" captionTargetId="figure@6.[276,1338,178,1608]" captionTargetPageId="6" captionText="Fig. 6. Comparison of right jugal (lateral view) and jugal sinus in tyrannosaurids. A) Thanatotheristes holotype (TMP 2010.5.7). B) Daspletosaurus torosus (TMP 2001.36.1).Black line represents position of transverse CT slice in the top right.Note the elliptical shape and extensive sinus inflation in the suborbital region cross section of Thanatotheristes, constrasting with the tear-drop shaped and constricted sinus of Daspletosaurus. Abbreviation: jse, jugal sinus endocast. Scale bars equal 3 cm." figureDoi="http://doi.org/10.5281/zenodo.3672017" httpUri="https://zenodo.org/record/3672017/files/figure.png" pageId="5" pageNumber="6">Fig. 6B</figureCitation>
|
|
), where the suborbital region of the jugal sinus is dorsoventrally-constricted and hourglass-shaped in lateral view. The lateral surface of the jugal is inflated into a rounded bulge as in
|
|
<taxonomicName box="[805,949,1693,1713]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis box="[805,949,1693,1713]" italics="true" pageId="5" pageNumber="6">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
and most other tyrannosaurids, but unlike in
|
|
<taxonomicName class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|
<emphasis italics="true" pageId="5" pageNumber="6">Gorgosaurus</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<emphasis box="[917,1049,1721,1741]" italics="true" pageId="5" pageNumber="6">Bistahieversor</emphasis>
|
|
where it is raised into a shelf-like crest.
|
|
</paragraph>
|
|
<paragraph blockId="5.[805,1475,969,1965]" pageId="5" pageNumber="6">
|
|
<emphasis bold="true" box="[837,954,1750,1769]" pageId="5" pageNumber="6">Postorbital</emphasis>
|
|
-The postorbital of
|
|
<collectionCode box="[1160,1205,1751,1770]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="5" pageNumber="6">TMP</collectionCode>
|
|
<date box="[1211,1295,1750,1769]" pageId="5" pageNumber="6" value="2010-05-07">2010.5.7</date>
|
|
is represented by a fragment of the ventral ramus coming from the region immediately below the caudal inflection point at the dorsal end of the jugal contact (
|
|
<figureCitation box="[893,947,1834,1853]" captionStart="Fig" captionStartId="7.[88,113,1683,1697]" captionTargetBox="[96,751,182,1649]" captionTargetId="figure@7.[93,754,178,1653]" captionTargetPageId="7" captionText="Fig. 7. Right postorbital fragment of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view, demonstrating arced subcutaneous ridges, B) and medial view.Note prominent bracing ridge for jugal contact on the medial surface of the bone. Abbreviations: br, bracing ridge for jugal contact; djc, dorsal extent of jugal contact; scr, subcutaneous ridges; sr, smooth region along lateral surface posterior margin. Scale bar equals 2 cm." figureDoi="http://doi.org/10.5281/zenodo.3672019" httpUri="https://zenodo.org/record/3672019/files/figure.png" pageId="5" pageNumber="6">Fig. 7</figureCitation>
|
|
). In lateral view, the postorbital is sculptured with a series of parallel and arced ridges and grooves as in other tyrannosaurids. These ridges are separated from the caudal edge of the postorbital by a narrow surface (
|
|
<quantity box="[1140,1214,1918,1937]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="5" pageNumber="6" unit="mm" value="10.0">10 mm</quantity>
|
|
) of smooth bone (
|
|
<figureCitation box="[1405,1474,1918,1937]" captionStart="Fig" captionStartId="7.[88,113,1683,1697]" captionTargetBox="[96,751,182,1649]" captionTargetId="figure@7.[93,754,178,1653]" captionTargetPageId="7" captionText="Fig. 7. Right postorbital fragment of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view, demonstrating arced subcutaneous ridges, B) and medial view.Note prominent bracing ridge for jugal contact on the medial surface of the bone. Abbreviations: br, bracing ridge for jugal contact; djc, dorsal extent of jugal contact; scr, subcutaneous ridges; sr, smooth region along lateral surface posterior margin. Scale bar equals 2 cm." figureDoi="http://doi.org/10.5281/zenodo.3672019" httpUri="https://zenodo.org/record/3672019/files/figure.png" pageId="5" pageNumber="6">Fig. 7A</figureCitation>
|
|
and
|
|
<figureCitation box="[849,914,1946,1965]" captionStart="Fig" captionStartId="4.[113,138,1672,1686]" captionTargetBox="[202,1409,184,1640]" captionTargetId="figure@4.[200,1413,178,1642]" captionTargetPageId="4" captionText="Fig. 4. Right maxilla of Thanatotheristes holotype (TMP 2010.5.7) in lateral view with A) known fragments overlain on hypothesized reconstruction and B) subcutaneous surface magnified. The subcutaneous surface is strongly ornamented and possesses autapomorphic dorsoventral ridges (arrows). Extent of antorbital fossa indicated by black dash line. Large healed bite mark is indicated by white dash line. Abbreviations: aofo, antorbital fossa; mf, maxillary fenestra; pmf, promaxillary fenestra; pnf, posterior-most neurovascular foramen; scs, subcutaneous surface; sppm, subnarial process of premaxilla. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672013" httpUri="https://zenodo.org/record/3672013/files/figure.png" pageId="5" pageNumber="6">Fig. S4</figureCitation>
|
|
).
|
|
</paragraph>
|
|
<caption ID-DOI="http://doi.org/10.5281/zenodo.3672017" ID-Zenodo-Dep="3672017" httpUri="https://zenodo.org/record/3672017/files/figure.png" pageId="6" pageNumber="7" startId="6.[113,138,1638,1652]" targetBox="[280,1335,183,1607]" targetPageId="6">
|
|
<paragraph blockId="6.[113,1500,1637,1698]" pageId="6" pageNumber="7">
|
|
<emphasis bold="true" box="[113,161,1638,1652]" pageId="6" pageNumber="7">Fig. 6.</emphasis>
|
|
Comparison of right jugal (lateral view) and jugal sinus in tyrannosaurids. A)
|
|
<taxonomicName authority="Voris & Therrien & Zelenitsky & Brown, 2020" authorityName="Voris & Therrien & Zelenitsky & Brown" authorityYear="2020" box="[768,892,1637,1652]" class="Reptilia" family="Tyrannosauridae" genus="Thanatotheristes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="3" phylum="Chordata" rank="genus" status="gen. nov.">
|
|
<emphasis box="[768,892,1637,1652]" italics="true" pageId="6" pageNumber="7">Thanatotheristes</emphasis>
|
|
</taxonomicName>
|
|
holotype (TMP 2010.5.7). B)
|
|
<taxonomicName authorityName="Russell" authorityYear="1970" box="[1115,1287,1637,1652]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="torosus">
|
|
<emphasis box="[1115,1287,1637,1652]" italics="true" pageId="6" pageNumber="7">Daspletosaurus torosus</emphasis>
|
|
</taxonomicName>
|
|
(TMP 2001.36.1). Black line represents position of transverse CT slice in the top right. Note the elliptical shape and extensive sinus inflation in the suborbital region cross section of
|
|
<taxonomicName authority="Voris & Therrien & Zelenitsky & Brown, 2020" authorityName="Voris & Therrien & Zelenitsky & Brown" authorityYear="2020" box="[1274,1398,1660,1675]" class="Reptilia" family="Tyrannosauridae" genus="Thanatotheristes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="3" phylum="Chordata" rank="genus" status="gen. nov.">
|
|
<emphasis box="[1274,1398,1660,1675]" italics="true" pageId="6" pageNumber="7">Thanatotheristes</emphasis>
|
|
</taxonomicName>
|
|
, constrasting with the tear-drop shaped and constricted sinus of
|
|
<taxonomicName box="[519,633,1683,1698]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|
<emphasis box="[519,633,1683,1698]" italics="true" pageId="6" pageNumber="7">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
. Abbreviation: jse, jugal sinus endocast. Scale bars equal 3 cm.
