1150 lines
216 KiB
XML
1150 lines
216 KiB
XML
<document id="91725BB6CB722DAA40DB806E67AD3BC8" ID-DOI="10.2110/palo.2017.076" ID-GBIF-Dataset="bba5c695-619c-4f8e-a634-c29f65a72f1c" ID-Zenodo-Dep="3749641" IM.bibliography_requiresApprovalFor="plazi" IM.metadata_requiresApprovalFor="plazi" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_requiresApprovalFor="plazi" checkinTime="1586775310973" checkinUser="jeremy" docAuthor="Matthew A. McLain, David Nelsen, Keith Snyder, Christopher T. Griffin, Bethania Siviero, Leonard R. Brand & Arthur V. Chadwick" docDate="2018" docId="038466034D5FDE59FBD84DEDFA16F78C" docLanguage="en" docName="McLainetal2018.pdf.imf" docOrigin="Palaios 33" docStyle="DocumentStyle{}" docTitle="Tyrannosaurus rex Osborn 1905" docType="treatment" docVersion="12" lastPageNumber="171" masterDocId="FFBD1E7B4D5EDE51FFC14B75FF8CF360" masterDocTitle="Tyrannosaur cannibalism: A case of a tooth-traced tyrannosaurid bone in the Lance Formation (Maastrichtian), Wyoming" masterLastPageNumber="173" masterPageNumber="164" pageNumber="164" updateTime="1698737307372" updateUser="plazi">
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<mods:title id="86AF7346689873BBF6A038F6FD84FBA1">Tyrannosaur cannibalism: A case of a tooth-traced tyrannosaurid bone in the Lance Formation (Maastrichtian), Wyoming</mods:title>
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<mods:name id="D1C7541CEC6D63E7A72E2D509EF35A04" type="personal">
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<mods:roleTerm id="9294056D4CEE5D4B367C79A1F68B9ED2">Author</mods:roleTerm>
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<mods:namePart id="78FDE2CE1023C6805BAE613FA0E2E8BD">Matthew A. McLain</mods:namePart>
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<mods:affiliation id="2A9122A2CA109745F26C4E0CA564BECB">Department of Biological and Physical Sciences, The Master’s University, Santa Clarita, California</mods:affiliation>
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<mods:nameIdentifier id="389A4DC88F76E47270BF9E74204E851B" type="email">mmclain@masters.edu</mods:nameIdentifier>
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<mods:namePart id="E952BC42D8FA5EEAC18146B20ECED375">David Nelsen</mods:namePart>
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<mods:affiliation id="0EC1A2C407AEDC2CA43F6C5F8527C371">Department of Biology, Southern Adventist University, Chattanooga, Tennessee, USA</mods:affiliation>
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</mods:name>
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<mods:name id="A6554D6D2F78CCE6A2CB85982DB0A2CD" type="personal">
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<mods:roleTerm id="89EA3F332BDA2A18CDACA4C186005E6D">Author</mods:roleTerm>
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<mods:namePart id="4D592D7F27381019FF8E418BBC5A0FF8">Keith Snyder</mods:namePart>
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<mods:affiliation id="452B66C6DF9589896660C3090C1A6259">Department of Biology, Southern Adventist University, Chattanooga, Tennessee, USA</mods:affiliation>
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</mods:name>
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<mods:roleTerm id="7D76B5B5C9CF77C9822006D447B5F30C">Author</mods:roleTerm>
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<mods:namePart id="5DF3DCB9438C58A03227BEB504D99B2F">Christopher T. Griffin</mods:namePart>
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<mods:affiliation id="5CD34931F2959C2BEDFAAEA4C9498A9B">Department of Geosciences, Virginia Tech, Blacksburg, Virginia, USA</mods:affiliation>
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</mods:name>
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<mods:name id="87E6B75BF0F78335DA669FA140197F26" type="personal">
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<mods:role id="5DB358843E75A241DCA84509A90232AD">
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<mods:roleTerm id="AA4B3E578CDE46F36B5083875BB6DC25">Author</mods:roleTerm>
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</mods:role>
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<mods:namePart id="3C8EEF87453D292FBAE3F71B609BC5E1">Bethania Siviero</mods:namePart>
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<mods:affiliation id="424D44E9934680E1514867850A056F0C">Department of Earth and Biological Sciences, Loma Linda University, Loma Linda, California, USA</mods:affiliation>
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</mods:name>
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<mods:name id="BCF2B071B9CC05A1A248C09502439EB7" type="personal">
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<mods:roleTerm id="7ABD8D6C4E0F1DBCC695118B4CA69710">Author</mods:roleTerm>
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<mods:namePart id="E4F2196277891020BFCE1567D665E37C">Leonard R. Brand</mods:namePart>
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<mods:affiliation id="B3444D09D2116492E6178941174C1B6C">Department of Earth and Biological Sciences, Loma Linda University, Loma Linda, California, USA</mods:affiliation>
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<mods:roleTerm id="E3C5BE963BFB4D22282EBD8646096323">Author</mods:roleTerm>
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<mods:namePart id="97D2A3838010D4BB65AF12F5245F5D0F">Arthur V. Chadwick</mods:namePart>
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<mods:affiliation id="7F8D4482B394ADC3BE9528A26BA6D192">Department of Biological Sciences, Southwestern Adventist University, Keene, Texas, USA</mods:affiliation>
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<mods:title id="BA6080DD70B39F87EED501636A8E49AC">Palaios</mods:title>
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<mods:date id="6936F3826E0505B5FC4D2A034081993D">2018</mods:date>
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<mods:detail id="B24DC211162E6BE03D5C01454C5A4C7B" type="volume">
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<mods:number id="C3D721459D6E36C763119E96A94606F0">33</mods:number>
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<mods:start id="66322A18F5ED1BCC565271E7FE26786D">164</mods:start>
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<mods:classification id="9EB6B6CCF48C6FE789DB5B7B2041FFE4">journal article</mods:classification>
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<mods:identifier id="F9FFE8614D583BDF85F957254A57B298" type="DOI">10.2110/palo.2017.076</mods:identifier>
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<treatment id="038466034D5FDE59FBD84DEDFA16F78C" ID-DOI="http://doi.org/10.5281/zenodo.4913673" ID-GBIF-Taxon="190305150" ID-Zenodo-Dep="4913673" LSID="urn:lsid:plazi:treatment:038466034D5FDE59FBD84DEDFA16F78C" httpUri="http://treatment.plazi.org/id/038466034D5FDE59FBD84DEDFA16F78C" lastPageId="8" lastPageNumber="171" pageId="1" pageNumber="164">
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<subSubSection id="C337849E4D5FDE50FBD84DEDFA8AF45D" pageId="1" pageNumber="164" type="description">
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<paragraph id="8B92D7154D5FDE50FBD84DEDFB60F5CB" blockId="1.[993,1316,1660,1882]" box="[1049,1260,1688,1708]" pageId="1" pageNumber="164">
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<taxonomicName id="4C2DAC964D5FDE50FBD84DEDFB60F5CB" authority="Owen 1842" authorityName="Owen" authorityYear="1842" box="[1049,1260,1688,1708]" class="Dinosauria" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="164" phylum="Chordata" rank="class">
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||
Dinosauria
|
||
<bibRefCitation id="EFBCAAE44D5FDE50FB424DEDFB60F5CB" author="OWEN, R." box="[1155,1260,1688,1707]" journalOrPublisher="Report of the British Association for the Advancement of Science" pageId="1" pageNumber="164" pagination="60 - 204" part="11" refId="ref10028" refString="OWEN, R., 1842, Report on British fossil reptiles, part II: Report of the British Association for the Advancement of Science, v. 11, p. 60 - 204." title="Report on British fossil reptiles, part II" type="journal article" year="1842">Owen 1842</bibRefCitation>
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</taxonomicName>