|
|
</paragraph>
|
|
</caption>
|
|
<paragraph blockId="6.[113,783,1771,1958]" lastBlockId="6.[831,1501,1771,1958]" pageId="6" pageNumber="7">
|
|
In medial view, the contact surface for the jugal is braced anteriorly by a robust ridge that extends dorsally to the posterior margin of the ventral ramus (
|
|
<figureCitation box="[412,481,1827,1847]" captionStart="Fig" captionStartId="7.[88,113,1683,1697]" captionTargetBox="[96,751,182,1649]" captionTargetId="figure@7.[93,754,178,1653]" captionTargetPageId="7" captionText="Fig. 7. Right postorbital fragment of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view, demonstrating arced subcutaneous ridges, B) and medial view.Note prominent bracing ridge for jugal contact on the medial surface of the bone. Abbreviations: br, bracing ridge for jugal contact; djc, dorsal extent of jugal contact; scr, subcutaneous ridges; sr, smooth region along lateral surface posterior margin. Scale bar equals 2 cm." figureDoi="http://doi.org/10.5281/zenodo.3672019" httpUri="https://zenodo.org/record/3672019/files/figure.png" pageId="6" pageNumber="7">Fig. 7B</figureCitation>
|
|
). Whereas this morphology is seen in some specimens of
|
|
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[384,502,1854,1874]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|
<emphasis box="[384,502,1854,1874]" italics="true" pageId="6" pageNumber="7">Tarbosaurus</emphasis>
|
|
</taxonomicName>
|
|
(
|
|
<collectionCode box="[512,564,1855,1874]" country="Poland" httpUri="http://grbio.org/cool/yd4q-n37a" name="Zoological Institute of Paleobiology, Polish Academy of Sciences" pageId="6" pageNumber="7">ZPAL</collectionCode>
|
|
MgD I/4; see
|
|
<bibRefCitation author="Hurum, J. H. & Sabath, K." journalOrPublisher="Acta Palaeontologica Polonica" pageId="6" pageNumber="7" pagination="190" part="48" refId="ref11786" refString="Hurum, J. H., Sabath, K., 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontologica Polonica 48, 161 e 190." title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath, 2003</bibRefCitation>
|
|
:fig. 8) and
|
|
<emphasis box="[404,545,1882,1902]" italics="true" pageId="6" pageNumber="7">Teratophoneus</emphasis>
|
|
(
|
|
<collectionCode box="[558,629,1883,1902]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34862" lsid="urn:lsid:biocol.org:col:34862" name="Utah Museum of Natural History" pageId="6" pageNumber="7" type="Museum">UMNH</collectionCode>
|
|
<collectionCode box="[635,662,1883,1902]" pageId="6" pageNumber="7">VP</collectionCode>
|
|
16690), the bracing ridge usually terminates at mid-width on the ventral ramus in most tyrannosaurids, or extends to the posterior margin of the ventral ramus as an indistinct, narrow structure in others (e.g.
|
|
<taxonomicName box="[831,978,1798,1818]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|
<emphasis box="[831,978,1798,1818]" italics="true" pageId="6" pageNumber="7">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<collectionCode box="[990,1040,1800,1819]" country="Canada" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34935" lsid="urn:lsid:biocol.org:col:34935" name="Canadian Museum of Nature" pageId="6" pageNumber="7" type="Museum">CMN</collectionCode>
|
|
8506).
|
|
</paragraph>
|
|
<paragraph blockId="6.[831,1501,1771,1958]" lastBlockId="7.[805,1475,178,1956]" lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7">
|
|
<emphasis bold="true" box="[863,968,1827,1846]" pageId="6" pageNumber="7">Skull roof</emphasis>
|
|
-The skull roof of
|
|
<collectionCode box="[1162,1207,1828,1847]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="6" pageNumber="7">TMP</collectionCode>
|
|
<date box="[1213,1297,1827,1846]" pageId="6" pageNumber="7" value="2010-05-07">2010.5.7</date>
|
|
consists of a partial left frontal preserved in articulation with a narrow (~
|
|
<quantity box="[1376,1435,1855,1875]" metricMagnitude="-3" metricUnit="m" metricValue="5.0" pageId="6" pageNumber="7" unit="mm" value="5.0">5 mm</quantity>
|
|
wide) fragment of the right frontal, the posteroventral portion of the prefrontal, and small fragments of the postorbital and caudal ramus of the lacrimal (
|
|
<figureCitation box="[994,1062,1939,1958]" captionStart="Fig" captionStartId="9.[88,113,1813,1827]" captionTargetBox="[94,753,182,1782]" captionTargetId="figure@9.[93,754,178,1783]" captionTargetPageId="9" captionText="Fig. 8. Comparison of left frontalin dorsalviewin tyrannosaurids.A)Thanatotheristes TMP 2010.5.7. White dash line represents nondiagnostic postorbital fragment attached to the frontal. B) Subcutaneous surface of frontal of Daspletosaurus torosus TMP 2001.036.0001. Abbreviations: apc, anterior postorbital contact expansion; aesc, autapomorphic anterior extension of sagittal crest; ifs, interfrontal suture;ls, lacrimal socket; np, nasal process; os, orbital slot; PA, parietal; PF, prefrontal; pfl, prefrontolacrimal process; pfn, prefrontonasal process; po, postorbital fragment; sc, sagittal crest; stf, supratemporal fossa; str, supratemporal ridge. Scale bars equal 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672023" httpUri="https://zenodo.org/record/3672023/files/figure.png" pageId="6" pageNumber="7">Fig. 8A</figureCitation>
|
|
and
|
|
<figureCitation box="[1115,1180,1939,1958]" captionStart="Fig" captionStartId="5.[88,113,874,888]" captionTargetBox="[329,1235,181,841]" captionTargetId="figure@5.[326,1236,178,844]" captionTargetPageId="5" captionText="Fig. 5. Left maxilla of referred specimen of Thanatotheristes (TMP 2018.16.1) in lateral view. Note the presence of pronounced vertical ridges (arrows) on subcutaneous surface, an autapomorphy of the taxon. Abbreviations: dvr, dorsoventral ridges; sc-afos, subcutaneous-antorbital fossa boundary. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672015" httpUri="https://zenodo.org/record/3672015/files/figure.png" pageId="6" pageNumber="7">Fig. S5</figureCitation>
|
|
). The left frontal is missing the prefrontal socket, nasal ramus, most of the sagittal crest, laterosphenoid contact surface, and most of the cerebral fossa on the ventral surface. The contact surface for the postorbital is deep, consistent with that of other derived tyrannosaurines (e.g.,
|
|
<taxonomicName box="[805,952,178,198]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
|
<emphasis box="[805,952,178,198]" italics="true" pageId="7" pageNumber="8">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[962,1080,178,198]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
|
<emphasis box="[962,1080,178,198]" italics="true" pageId="7" pageNumber="8">Tarbosaurus</emphasis>
|
|
</taxonomicName>
|
|
, and
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1133,1276,178,198]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1133,1276,178,198]" italics="true" pageId="7" pageNumber="8">Tyrannosaurus</emphasis>
|
|
</taxonomicName>
|
|
) but deeper than in other tyrannosaurids (e.g.,
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1171,1304,206,226]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1171,1304,206,226]" italics="true" pageId="7" pageNumber="8">Albertosaurus</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<taxonomicName authority=", Teratophoneus" authorityName="Teratophoneus" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1348,1470,206,226]" italics="true" pageId="7" pageNumber="8">Gorgosaurus</emphasis>
|
|
,
|
|
<emphasis box="[805,946,234,254]" italics="true" pageId="7" pageNumber="8">Teratophoneus</emphasis>
|
|
</taxonomicName>
|
|
).