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</paragraph>
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<paragraph id="8B92D7154D5FDE50FBD94DC0FB61F5A8" blockId="1.[993,1316,1660,1882]" box="[1048,1261,1717,1737]" pageId="1" pageNumber="164">
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||
<taxonomicName id="4C2DAC964D5FDE50FBD94DC0FB61F5A8" authority="Seeley 1887" authorityName="Seeley" authorityYear="1887" box="[1048,1261,1717,1737]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="1" pageNumber="164" phylum="Chordata" rank="order">
|
||
Saurischia
|
||
<bibRefCitation id="EFBCAAE44D5FDE50FBBC4DC0FB61F5A8" author="SEELEY, H. G." box="[1149,1261,1717,1737]" journalOrPublisher="Proceedings of the Royal Society of London" pageId="1" pageNumber="164" pagination="165 - 171" part="43" refId="ref10623" refString="SEELEY, H. G., 1887, On the classification of the fossil animals commonly named Dinosauria: Proceedings of the Royal Society of London, v. 43, p. 165 - 171." title="On the classification of the fossil animals commonly named Dinosauria" type="journal article" year="1887">Seeley 1887</bibRefCitation>
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</taxonomicName>
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</paragraph>
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||
<paragraph id="8B92D7154D5FDE50FBD64DA7FB62F586" blockId="1.[993,1316,1660,1882]" box="[1047,1262,1746,1766]" pageId="1" pageNumber="164">
|
||
Theropoda
|
||
<bibRefCitation id="EFBCAAE44D5FDE50FB414DA7FB62F586" author="MARSH, O. C." box="[1152,1262,1746,1766]" journalOrPublisher="American Journal of Science" pageId="1" pageNumber="164" pagination="417 - 423" part="3" refId="ref9657" refString="MARSH, O. C., 1881, Principal characters of American Jurassic dinosaurs, part V: American Journal of Science, v. 3, p. 417 - 423." title="Principal characters of American Jurassic dinosaurs, part V" type="journal article" year="1881">Marsh 1881</bibRefCitation>
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||
</paragraph>
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||
<paragraph id="8B92D7154D5FDE50FC344D9AFA9CF463" blockId="1.[993,1316,1660,1882]" box="[1013,1296,1775,1795]" pageId="1" pageNumber="164">
|
||
Tyrannosauroidea
|
||
<bibRefCitation id="EFBCAAE44D5FDE50FB5D4D9AFA9CF463" author="WALKER, A. D." box="[1180,1296,1775,1795]" journalOrPublisher="Philosophical Transactions of the Royal Society of London B" pageId="1" pageNumber="164" pagination="53 - 134" part="248" refId="ref10880" refString="WALKER, A. D., 1964, Triassic reptiles from the Elgin area, Ornithosuchus and the origin of carnosaurs: Philosophical Transactions of the Royal Society of London B, v. 248, p. 53 - 134." title="Triassic reptiles from the Elgin area, Ornithosuchus and the origin of carnosaurs" type="journal article" year="1964">Walker 1964</bibRefCitation>
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||
</paragraph>
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||
<paragraph id="8B92D7154D5FDE50FC384C79FA87F440" blockId="1.[993,1316,1660,1882]" box="[1017,1291,1804,1824]" pageId="1" pageNumber="164">
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||
<taxonomicName id="4C2DAC964D5FDE50FC384C79FA87F440" authority="Osborn 1906" authorityName="Osborn" authorityYear="1906" box="[1017,1291,1804,1824]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="164" phylum="Chordata" rank="family">
|
||
Tyrannosauridae
|
||
<bibRefCitation id="EFBCAAE44D5FDE50FB544C79FA87F440" author="OSBORN, H. F." box="[1173,1291,1804,1824]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="1" pageNumber="164" pagination="281 - 296" part="22" refId="ref9989" refString="OSBORN, H. F., 1906, Tyrannosaurus, Upper Cretaceous carnivorous dinosaur (second communication): Bulletin of the American Museum of Natural History, v. 22, p. 281 - 296." title="Tyrannosaurus, Upper Cretaceous carnivorous dinosaur (second communication)" type="journal article" year="1906">Osborn 1906</bibRefCitation>
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||
</taxonomicName>
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||
</paragraph>