|
|
</paragraph>
|
|
<paragraph blockId="7.[805,1475,178,1956]" pageId="7" pageNumber="8">
|
|
The dorsal surface of the forehead region (sensu
|
|
<bibRefCitation author="Carr, T. D. & Williamson, T. E." journalOrPublisher="Zoological Journal of the Linnean Society" pageId="7" pageNumber="8" pagination="523" part="142" refId="ref10143" refString="Carr, T. D., Williamson, T. E., 2004. Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America. Zoological Journal of the Linnean Society 142, 479 e 523." title="Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America" type="journal article" year="2004">Carr and Williamson, 2004</bibRefCitation>
|
|
) in
|
|
<collectionCode box="[1033,1078,291,310]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
<date box="[1088,1172,291,310]" pageId="7" pageNumber="8" value="2010-05-07">2010.5.7</date>
|
|
slopes medially towards the interfrontal suture as in most mature tyrannosaurines, except for
|
|
<emphasis box="[805,946,345,365]" italics="true" pageId="7" pageNumber="8">Teratophoneus</emphasis>
|
|
, where it slopes away from the interfrontal suture, as well as in
|
|
<taxonomicName box="[915,1011,373,393]" class="Reptilia" family="Tyrannosauridae" genus="Lythronax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
|
<emphasis box="[915,1011,373,393]" italics="true" pageId="7" pageNumber="8">Lythronax</emphasis>
|
|
</taxonomicName>
|
|
and some specimens of
|
|
<taxonomicName box="[1272,1419,373,393]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1272,1419,373,393]" italics="true" pageId="7" pageNumber="8">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
(e.g.,
|
|
<collectionCode box="[805,850,403,422]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
1980.16.924), where it is flat. The dorsomedial surface is raised into a low, broad, and rounded extension of the sagittal crest located anterior to the supratemporal ridge (
|
|
<figureCitation box="[1254,1321,458,477]" captionStart="Fig" captionStartId="9.[88,113,1813,1827]" captionTargetBox="[94,753,182,1782]" captionTargetId="figure@9.[93,754,178,1783]" captionTargetPageId="9" captionText="Fig. 8. Comparison of left frontalin dorsalviewin tyrannosaurids.A)Thanatotheristes TMP 2010.5.7. White dash line represents nondiagnostic postorbital fragment attached to the frontal. B) Subcutaneous surface of frontal of Daspletosaurus torosus TMP 2001.036.0001. Abbreviations: apc, anterior postorbital contact expansion; aesc, autapomorphic anterior extension of sagittal crest; ifs, interfrontal suture;ls, lacrimal socket; np, nasal process; os, orbital slot; PA, parietal; PF, prefrontal; pfl, prefrontolacrimal process; pfn, prefrontonasal process; po, postorbital fragment; sc, sagittal crest; stf, supratemporal fossa; str, supratemporal ridge. Scale bars equal 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672023" httpUri="https://zenodo.org/record/3672023/files/figure.png" pageId="7" pageNumber="8">Fig. 8A</figureCitation>
|
|
and
|
|
<figureCitation box="[1371,1435,458,477]" captionStart="Fig" captionStartId="5.[88,113,874,888]" captionTargetBox="[329,1235,181,841]" captionTargetId="figure@5.[326,1236,178,844]" captionTargetPageId="5" captionText="Fig. 5. Left maxilla of referred specimen of Thanatotheristes (TMP 2018.16.1) in lateral view. Note the presence of pronounced vertical ridges (arrows) on subcutaneous surface, an autapomorphy of the taxon. Abbreviations: dvr, dorsoventral ridges; sc-afos, subcutaneous-antorbital fossa boundary. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672015" httpUri="https://zenodo.org/record/3672015/files/figure.png" pageId="7" pageNumber="8">Fig. S5</figureCitation>
|
|
). In all other tyrannosaurids, the sagittal crest does not extend beyond the supratemporal ridge. The supratemporal ridge in
|
|
<collectionCode box="[1339,1384,514,533]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
<date box="[1390,1474,514,533]" pageId="7" pageNumber="8" value="2010-05-07">2010.5.7</date>
|
|
is sinuous and primarily mediolaterally-oriented, extending from the sagittal crest to the anterior margin of the postorbital contact. Whereas the supratemporal ridge is also sinuous and mediolaterally-oriented in most tyrannosaurids, it is sinuous and anterolaterally-oriented in
|
|
<taxonomicName authority="(Carr et al., 2017)" baseAuthorityName="Carr" baseAuthorityYear="2017" box="[1074,1470,652,673]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="horneri">
|
|
<emphasis box="[1074,1297,652,672]" italics="true" pageId="7" pageNumber="8">Daspletosaurus horneri</emphasis>
|
|
(
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[1310,1461,653,673]" journalOrPublisher="Scientific Reports" pageId="7" pageNumber="8" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
)
|
|
</taxonomicName>
|
|
, arcuate and laterally-to anterolaterally-oriented in
|
|
<taxonomicName authority="(Carr et al., 2017)" baseAuthorityName="Carr" baseAuthorityYear="2017" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="torosus">
|
|
<emphasis box="[1321,1417,680,700]" italics="true" pageId="7" pageNumber="8">D. torosus</emphasis>
|
|
(
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." journalOrPublisher="Scientific Reports" pageId="7" pageNumber="8" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
)
|
|
</taxonomicName>
|
|
, and straight and mediolaterally-oriented in
|
|
<emphasis italics="true" pageId="7" pageNumber="8">
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
|
rex
|
|
</emphasis>
|
|
(
|
|
<collectionCode box="[918,988,737,756]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34925" lsid="urn:lsid:biocol.org:col:34925" name="American Museum of Natural History" pageId="7" pageNumber="8">AMNH</collectionCode>
|
|
<collectionCode box="[994,1045,737,756]" pageId="7" pageNumber="8">FARB</collectionCode>
|
|
5117).
|
|
</paragraph>
|
|
<paragraph blockId="7.[805,1475,178,1956]" pageId="7" pageNumber="8">
|
|
The dorsal margin of the lacrimal slot on the frontal of
|
|
<collectionCode box="[1429,1474,766,785]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
<date box="[805,889,793,812]" pageId="7" pageNumber="8" value="2010-05-07">2010.5.7</date>
|
|
is oriented at an angle of approximately 60 to the interfrontal suture in dorsal view (
|
|
<figureCitation box="[1116,1190,821,840]" captionStart="Fig" captionStartId="9.[88,113,1813,1827]" captionTargetBox="[94,753,182,1782]" captionTargetId="figure@9.[93,754,178,1783]" captionTargetPageId="9" captionText="Fig. 8. Comparison of left frontalin dorsalviewin tyrannosaurids.A)Thanatotheristes TMP 2010.5.7. White dash line represents nondiagnostic postorbital fragment attached to the frontal. B) Subcutaneous surface of frontal of Daspletosaurus torosus TMP 2001.036.0001. Abbreviations: apc, anterior postorbital contact expansion; aesc, autapomorphic anterior extension of sagittal crest; ifs, interfrontal suture;ls, lacrimal socket; np, nasal process; os, orbital slot; PA, parietal; PF, prefrontal; pfl, prefrontolacrimal process; pfn, prefrontonasal process; po, postorbital fragment; sc, sagittal crest; stf, supratemporal fossa; str, supratemporal ridge. Scale bars equal 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672023" httpUri="https://zenodo.org/record/3672023/files/figure.png" pageId="7" pageNumber="8">Fig. 8A</figureCitation>
|
|
). In contrast, this margin is mediolaterally-oriented in most other tyrannosaurids, except in adult
|
|
<emphasis box="[866,1048,875,895]" italics="true" pageId="7" pageNumber="8">
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[866,1009,875,895]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
|
rex
|
|
</emphasis>
|
|
(e.g.,
|
|
<collectionCode box="[1111,1178,877,896]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34925" lsid="urn:lsid:biocol.org:col:34925" name="American Museum of Natural History" pageId="7" pageNumber="8">AMNH</collectionCode>
|
|
5027,
|
|
<collectionCode box="[1250,1317,877,896]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34925" lsid="urn:lsid:biocol.org:col:34925" name="American Museum of Natural History" pageId="7" pageNumber="8">AMNH</collectionCode>
|
|
5117) where it forms an angle of approximately 70.