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||
<paragraph id="8B92D7154D5FDE50FC3E4C5EFA8AF45D" blockId="1.[993,1316,1660,1882]" box="[1023,1286,1833,1853]" pageId="1" pageNumber="164">
|
||
<taxonomicName id="4C2DAC964D5FDE50FC3E4C5EFA8AF45D" authority="Osborn 1905" authorityName="Osborn" authorityYear="1905" box="[1023,1286,1833,1853]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="164" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D5FDE50FC3E4C5EFB0BF45D" box="[1023,1159,1835,1853]" italics="true" pageId="1" pageNumber="164">Tyrannosaurus</emphasis>
|
||
<bibRefCitation id="EFBCAAE44D5FDE50FB4E4C5CFA8AF45D" author="OSBORN, H. F." box="[1167,1286,1833,1853]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="1" pageNumber="164" pagination="259 - 265" part="21" refId="ref9954" refString="OSBORN, H. F., 1905, Tyrannosaurus and other Cretaceous carnivorous dinosaurs: Bulletin of the American Museum of Natural History, v. 21, p. 259 - 265." title="Tyrannosaurus and other Cretaceous carnivorous dinosaurs" type="journal article" year="1905">Osborn 1905</bibRefCitation>
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</taxonomicName>
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</paragraph>
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</subSubSection>
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<subSubSection id="C337849E4D5FDE50FC2C4C3DFA94F43A" box="[1005,1304,1862,1882]" pageId="1" pageNumber="164" type="nomenclature">
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||
<paragraph id="8B92D7154D5FDE50FC2C4C3DFA94F43A" blockId="1.[993,1316,1660,1882]" box="[1005,1304,1862,1882]" pageId="1" pageNumber="164">
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||
<taxonomicName id="4C2DAC964D5FDE50FC2C4C3DFA94F43A" authority="Osborn 1905" authorityName="Osborn" authorityYear="1905" box="[1005,1304,1862,1882]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="164" phylum="Chordata" rank="species" species="rex">
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||
<emphasis id="B9590B074D5FDE50FC2C4C3DFB16F43A" box="[1005,1178,1864,1882]" italics="true" pageId="1" pageNumber="164">Tyrannosaurus rex</emphasis>
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<bibRefCitation id="EFBCAAE44D5FDE50FB604C33FA94F43A" author="OSBORN, H. F." box="[1185,1304,1862,1882]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="1" pageNumber="164" pagination="259 - 265" part="21" refId="ref9954" refString="OSBORN, H. F., 1905, Tyrannosaurus and other Cretaceous carnivorous dinosaurs: Bulletin of the American Museum of Natural History, v. 21, p. 259 - 265." title="Tyrannosaurus and other Cretaceous carnivorous dinosaurs" type="journal article" year="1905">Osborn 1905</bibRefCitation>
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</taxonomicName>
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</paragraph>
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</subSubSection>
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<subSubSection id="C337849E4D5FDE53FC8B4C0FFD19F729" lastPageId="2" lastPageNumber="165" pageId="1" pageNumber="164" type="materials_examined">
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<paragraph id="8B92D7154D5FDE53FC8B4C0FFD19F729" blockId="1.[842,1395,1914,1934]" lastBlockId="2.[165,661,1077,1097]" lastPageId="2" lastPageNumber="165" pageId="1" pageNumber="164">
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<emphasis id="B9590B074D5FDE50FC8B4C0FFC37F4EE" bold="true" box="[842,955,1914,1934]" pageId="1" pageNumber="164">Material.—</emphasis>
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<materialsCitation id="3B45DD484D5FDE53FC7A4C0FFD19F729" ID-GBIF-Occurrence="3396390314" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara County" lastPageId="2" lastPageNumber="165" pageId="1" pageNumber="164" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">
|
||
SWAU HRS13997, a partial right metatarsal IV.
|
||
<emphasis id="B9590B074D5CDE53FF644F40FE9DF729" bold="true" box="[165,273,1077,1097]" pageId="2" pageNumber="165">Locality.—</emphasis>
|
||
Niobrara County, eastern Wyoming, USA.