|
|
</paragraph>
|
|
<paragraph blockId="7.[805,1475,178,1956]" pageId="7" pageNumber="8">
|
|
In ventral view, the prefrontal-frontal contact is wider and more complex in
|
|
<collectionCode box="[920,965,961,980]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
<date box="[970,1054,960,979]" pageId="7" pageNumber="8" value="2010-05-07">2010.5.7</date>
|
|
than in all other tyrannosaurids. Whereas the prefrontal typically articulates with the frontal via a single medial prong in tyrannosaurids, it articulates via two prongs, a large medial prong and a shorter, lateral prong, in
|
|
<collectionCode box="[1336,1381,1045,1064]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
<date box="[1390,1474,1044,1063]" pageId="7" pageNumber="8" value="2010-05-07">2010.5.7</date>
|
|
(
|
|
<figureCitation box="[813,876,1072,1091]" captionStart="Fig" captionStartId="5.[88,113,874,888]" captionTargetBox="[329,1235,181,841]" captionTargetId="figure@5.[326,1236,178,844]" captionTargetPageId="5" captionText="Fig. 5. Left maxilla of referred specimen of Thanatotheristes (TMP 2018.16.1) in lateral view. Note the presence of pronounced vertical ridges (arrows) on subcutaneous surface, an autapomorphy of the taxon. Abbreviations: dvr, dorsoventral ridges; sc-afos, subcutaneous-antorbital fossa boundary. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672015" httpUri="https://zenodo.org/record/3672015/files/figure.png" pageId="7" pageNumber="8">Fig. S5</figureCitation>
|
|
). In anterior view, the prefrontal of
|
|
<collectionCode box="[1232,1277,1072,1091]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
<date box="[1282,1365,1072,1091]" pageId="7" pageNumber="8" value="2010-05-07">2010.5.7</date>
|
|
is strongly arced dorsomedially, as in
|
|
<taxonomicName box="[1073,1220,1099,1119]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1073,1220,1099,1119]" italics="true" pageId="7" pageNumber="8">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
where it is related to the presence of a broad, tear drop-shaped prefrontal (
|
|
<figureCitation box="[1355,1431,1128,1147]" captionStart="Fig" captionStartId="9.[88,113,1813,1827]" captionTargetBox="[94,753,182,1782]" captionTargetId="figure@9.[93,754,178,1783]" captionTargetPageId="9" captionText="Fig. 8. Comparison of left frontalin dorsalviewin tyrannosaurids.A)Thanatotheristes TMP 2010.5.7. White dash line represents nondiagnostic postorbital fragment attached to the frontal. B) Subcutaneous surface of frontal of Daspletosaurus torosus TMP 2001.036.0001. Abbreviations: apc, anterior postorbital contact expansion; aesc, autapomorphic anterior extension of sagittal crest; ifs, interfrontal suture;ls, lacrimal socket; np, nasal process; os, orbital slot; PA, parietal; PF, prefrontal; pfl, prefrontolacrimal process; pfn, prefrontonasal process; po, postorbital fragment; sc, sagittal crest; stf, supratemporal fossa; str, supratemporal ridge. Scale bars equal 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672023" httpUri="https://zenodo.org/record/3672023/files/figure.png" pageId="7" pageNumber="8">Fig. 8B</figureCitation>
|
|
). In contrast, tyrannosaurid taxa in which the arcing of the prefrontal is subtle, extending more dorsally than medially (e.g., albertosaurines,
|
|
<taxonomicName authority=", Teratophoneus" authorityName="Teratophoneus" box="[864,1121,1210,1231]" class="Reptilia" family="Tyrannosauridae" genus="Lythronax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
|
<emphasis box="[864,960,1210,1230]" italics="true" pageId="7" pageNumber="8">Lythronax</emphasis>
|
|
,
|
|
<emphasis box="[980,1121,1210,1230]" italics="true" pageId="7" pageNumber="8">Teratophoneus</emphasis>
|
|
</taxonomicName>
|
|
), possess a narrow, lens-shaped prefrontal. As such, it can be surmised that
|
|
<collectionCode box="[1235,1280,1240,1259]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
<date box="[1285,1369,1239,1258]" pageId="7" pageNumber="8" value="2010-05-07">2010.5.7</date>
|
|
possessed a broad, tear drop-shaped prefrontal.
|
|
</paragraph>
|
|
<paragraph blockId="7.[805,1475,178,1956]" pageId="7" pageNumber="8">
|
|
<emphasis bold="true" box="[837,939,1295,1314]" pageId="7" pageNumber="8">Braincase</emphasis>
|
|
-The braincase of
|
|
<collectionCode box="[1135,1180,1295,1314]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
<date box="[1188,1271,1295,1314]" pageId="7" pageNumber="8" value="2010-05-07">2010.5.7</date>
|
|
is represented by a single fragment of the right laterosphenoid, specifically the antotic region and medial portion of the capitate process, with small fragments of the parietal and right frontal preserved in articulation (
|
|
<figureCitation box="[813,878,1407,1426]" captionStart="Fig" captionStartId="6.[113,138,1638,1652]" captionTargetBox="[280,1335,183,1607]" captionTargetId="figure@6.[276,1338,178,1608]" captionTargetPageId="6" captionText="Fig. 6. Comparison of right jugal (lateral view) and jugal sinus in tyrannosaurids. A) Thanatotheristes holotype (TMP 2010.5.7). B) Daspletosaurus torosus (TMP 2001.36.1).Black line represents position of transverse CT slice in the top right.Note the elliptical shape and extensive sinus inflation in the suborbital region cross section of Thanatotheristes, constrasting with the tear-drop shaped and constricted sinus of Daspletosaurus. Abbreviation: jse, jugal sinus endocast. Scale bars equal 3 cm." figureDoi="http://doi.org/10.5281/zenodo.3672017" httpUri="https://zenodo.org/record/3672017/files/figure.png" pageId="7" pageNumber="8">Fig. S6</figureCitation>
|
|
).
|
|
</paragraph>
|
|
<paragraph blockId="7.[805,1475,178,1956]" pageId="7" pageNumber="8">
|
|
The antotic region of
|
|
<collectionCode box="[1069,1114,1435,1454]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
<date box="[1124,1208,1434,1453]" pageId="7" pageNumber="8" value="2010-05-07">2010.5.7</date>
|
|
is broader than in other tyrannosaurids (
|
|
<figureCitation box="[971,1036,1462,1481]" captionStart="Fig" captionStartId="6.[113,138,1638,1652]" captionTargetBox="[280,1335,183,1607]" captionTargetId="figure@6.[276,1338,178,1608]" captionTargetPageId="6" captionText="Fig. 6. Comparison of right jugal (lateral view) and jugal sinus in tyrannosaurids. A) Thanatotheristes holotype (TMP 2010.5.7). B) Daspletosaurus torosus (TMP 2001.36.1).Black line represents position of transverse CT slice in the top right.Note the elliptical shape and extensive sinus inflation in the suborbital region cross section of Thanatotheristes, constrasting with the tear-drop shaped and constricted sinus of Daspletosaurus. Abbreviation: jse, jugal sinus endocast. Scale bars equal 3 cm." figureDoi="http://doi.org/10.5281/zenodo.3672017" httpUri="https://zenodo.org/record/3672017/files/figure.png" pageId="7" pageNumber="8">Fig. S6</figureCitation>
|
|
). The antotic crest descends ventrally from the capitate process as a prominent, weakly rugose ridge that bifurcates to form a second, more subtle and posteriorly-extending ridge (leading to the otosphenoidal crest in some tyrannosaurids:
|
|
<bibRefCitation author="Bever, G. S. & Brusatte, S. L. & Carr, T. D. & Xu, X. & Balanoff, A. M. & Norell, M. A." box="[805,1001,1574,1593]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="7" pageNumber="8" pagination="72" part="376" refId="ref9397" refString="Bever, G. S., Brusatte, S. L., Carr, T. D., Xu, X., Balanoff, A. M., Norell, M. A., 2013. The braincase anatomy of the Late Cretaceous dinosaur Alioramus (Theropoda: Tyrannosauroidea). Bulletin of the American Museum of Natural History 376, 1 e 72." title="The braincase anatomy of the Late Cretaceous dinosaur Alioramus (Theropoda: Tyrannosauroidea)" type="journal article" year="2013">Bever et al., 2013</bibRefCitation>
|
|
;
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[1018,1195,1574,1593]" journalOrPublisher="Scientific Reports" pageId="7" pageNumber="8" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
). The morphology of the posteriorly-extending ridge contrasts with that of
|
|
<taxonomicName box="[1322,1469,1601,1621]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1322,1469,1601,1621]" italics="true" pageId="7" pageNumber="8">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
, in which it is raised into a wide shelf that extends laterally to the edge of the dorsotemporal fenestra (
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[1175,1333,1658,1677]" journalOrPublisher="Scientific Reports" pageId="7" pageNumber="8" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
).
|
|
</paragraph>
|
|
<caption ID-DOI="http://doi.org/10.5281/zenodo.3672019" ID-Zenodo-Dep="3672019" httpUri="https://zenodo.org/record/3672019/files/figure.png" pageId="7" pageNumber="8" startId="7.[88,113,1683,1697]" targetBox="[96,751,182,1649]" targetPageId="7">
|
|
<paragraph blockId="7.[88,759,1682,1812]" pageId="7" pageNumber="8">
|
|
<emphasis bold="true" box="[88,139,1683,1697]" pageId="7" pageNumber="8">Fig. 7.</emphasis>
|
|
Right postorbital fragment of
|
|
<taxonomicName authority="Voris & Therrien & Zelenitsky & Brown, 2020" authorityName="Voris & Therrien & Zelenitsky & Brown" authorityYear="2020" box="[395,519,1682,1697]" class="Reptilia" family="Tyrannosauridae" genus="Thanatotheristes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="3" phylum="Chordata" rank="genus" status="gen. nov.">
|
|
<emphasis box="[395,519,1682,1697]" italics="true" pageId="7" pageNumber="8">Thanatotheristes</emphasis>
|
|
</taxonomicName>
|
|
holotype (TMP 2010.5.7). A) Specimen in lateral view, demonstrating arced subcutaneous ridges, B) and medial view. Note prominent bracing ridge for jugal contact on the medial surface of the bone. Abbreviations: br, bracing ridge for jugal contact; djc, dorsal extent of jugal contact; scr, subcutaneous ridges; sr, smooth region along lateral surface posterior margin. Scale bar equals 2 cm.
|
|
</paragraph>
|
|
</caption>
|
|
<paragraph blockId="7.[805,1475,178,1956]" pageId="7" pageNumber="8">
|
|
<emphasis bold="true" box="[837,935,1685,1704]" pageId="7" pageNumber="8">Quadrate</emphasis>
|
|
-The right quadrate of
|
|
<collectionCode box="[1184,1229,1686,1705]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
<date box="[1237,1321,1685,1704]" pageId="7" pageNumber="8" value="2010-05-07">2010.5.7</date>
|
|
is represented by a fragmentary mandibular ramus that includes a complete medial condyle and the medial half of the lateral condyle (
|
|
<figureCitation box="[1380,1432,1741,1761]" captionStart="Fig" captionStartId="9.[805,830,875,889]" captionTargetBox="[815,1469,182,843]" captionTargetId="figure@9.[810,1474,178,846]" captionTargetPageId="9" captionText="Fig. 9. Articulated right quadrate and surangular of Thanatotheristes holotype in lateral view. Abbreviations: anc, angular contact surface; ls, lateral shelf; ngf, non-glenoid fossa; pp, pneumatic pocket; qu, quadrate; sf, surangular foramen. Scale bar equals 2 cm." figureDoi="http://doi.org/10.5281/zenodo.3672021" httpUri="https://zenodo.org/record/3672021/files/figure.png" pageId="7" pageNumber="8">Fig. 9</figureCitation>
|
|
and
|
|
<figureCitation box="[805,872,1769,1788]" captionStart="Fig" captionStartId="7.[88,113,1683,1697]" captionTargetBox="[96,751,182,1649]" captionTargetId="figure@7.[93,754,178,1653]" captionTargetPageId="7" captionText="Fig. 7. Right postorbital fragment of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view, demonstrating arced subcutaneous ridges, B) and medial view.Note prominent bracing ridge for jugal contact on the medial surface of the bone. Abbreviations: br, bracing ridge for jugal contact; djc, dorsal extent of jugal contact; scr, subcutaneous ridges; sr, smooth region along lateral surface posterior margin. Scale bar equals 2 cm." figureDoi="http://doi.org/10.5281/zenodo.3672019" httpUri="https://zenodo.org/record/3672019/files/figure.png" pageId="7" pageNumber="8">Fig. S7</figureCitation>
|
|
). The quadrate pneumatic foramen, located on the ventral anterior face of the quadrate, is ventrally and laterally bounded by a shallow rectangular fossa. The morphology of this fossa is consistent with that of most tyrannosaurids, except in
|
|
<taxonomicName box="[1284,1406,1852,1872]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1284,1406,1852,1872]" italics="true" pageId="7" pageNumber="8">Gorgosaurus</emphasis>
|
|
</taxonomicName>
|
|
where the ventral margin of the fossa inflects dorsally and converges with the lateral wall of the pneumatopore to form a triangular secondary fossa (e.g.
|
|
<collectionCode box="[910,955,1937,1956]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="7" pageNumber="8">TMP</collectionCode>
|
|
1991.36.500).
|
|
</paragraph>
|
|
<paragraph blockId="8.[113,784,179,1984]" pageId="8" pageNumber="9">
|
|
<emphasis bold="true" box="[145,229,179,198]" pageId="8" pageNumber="9">Dentary</emphasis>
|
|
-The anterior ramus of both dentaries are preserved in
|
|
<collectionCode box="[113,158,208,227]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[163,247,207,226]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
. The left bone preserves the first seven alveoli (
|
|
<figureCitation box="[712,775,207,226]" captionStart="Fig" captionStartId="9.[88,113,1813,1827]" captionTargetBox="[94,753,182,1782]" captionTargetId="figure@9.[93,754,178,1783]" captionTargetPageId="9" captionText="Fig. 8. Comparison of left frontalin dorsalviewin tyrannosaurids.A)Thanatotheristes TMP 2010.5.7. White dash line represents nondiagnostic postorbital fragment attached to the frontal. B) Subcutaneous surface of frontal of Daspletosaurus torosus TMP 2001.036.0001. Abbreviations: apc, anterior postorbital contact expansion; aesc, autapomorphic anterior extension of sagittal crest; ifs, interfrontal suture;ls, lacrimal socket; np, nasal process; os, orbital slot; PA, parietal; PF, prefrontal; pfl, prefrontolacrimal process; pfn, prefrontonasal process; po, postorbital fragment; sc, sagittal crest; stf, supratemporal fossa; str, supratemporal ridge. Scale bars equal 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672023" httpUri="https://zenodo.org/record/3672023/files/figure.png" pageId="8" pageNumber="9">Fig. S8</figureCitation>
|
|
) and the right preserves the first 12 alveoli (
|
|
<figureCitation box="[550,613,235,254]" captionStart="Fig" captionStartId="10.[113,138,1695,1709]" captionTargetBox="[203,1411,182,1665]" captionTargetId="figure@10.[200,1413,178,1666]" captionTargetPageId="10" captionText="Fig. 10. Right dentary of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view. B) Specimen in dorsal view. Numbers correspond to dentary alveolus position. Comparison of dentary symphyseal region between C) Thanatotheristes and D) Daspletosaurus torosus (TMP 2001.36.1). Note the absence of bony projections on the symphyseal surface and a rugose knob ventral to Meckelian foramen in Thanatotheristes contrasting with Daspletosaurus. Abbreviations: apr, anteroposterior ridges; bp, bony projections; dc, dentary chin; mf, meckelian foramen; mg, meckelian groove; rk, rugose knob. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672025" httpUri="https://zenodo.org/record/3672025/files/figure.png" pageId="8" pageNumber="9">Fig. 10</figureCitation>
|
|
). Two additional fragments of the right dentary are also present, one of the posterior alveolar margin that preserves three additional alveoli (resulting in a total tooth count of 15 teeth) and the other of the splenial joint surface (
|
|
<figureCitation box="[204,273,347,366]" captionStart="Fig" captionStartId="9.[805,830,875,889]" captionTargetBox="[815,1469,182,843]" captionTargetId="figure@9.[810,1474,178,846]" captionTargetPageId="9" captionText="Fig. 9. Articulated right quadrate and surangular of Thanatotheristes holotype in lateral view. Abbreviations: anc, angular contact surface; ls, lateral shelf; ngf, non-glenoid fossa; pp, pneumatic pocket; qu, quadrate; sf, surangular foramen. Scale bar equals 2 cm." figureDoi="http://doi.org/10.5281/zenodo.3672021" httpUri="https://zenodo.org/record/3672021/files/figure.png" pageId="8" pageNumber="9">Fig. S9</figureCitation>
|
|
). The dorsal margin of both dentaries has been moderately crushed, such that many teeth are broken and fragmentary. Deformation to the main body of the dentaries, however, appears minimal as the Meckelian canal is circular in cross section.