|
||
</materialsCitation>
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||
</paragraph>
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||
</subSubSection>
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||
<caption id="DF52879D4D5CDE53FE604882FB52F767" ID-DOI="http://doi.org/10.5281/zenodo.3749643" ID-Zenodo-Dep="3749643" box="[417,1246,1014,1032]" httpUri="https://zenodo.org/record/3749643/files/figure.png" pageId="2" pageNumber="165" startId="2.[417,443,1015,1031]" targetBox="[144,1515,176,986]" targetPageId="2">
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<paragraph id="8B92D7154D5CDE53FE604882FB52F767" blockId="2.[417,1246,1014,1032]" box="[417,1246,1014,1032]" pageId="2" pageNumber="165">
|
||
FIG. 1.—A
|
||
<taxonomicName id="4C2DAC964D5CDE53FE3F4882FD1AF767" authorityName="Osborn" authorityYear="1905" box="[510,662,1015,1031]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D5CDE53FE3F4882FD1AF767" box="[510,662,1015,1031]" italics="true" pageId="2" pageNumber="165">Tyrannosaurus rex</emphasis>
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</taxonomicName>
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||
metatarsal IV (
|
||
<materialsCitation id="3B45DD484D5CDE53FCD44882FC22F768" ID-GBIF-Occurrence="3396390301" box="[789,942,1014,1032]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="2" pageNumber="165" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
) in four views. Scale bar = 100 mm.
|
||
</paragraph>
|
||
</caption>
|
||
<subSubSection id="C337849E4D5CDE53FF644F07FE5CF680" pageId="2" pageNumber="165" type="diagnosis">
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||
<paragraph id="8B92D7154D5CDE53FF644F07FE5CF680" blockId="2.[141,815,1138,1504]" pageId="2" pageNumber="165">
|
||
<emphasis id="B9590B074D5CDE53FF644F07FE92F7E6" bold="true" box="[165,286,1138,1158]" pageId="2" pageNumber="165">Diagnosis.—</emphasis>
|
||
The hollow core of
|
||
<materialsCitation id="3B45DD484D5CDE53FE194F06FD0EF7E6" ID-GBIF-Occurrence="3396390317" box="[472,642,1139,1158]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="2" pageNumber="165" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
is evidence that it came from a theropod, and the large size of the bone and thickness of the bone walls coupled with its presence in the Lance Formation suggests that it came from a tyrannosaurid. We have identified
|
||
<materialsCitation id="3B45DD484D5CDE53FD844FBCFD61F7BC" ID-GBIF-Occurrence="3396390332" box="[581,749,1225,1244]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="2" pageNumber="165" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
to be a partial right metatarsal IV as evidenced by the flattened region on the medial surface that is the contacting surface with the distinctive metatarsal III of tyrannosaurids, which narrows to a point medially (
|
||
<bibRefCitation id="EFBCAAE44D5CDE53FD774E6AFCA8F653" author="HOLTZ, T. R." box="[694,804,1311,1331]" editor="D. B. Weishampel & P. Dodson & H. Osmolska" journalOrPublisher="University of California Press, Berkeley, CA" pageId="2" pageNumber="165" pagination="111 - 136" refId="ref8878" refString="HOLTZ, T. R., 2004, Tyrannosauroidea, in D. B. Weishampel, P. Dodson, and H. Osmolska (eds.), The Dinosauria: University of California Press, Berkeley, CA, p. 111 - 136." title="Tyrannosauroidea" type="book chapter" volumeTitle="The Dinosauria" year="2004">Holtz 2004</bibRefCitation>
|
||
). Since the only tyrannosaurids currently recognized in the Lance Formation are
|
||
<taxonomicName id="4C2DAC964D5CDE53FF6F4E2FFED6F60C" authorityName="Osborn" authorityYear="1905" box="[174,346,1370,1388]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D5CDE53FF6F4E2FFED6F60C" box="[174,346,1370,1388]" italics="true" pageId="2" pageNumber="165">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