|
|
</paragraph>
|
|
<paragraph blockId="8.[113,784,179,1984]" pageId="8" pageNumber="9">
|
|
The dentary of
|
|
<collectionCode box="[297,342,459,478]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[348,432,458,477]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
is nearly of the same depth as that of the largest
|
|
<taxonomicName box="[247,394,485,505]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[247,394,485,505]" italics="true" pageId="8" pageNumber="9">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
individuals known but much narrower than that of similar-sized or larger individuals (
|
|
<figureCitation box="[583,645,514,533]" captionStart="Fig" captionStartId="10.[113,138,1695,1709]" captionTargetBox="[203,1411,182,1665]" captionTargetId="figure@10.[200,1413,178,1666]" captionTargetPageId="10" captionText="Fig. 10. Right dentary of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view. B) Specimen in dorsal view. Numbers correspond to dentary alveolus position. Comparison of dentary symphyseal region between C) Thanatotheristes and D) Daspletosaurus torosus (TMP 2001.36.1). Note the absence of bony projections on the symphyseal surface and a rugose knob ventral to Meckelian foramen in Thanatotheristes contrasting with Daspletosaurus. Abbreviations: apr, anteroposterior ridges; bp, bony projections; dc, dentary chin; mf, meckelian foramen; mg, meckelian groove; rk, rugose knob. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672025" httpUri="https://zenodo.org/record/3672025/files/figure.png" pageId="8" pageNumber="9">Fig. 10</figureCitation>
|
|
). The dentary chin (sensu
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[238,398,542,561]" journalOrPublisher="Scientific Reports" pageId="8" pageNumber="9" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
), a subangular to subrounded prominence on the anteroventral margin of the dentary, is located ventral to the third interdental plate as in some
|
|
<taxonomicName box="[519,666,596,616]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[519,666,596,616]" italics="true" pageId="8" pageNumber="9">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
individuals (ventral to the third alveolus in other
|
|
<taxonomicName box="[508,655,624,644]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[508,655,624,644]" italics="true" pageId="8" pageNumber="9">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
individuals) and
|
|
<emphasis box="[163,304,652,672]" italics="true" pageId="8" pageNumber="9">Teratophoneus</emphasis>
|
|
, but ventral to the fourth alveolus in other tyrannosaurids (
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[280,438,681,701]" journalOrPublisher="Scientific Reports" pageId="8" pageNumber="9" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
). The dentary chin is more prominent in
|
|
<collectionCode box="[198,243,710,729]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[249,333,709,728]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
than in either
|
|
<emphasis box="[484,619,708,728]" italics="true" pageId="8" pageNumber="9">Bistahieversor</emphasis>
|
|
,
|
|
<taxonomicName box="[630,777,708,728]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[630,777,708,728]" italics="true" pageId="8" pageNumber="9">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
, or
|
|
<taxonomicName box="[141,237,736,756]" class="Reptilia" family="Tyrannosauridae" genus="Lythronax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[141,237,736,756]" italics="true" pageId="8" pageNumber="9">Lythronax</emphasis>
|
|
</taxonomicName>
|
|
, being similar to that observed in albertosaurines and
|
|
<emphasis box="[113,254,764,784]" italics="true" pageId="8" pageNumber="9">Teratophoneus</emphasis>
|
|
. The anterior margin of the dentary is straight and inclined anterodorsally at approximately 45, as in albertosaurines and
|
|
<emphasis box="[160,301,820,840]" italics="true" pageId="8" pageNumber="9">Teratophoneus</emphasis>
|
|
, but inflects dorsally at the anterior tip of the bone to meet the alveolar margin at an angle of approximately 90 (
|
|
<figureCitation box="[121,183,877,896]" captionStart="Fig" captionStartId="10.[113,138,1695,1709]" captionTargetBox="[203,1411,182,1665]" captionTargetId="figure@10.[200,1413,178,1666]" captionTargetPageId="10" captionText="Fig. 10. Right dentary of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view. B) Specimen in dorsal view. Numbers correspond to dentary alveolus position. Comparison of dentary symphyseal region between C) Thanatotheristes and D) Daspletosaurus torosus (TMP 2001.36.1). Note the absence of bony projections on the symphyseal surface and a rugose knob ventral to Meckelian foramen in Thanatotheristes contrasting with Daspletosaurus. Abbreviations: apr, anteroposterior ridges; bp, bony projections; dc, dentary chin; mf, meckelian foramen; mg, meckelian groove; rk, rugose knob. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672025" httpUri="https://zenodo.org/record/3672025/files/figure.png" pageId="8" pageNumber="9">Fig.10</figureCitation>
|
|
), as in derived tyrannosaurines (
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[508,660,877,896]" journalOrPublisher="Scientific Reports" pageId="8" pageNumber="9" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
). Despite its narrowness, the overall dentary morphology is consistent with that of derived tyrannosaurines.
|
|
</paragraph>
|
|
<paragraph blockId="8.[113,784,179,1984]" pageId="8" pageNumber="9">
|
|
The subcutaneous surface of the dentary is smooth posteriorly and becomes increasingly rugose anteroventrally. The anterior alveolar margin is excavated by shallow, dorsoventrallyoriented neurovascular sulci that originate from the alveolar row of neurovascular foramina. Whereas similar subcutaneous texturing is observed in nearly all tyrannosaurids, the neurovascular sulci of the anterior alveolar margin tend to be deeper in larger, derived tyrannosaurines, such as
|
|
<taxonomicName box="[583,734,1154,1174]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[583,734,1154,1174]" italics="true" pageId="8" pageNumber="9">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[113,260,1182,1202]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[113,260,1182,1202]" italics="true" pageId="8" pageNumber="9">Tyrannosaurus</emphasis>
|
|
</taxonomicName>
|
|
.
|
|
</paragraph>
|
|
<paragraph blockId="8.[113,784,179,1984]" pageId="8" pageNumber="9">
|
|
In
|
|
<collectionCode box="[169,214,1212,1231]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[220,303,1211,1230]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
, the medial surface of the dentary is incised by a deep Meckelian groove that shallows anteriorly into a sulcus, a characteristic previously interpreted as an indicator of immaturity (
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[121,283,1295,1314]" journalOrPublisher="Scientific Reports" pageId="8" pageNumber="9" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
). The ventral bar is swollen and cylindrical posteriorly and flattens anteriorly towards the large (
|
|
<quantity box="[617,689,1323,1342]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="8" pageNumber="9" unit="mm" value="10.0">10 mm</quantity>
|
|
in diameter), circular Meckelian foramen. The area ventral to the Meckelian foramen is flat and weakly rugose in
|
|
<collectionCode box="[489,534,1379,1398]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[541,624,1379,1398]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
(
|
|
<figureCitation box="[639,719,1379,1398]" captionStart="Fig" captionStartId="10.[113,138,1695,1709]" captionTargetBox="[203,1411,182,1665]" captionTargetId="figure@10.[200,1413,178,1666]" captionTargetPageId="10" captionText="Fig. 10. Right dentary of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view. B) Specimen in dorsal view. Numbers correspond to dentary alveolus position. Comparison of dentary symphyseal region between C) Thanatotheristes and D) Daspletosaurus torosus (TMP 2001.36.1). Note the absence of bony projections on the symphyseal surface and a rugose knob ventral to Meckelian foramen in Thanatotheristes contrasting with Daspletosaurus. Abbreviations: apr, anteroposterior ridges; bp, bony projections; dc, dentary chin; mf, meckelian foramen; mg, meckelian groove; rk, rugose knob. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672025" httpUri="https://zenodo.org/record/3672025/files/figure.png" pageId="8" pageNumber="9">Fig. 10C</figureCitation>
|
|
), as in albertosaurines,
|
|
<taxonomicName box="[283,379,1405,1425]" class="Reptilia" family="Tyrannosauridae" genus="Lythronax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[283,379,1405,1425]" italics="true" pageId="8" pageNumber="9">Lythronax</emphasis>
|
|
</taxonomicName>
|
|
, and
|
|
<emphasis box="[442,583,1405,1425]" italics="true" pageId="8" pageNumber="9">Teratophoneus</emphasis>
|
|
, but in contrast to other derived tyrannosaurines (
|
|
<taxonomicName box="[484,631,1433,1453]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[484,631,1433,1453]" italics="true" pageId="8" pageNumber="9">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[660,778,1433,1453]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[660,778,1433,1453]" italics="true" pageId="8" pageNumber="9">Tarbosaurus</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[113,256,1461,1481]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[113,256,1461,1481]" italics="true" pageId="8" pageNumber="9">Tyrannosaurus</emphasis>
|
|
</taxonomicName>
|
|
,
|
|
<taxonomicName box="[268,447,1461,1481]" class="Reptilia" family="Tyrannosauridae" genus="Zhuchengtyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[268,447,1461,1481]" italics="true" pageId="8" pageNumber="9">Zhuchengtyrannus</emphasis>
|
|
</taxonomicName>
|
|
) where it is raised into a swollen and rugose knob (
|
|
<figureCitation box="[299,382,1490,1510]" captionStart="Fig" captionStartId="10.[113,138,1695,1709]" captionTargetBox="[203,1411,182,1665]" captionTargetId="figure@10.[200,1413,178,1666]" captionTargetPageId="10" captionText="Fig. 10. Right dentary of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view. B) Specimen in dorsal view. Numbers correspond to dentary alveolus position. Comparison of dentary symphyseal region between C) Thanatotheristes and D) Daspletosaurus torosus (TMP 2001.36.1). Note the absence of bony projections on the symphyseal surface and a rugose knob ventral to Meckelian foramen in Thanatotheristes contrasting with Daspletosaurus. Abbreviations: apr, anteroposterior ridges; bp, bony projections; dc, dentary chin; mf, meckelian foramen; mg, meckelian groove; rk, rugose knob. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672025" httpUri="https://zenodo.org/record/3672025/files/figure.png" pageId="8" pageNumber="9">Fig. 10D</figureCitation>
|
|
;
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." box="[394,553,1490,1510]" journalOrPublisher="Scientific Reports" pageId="8" pageNumber="9" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
). The symphyseal surface of
|
|
<collectionCode box="[200,245,1519,1538]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[259,343,1518,1537]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
is rugose (
|
|
<figureCitation box="[476,562,1518,1537]" captionStart="Fig" captionStartId="10.[113,138,1695,1709]" captionTargetBox="[203,1411,182,1665]" captionTargetId="figure@10.[200,1413,178,1666]" captionTargetPageId="10" captionText="Fig. 10. Right dentary of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view. B) Specimen in dorsal view. Numbers correspond to dentary alveolus position. Comparison of dentary symphyseal region between C) Thanatotheristes and D) Daspletosaurus torosus (TMP 2001.36.1). Note the absence of bony projections on the symphyseal surface and a rugose knob ventral to Meckelian foramen in Thanatotheristes contrasting with Daspletosaurus. Abbreviations: apr, anteroposterior ridges; bp, bony projections; dc, dentary chin; mf, meckelian foramen; mg, meckelian groove; rk, rugose knob. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672025" httpUri="https://zenodo.org/record/3672025/files/figure.png" pageId="8" pageNumber="9">Fig. 10C</figureCitation>
|
|
), bearing prominent anteroposteriorly-oriented ridges as in
|
|
<taxonomicName authorityName="Maleev" authorityYear="1955" box="[520,638,1545,1565]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[520,638,1545,1565]" italics="true" pageId="8" pageNumber="9">Tarbosaurus</emphasis>
|
|
</taxonomicName>
|
|
and
|
|
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis italics="true" pageId="8" pageNumber="9">Tyrannosaurus</emphasis>
|
|
</taxonomicName>
|
|
, but lacks the large and deeply interlocking bony projections observed in subadult and adult individuals of
|
|
<taxonomicName box="[636,783,1601,1621]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[636,783,1601,1621]" italics="true" pageId="8" pageNumber="9">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
(
|
|
<figureCitation box="[121,209,1630,1649]" captionStart="Fig" captionStartId="10.[113,138,1695,1709]" captionTargetBox="[203,1411,182,1665]" captionTargetId="figure@10.[200,1413,178,1666]" captionTargetPageId="10" captionText="Fig. 10. Right dentary of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view. B) Specimen in dorsal view. Numbers correspond to dentary alveolus position. Comparison of dentary symphyseal region between C) Thanatotheristes and D) Daspletosaurus torosus (TMP 2001.36.1). Note the absence of bony projections on the symphyseal surface and a rugose knob ventral to Meckelian foramen in Thanatotheristes contrasting with Daspletosaurus. Abbreviations: apr, anteroposterior ridges; bp, bony projections; dc, dentary chin; mf, meckelian foramen; mg, meckelian groove; rk, rugose knob. Scale bar equals 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672025" httpUri="https://zenodo.org/record/3672025/files/figure.png" pageId="8" pageNumber="9">Fig. 10D</figureCitation>
|
|
). Although development of the rugose knob and the symphyseal rugosities may be related to ontogeny (
|
|
<bibRefCitation author="Carr, T. D. & Varricchio, D. J. & Sedlmayr, J. C. & Roberts, E. M. & Moore, J. R." journalOrPublisher="Scientific Reports" pageId="8" pageNumber="9" pagination="11" part="7" refId="ref10327" refString="Carr, T. D., Varricchio, D. J., Sedlmayr, J. C., Roberts, E. M., Moore, J. R., 2017. A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system. Scientific Reports 7, 1 e 11. https: // doi. org / 10.1038 / srep 44942." title="A new tyrannosaur with evidence for anagenesis and crocodile-like facial sensory system" type="journal article" year="2017">Carr et al., 2017</bibRefCitation>
|
|
), the fact that they are present in
|
|
<taxonomicName box="[517,664,1684,1704]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[517,664,1684,1704]" italics="true" pageId="8" pageNumber="9">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
individuals of similar size to
|
|
<collectionCode box="[291,336,1714,1733]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[345,428,1713,1732]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
suggests that their absence in the latter is not due to ontogeny in this specimen.
|
|
</paragraph>
|
|
<paragraph blockId="8.[113,784,179,1984]" pageId="8" pageNumber="9">
|
|
<emphasis bold="true" box="[145,262,1769,1788]" pageId="8" pageNumber="9">Surangular</emphasis>
|
|
-Only the posterior-most portion of the right surangular is present in
|
|
<collectionCode box="[339,384,1798,1817]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[394,477,1797,1816]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
, which includes the articular ramus, the posterior third of the surangular shelf, and the posterior margin of the surangular foramen (
|
|
<figureCitation box="[479,534,1853,1872]" captionStart="Fig" captionStartId="9.[805,830,875,889]" captionTargetBox="[815,1469,182,843]" captionTargetId="figure@9.[810,1474,178,846]" captionTargetPageId="9" captionText="Fig. 9. Articulated right quadrate and surangular of Thanatotheristes holotype in lateral view. Abbreviations: anc, angular contact surface; ls, lateral shelf; ngf, non-glenoid fossa; pp, pneumatic pocket; qu, quadrate; sf, surangular foramen. Scale bar equals 2 cm." figureDoi="http://doi.org/10.5281/zenodo.3672021" httpUri="https://zenodo.org/record/3672021/files/figure.png" pageId="8" pageNumber="9">Fig. 9</figureCitation>
|
|
). A largely undiagnostic portion of the right articular is preserved in articulation with the surangular.