and possibly
|
||
<taxonomicName id="4C2DAC964D5CDE53FE164E2FFD23F60D" baseAuthorityName="Molnar" baseAuthorityYear="1980" box="[471,687,1369,1389]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="species" species="lancensis">
|
||
<emphasis id="B9590B074D5CDE53FE164E2FFD23F60D" box="[471,687,1369,1389]" italics="true" pageId="2" pageNumber="165">Nanotyrannus lancensis</emphasis>
|
||
</taxonomicName>
|
||
, it very likely belongs to one of these two. Because the individual was at the purported size of a mature
|
||
<taxonomicName id="4C2DAC964D5CDE53FEEF4EE1FD86F6C7" baseAuthorityName="Molnar" baseAuthorityYear="1980" box="[302,522,1427,1447]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="species" species="lancensis">
|
||
<emphasis id="B9590B074D5CDE53FEEF4EE1FD86F6C7" box="[302,522,1427,1447]" italics="true" pageId="2" pageNumber="165">Nanotyrannus lancensis</emphasis>
|
||
</taxonomicName>
|
||
, yet immature and still rapidly growing, we conclude that the bone is from a
|
||
<taxonomicName id="4C2DAC964D5CDE53FD914EC5FC8CF6A2" authorityName="Osborn" authorityYear="1905" box="[592,768,1456,1474]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D5CDE53FD914EC5FC8CF6A2" box="[592,768,1456,1474]" italics="true" pageId="2" pageNumber="165">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
(see histological description in Results).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C337849E4D5CDE55FF644D7CFAD9F6DF" lastPageId="4" lastPageNumber="167" pageId="2" pageNumber="165" type="description">
|
||
<paragraph id="8B92D7154D5CDE53FF644D7CFD8EF5F1" blockId="2.[141,814,1545,1681]" pageId="2" pageNumber="165">
|
||
<emphasis id="B9590B074D5CDE53FF644D7CFEBDF57D" bold="true" box="[165,305,1545,1565]" pageId="2" pageNumber="165">Description.—</emphasis>
|
||
Both the proximal and distal ends of
|
||
<materialsCitation id="3B45DD484D5CDE53FD454D7FFCA1F57D" ID-GBIF-Occurrence="3396390304" box="[644,813,1546,1565]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="2" pageNumber="165" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
are broken, resulting in a length of 280 mm for what remains of the element. In addition to the fractures at the distal and proximal ends, a section of the shaft is missing, which exposes the hollow core of the bone (
|
||
<figureCitation id="1316CB904D5CDE53FF554D08FF46F5F0" box="[148,202,1661,1681]" captionStart="FIG. 1" captionStartId="2.[417,443,1015,1031]" captionTargetBox="[144,1515,176,986]" captionTargetId="figure@2.[144,1515,176,986]" captionTargetPageId="2" captionText="FIG. 1.—A Tyrannosaurus rex metatarsal IV (SWAU HRS13997) in four views. Scale bar = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.3749643" httpUri="https://zenodo.org/record/3749643/files/figure.png" pageId="2" pageNumber="165">Fig. 1</figureCitation>
|
||
, online
|
||
<httpUri id="B5AC09BF4D5CDE53FED64D08FE79F5F1" box="[279,501,1661,1681]" pageId="2" pageNumber="165">Supplemental File fig. 1</httpUri>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D5CDE53FE6E4DC8FD87F5AD" blockId="2.[431,523,1725,1741]" box="[431,523,1725,1741]" pageId="2" pageNumber="165">
|
||
<heading id="D0DA60794D5CDE53FE6E4DC8FD87F5AD" allCaps="true" bold="true" box="[431,523,1725,1741]" centered="true" fontSize="7" level="3" pageId="2" pageNumber="165" reason="2">
|
||
<emphasis id="B9590B074D5CDE53FE6E4DC8FD87F5AD" bold="true" box="[431,523,1725,1741]" pageId="2" pageNumber="165">METHODS</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D5CDE53FF644D9FFC05F6C3" blockId="2.[141,815,1770,1934]" lastBlockId="2.[845,1518,1077,1934]" pageId="2" pageNumber="165">
|
||
After excavation,