|
|
</paragraph>
|
|
<paragraph blockId="8.[113,784,179,1984]" lastBlockId="8.[831,1501,179,896]" pageId="8" pageNumber="9">
|
|
The surangular shelf (sensu
|
|
<bibRefCitation author="Currie, P. J." box="[428,566,1937,1956]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="8" pageNumber="9" pagination="226" part="48" refId="ref10511" refString="Currie, P. J., 2003 b. Cranial anatomy of tyrannosaurid dinosaurs from the late Cretaceous Alberta, Canada. Acta Palaeontologica Polonica 48, 191 e 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the late Cretaceous Alberta, Canada" type="journal article" year="2003">Currie, 2003b</bibRefCitation>
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|
) is a deep and robust flange located anterior to the quadrate-surangular articulation. In
|
|
<collectionCode box="[831,876,180,199]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[881,964,179,198]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
, the surangular shelf projects dorsolaterally, such that its ventral surface is visible in lateral view (
|
|
<figureCitation box="[1273,1327,207,226]" captionStart="Fig" captionStartId="9.[805,830,875,889]" captionTargetBox="[815,1469,182,843]" captionTargetId="figure@9.[810,1474,178,846]" captionTargetPageId="9" captionText="Fig. 9. Articulated right quadrate and surangular of Thanatotheristes holotype in lateral view. Abbreviations: anc, angular contact surface; ls, lateral shelf; ngf, non-glenoid fossa; pp, pneumatic pocket; qu, quadrate; sf, surangular foramen. Scale bar equals 2 cm." figureDoi="http://doi.org/10.5281/zenodo.3672021" httpUri="https://zenodo.org/record/3672021/files/figure.png" pageId="8" pageNumber="9">Fig. 9</figureCitation>
|
|
). In contrast, the surangular shelf projects laterally in albertosaurines and alioramins, and ventrolaterally in derived tyrannosaurines, where it overhangs the dorsal margin of the surangular foramen (
|
|
<bibRefCitation author="Brusatte, S. L. & Benson, R. B. J. & Norell, M. A." journalOrPublisher="American Museum Novitates" pageId="8" pageNumber="9" pagination="53" part="3717" refId="ref9801" refString="Brusatte, S. L., Benson, R. B. J., Norell, M. A., 2011. The anatomy of Dryptosaurus aquilunguis (Dinosauria: Theropoda) and a review of its tyrannosauroid affinities. American Museum Novitates 3717, 1 e 53." title="The anatomy of Dryptosaurus aquilunguis (Dinosauria: Theropoda) and a review of its tyrannosauroid affinities" type="journal article" year="2011">Brusatte et al., 2011</bibRefCitation>
|
|
). The adductor fossa, located on the dorsal surface of the surangular shelf, is medially bounded by the coronoid process. In
|
|
<collectionCode box="[855,900,375,394]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[905,988,374,393]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
, the coronoid process extends to the posteromedial margin of the adductor fossa and converges on the triangular process anterodorsal to the glenoid fossa (
|
|
<figureCitation box="[1293,1364,430,449]" captionStart="Fig" captionStartId="7.[88,113,1683,1697]" captionTargetBox="[96,751,182,1649]" captionTargetId="figure@7.[93,754,178,1653]" captionTargetPageId="7" captionText="Fig. 7. Right postorbital fragment of Thanatotheristes holotype (TMP 2010.5.7). A) Specimen in lateral view, demonstrating arced subcutaneous ridges, B) and medial view.Note prominent bracing ridge for jugal contact on the medial surface of the bone. Abbreviations: br, bracing ridge for jugal contact; djc, dorsal extent of jugal contact; scr, subcutaneous ridges; sr, smooth region along lateral surface posterior margin. Scale bar equals 2 cm." figureDoi="http://doi.org/10.5281/zenodo.3672019" httpUri="https://zenodo.org/record/3672019/files/figure.png" pageId="8" pageNumber="9">Fig. S7</figureCitation>
|
|
), as seen in
|
|
<taxonomicName box="[831,927,457,477]" class="Reptilia" family="Tyrannosauridae" genus="Lythronax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[831,927,457,477]" italics="true" pageId="8" pageNumber="9">Lythronax</emphasis>
|
|
</taxonomicName>
|
|
(
|
|
<collectionCode box="[946,1017,458,477]" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34862" lsid="urn:lsid:biocol.org:col:34862" name="Utah Museum of Natural History" pageId="8" pageNumber="9" type="Museum">UMNH</collectionCode>
|
|
<collectionCode box="[1030,1057,459,478]" pageId="8" pageNumber="9">VP</collectionCode>
|
|
20200). In all other tyrannosaurids, the coronoid process terminates anterior to the posterior margin of adductor fossa.
|
|
</paragraph>
|
|
<paragraph blockId="8.[831,1501,179,896]" pageId="8" pageNumber="9">
|
|
A pneumatic pocket occurs posterior to the surangular foramen in all mature tyrannosaurines and most albertosaurine specimens. In
|
|
<collectionCode box="[856,901,598,617]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[906,989,598,617]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
, this pneumatic pocket is enclosed posterolaterally by a thick flange (
|
|
<figureCitation box="[1025,1083,626,645]" captionStart="Fig" captionStartId="9.[805,830,875,889]" captionTargetBox="[815,1469,182,843]" captionTargetId="figure@9.[810,1474,178,846]" captionTargetPageId="9" captionText="Fig. 9. Articulated right quadrate and surangular of Thanatotheristes holotype in lateral view. Abbreviations: anc, angular contact surface; ls, lateral shelf; ngf, non-glenoid fossa; pp, pneumatic pocket; qu, quadrate; sf, surangular foramen. Scale bar equals 2 cm." figureDoi="http://doi.org/10.5281/zenodo.3672021" httpUri="https://zenodo.org/record/3672021/files/figure.png" pageId="8" pageNumber="9">Fig. 9</figureCitation>
|
|
and
|
|
<figureCitation box="[1140,1209,626,645]" captionStart="Fig" captionStartId="9.[88,113,1813,1827]" captionTargetBox="[94,753,182,1782]" captionTargetId="figure@9.[93,754,178,1783]" captionTargetPageId="9" captionText="Fig. 8. Comparison of left frontalin dorsalviewin tyrannosaurids.A)Thanatotheristes TMP 2010.5.7. White dash line represents nondiagnostic postorbital fragment attached to the frontal. B) Subcutaneous surface of frontal of Daspletosaurus torosus TMP 2001.036.0001. Abbreviations: apc, anterior postorbital contact expansion; aesc, autapomorphic anterior extension of sagittal crest; ifs, interfrontal suture;ls, lacrimal socket; np, nasal process; os, orbital slot; PA, parietal; PF, prefrontal; pfl, prefrontolacrimal process; pfn, prefrontonasal process; po, postorbital fragment; sc, sagittal crest; stf, supratemporal fossa; str, supratemporal ridge. Scale bars equal 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3672023" httpUri="https://zenodo.org/record/3672023/files/figure.png" pageId="8" pageNumber="9">Fig. S8</figureCitation>
|
|
). In other derived tyrannosaurines (including
|
|
<taxonomicName box="[1050,1197,652,672]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="genus">
|
|
<emphasis box="[1050,1197,652,672]" italics="true" pageId="8" pageNumber="9">Daspletosaurus</emphasis>
|
|
</taxonomicName>
|
|
), the pneumatic pocket is enclosed posterolaterally and ventrolaterally by this same flange, and dorsolaterally by the posterior edge of the lateral surangular shelf.
|
|
</paragraph>
|
|
<paragraph blockId="8.[831,1501,179,896]" pageId="8" pageNumber="9">
|
|
The nonglenoid fossa is a crescentic depression located ventrolateral to the glenoid fossa (
|
|
<figureCitation box="[1185,1241,793,812]" captionStart="Fig" captionStartId="9.[805,830,875,889]" captionTargetBox="[815,1469,182,843]" captionTargetId="figure@9.[810,1474,178,846]" captionTargetPageId="9" captionText="Fig. 9. Articulated right quadrate and surangular of Thanatotheristes holotype in lateral view. Abbreviations: anc, angular contact surface; ls, lateral shelf; ngf, non-glenoid fossa; pp, pneumatic pocket; qu, quadrate; sf, surangular foramen. Scale bar equals 2 cm." figureDoi="http://doi.org/10.5281/zenodo.3672021" httpUri="https://zenodo.org/record/3672021/files/figure.png" pageId="8" pageNumber="9">Fig. 9</figureCitation>
|
|
;
|
|
<bibRefCitation author="Brusatte, S. L. & Carr, T. D. & Norell, M. A." box="[1256,1459,793,812]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="8" pageNumber="9" pagination="197" part="366" refId="ref9853" refString="Brusatte, S. L., Carr, T. D., Norell, M. A., 2012. The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bulletin of the American Museum of Natural History 366, 1 e 197." title="The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia" type="journal article" year="2012">Brusatte et al., 2012</bibRefCitation>
|
|
). In
|
|
<collectionCode box="[831,876,821,840]" country="South Africa" httpUri="http://grbio.org/cool/4q52-xvg6" name="Transvaal Museum" pageId="8" pageNumber="9">TMP</collectionCode>
|
|
<date box="[883,966,821,840]" pageId="8" pageNumber="9" value="2010-05-07">2010.5.7</date>
|
|
, the anteroventral margin of the nonglenoid fossa is bounded by a prominent and rugose crest. This crest is significantly lower in all other tyrannosaurids.
|
|
</paragraph>
|
|
</subSubSection>
|
|
</treatment>
|
|
</document> |