|
||
<materialsCitation id="3B45DD484D5CDE53FE8C4D9FFE75F59D" ID-GBIF-Occurrence="3396390302" box="[333,505,1770,1789]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="2" pageNumber="165" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
was prepared and photographed with a Canon Mark II 5D camera and an electronically coupled turntable. This allows for a 3DVR image of the bone to be created, which is available for viewing at http://fossil.swau.edu. The scratches on the bone were carefully observed and photographed using a Nikon SMZ 1500 with a Nikon DS-Fi1 Digital Sight camera, and they were measured using the Nikon NIS-Element Basic Research v. 4.30.02. The widths of the striations on the
|
||
<taxonomicName id="4C2DAC964D5CDE53FC514F27FBE0F706" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[912,1132,1106,1126]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="165" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5CDE53FC514F27FBE0F706" box="[912,1132,1106,1126]" italics="true" pageId="2" pageNumber="165">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
were measured and compared to denticle widths on theropod teeth obtained from nearby Lance Formation quarries (see online Supplementary File for the list of specimens measured). We compared
|
||
<materialsCitation id="3B45DD484D5CDE53FC6A4FDCFBDFF7DC" ID-GBIF-Occurrence="3396390329" box="[939,1107,1193,1212]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="2" pageNumber="165" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
to tyrannosaurid material from the LACM for identification purposes, specifically
|
||
<materialsCitation id="3B45DD484D5CDE53FB6D4FB3FABAF7B9" ID-GBIF-Occurrence="3396390336" box="[1196,1334,1221,1241]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/ContactSheet.php?QueryName=DetailedQuery&StartAt=1&QueryPage=%2Fdinosaur-institute%2Findex.php&Restriction=%28exists+%28SecDepartment_tab+where+%28SecDepartment+contains+%27Dinosaur+Institute%27%29%29%29&col_SpeSpecimenNoInteger=23845&LimitPerPage=20&Search=Search& " pageId="2" pageNumber="165" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
, a specimen from Montana originally considered to be from
|
||
<taxonomicName id="4C2DAC964D5CDE53FB0A4F97FAC7F796" authorityName="Osborn" authorityYear="1905" box="[1227,1355,1250,1270]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D5CDE53FB0A4F97FAC7F796" box="[1227,1355,1250,1270]" italics="true" pageId="2" pageNumber="165">Albertosaurus</emphasis>
|
||
</taxonomicName>
|
||
cf.
|
||
<taxonomicName id="4C2DAC964D5CDE53FAAF4F97FA62F796" authorityName="Osborn" authorityYear="1905" box="[1390,1518,1250,1270]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D5CDE53FAAF4F97FA62F796" box="[1390,1518,1250,1270]" italics="true" pageId="2" pageNumber="165">Albertosaurus</emphasis>
|
||
</taxonomicName>
|
||
(=
|
||
<taxonomicName id="4C2DAC964D5CDE53FCA34E75FC6EF672" box="[866,994,1280,1298]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D5CDE53FCA34E75FC6EF672" box="[866,994,1280,1298]" italics="true" pageId="2" pageNumber="165">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
)
|
||
<taxonomicName id="4C2DAC964D5CDE53FC364F8AFB6BF673" authority="(Molnar 1980)" baseAuthorityName="Molnar" baseAuthorityYear="1980" box="[1015,1255,1279,1299]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="species" species="lancensis">
|
||
<emphasis id="B9590B074D5CDE53FC364F8AFBC7F673" box="[1015,1099,1279,1299]" italics="true" pageId="2" pageNumber="165">lancensis</emphasis>
|
||
(
|
||
<bibRefCitation id="EFBCAAE44D5CDE53FB9E4F8AFB6CF673" author="MOLNAR, R. E." box="[1119,1248,1279,1299]" journalOrPublisher="Journal of Paleontology" pageId="2" pageNumber="165" pagination="102 - 108" part="54" refId="ref9732" refString="MOLNAR, R. E., 1980, An albertosaur from the Hell Creek Formation of Montana: Journal of Paleontology, v. 54, p. 102 - 108." title="An albertosaur from the Hell Creek Formation of Montana" type="journal article" year="1980">Molnar 1980</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, then described as a new species (
|
||
<taxonomicName id="4C2DAC964D5CDE53FC5A4E69FA9AF650" authority=", (Paul 1988))" authorityName=", (Paul" authorityYear="1988" baseAuthorityName="Paul" baseAuthorityYear="1988" box="[923,1302,1308,1328]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="species" species="megagracilis">
|
||
<emphasis id="B9590B074D5CDE53FC5A4E69FB14F650" box="[923,1176,1308,1328]" italics="true" pageId="2" pageNumber="165">Albertosaurus megagracilis</emphasis>
|
||
, (
|
||
<bibRefCitation id="EFBCAAE44D5CDE53FB684E69FA8BF650" author="PAUL, G. S." bookContentInfo="464 p." box="[1193,1287,1308,1328]" journalOrPublisher="Simon and Schuster, New York, New York" pageId="2" pageNumber="165" refId="ref10111" refString="PAUL, G. S., 1988, Predatory Dinosaurs of the World: A Complete Illustrated Guide: Simon and Schuster, New York, New York, 464 p." title="Predatory Dinosaurs of the World: A Complete Illustrated Guide" type="book" year="1988">Paul 1988</bibRefCitation>
|
||
))
|
||
</taxonomicName>
|
||
, which was later placed in a new genus
|
||
<taxonomicName id="4C2DAC964D5CDE53FC1F4E4FFA88F62C" authority="(Olshevsky 1995)" baseAuthorityName="Olshevsky" baseAuthorityYear="1995" box="[990,1284,1336,1356]" class="Reptilia" family="Tyrannosauridae" genus="Dinotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D5CDE53FC1F4E4FFBD6F62C" box="[990,1114,1338,1356]" italics="true" pageId="2" pageNumber="165">Dinotyrannus</emphasis>
|
||
(
|
||
<bibRefCitation id="EFBCAAE44D5CDE53FBA64E4DFB71F62C" author="OLSHEVSKY, G." box="[1127,1277,1336,1356]" journalOrPublisher="Kyoryugaku Saizensen" pageId="2" pageNumber="165" pagination="92 - 119" part="9 - 10" refId="ref9925" refString="OLSHEVSKY, G., 1995, The origin and evolution of tyrannosaurids: Kyoryugaku Saizensen, v. 9 - 10, p. 92 - 119." title="The origin and evolution of tyrannosaurids" type="journal article" year="1995">Olshevsky 1995</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, but is now considered to be a ‘‘subadult’’
|
||
<taxonomicName id="4C2DAC964D5CDE53FC254E22FA17F609" authority="(Carr and Williamson 2004)" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[996,1435,1365,1385]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="165" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D5CDE53FC254E22FB1DF609" box="[996,1169,1367,1385]" italics="true" pageId="2" pageNumber="165">Tyrannosaurus rex</emphasis>
|
||
(
|
||
<bibRefCitation id="EFBCAAE44D5CDE53FB5F4E23FA19F609" author="CARR, T. D. & WILLIAMSON, T. E." box="[1182,1429,1365,1385]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="2" pageNumber="165" pagination="479 - 523" part="142" refId="ref7675" refString="CARR, T. D. AND WILLIAMSON, T. E., 2004, Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America: Zoological Journal of the Linnean Society, v. 142, p. 479 - 523." title="Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America" type="book chapter" year="2004">Carr and Williamson 2004</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. We also consulted figured tyrannosaurid material in the literature (e.g.,
|
||
<bibRefCitation id="EFBCAAE44D5CDE53FA6A4E07FCF2F6C3" author="BROCHU, C. A." journalOrPublisher="Journal of Vertebrate Paleontology" pageId="2" pageNumber="165" pagination="1 - 138" part="22" refId="ref7504" refString="BROCHU, C. A., 2003, Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull: Journal of Vertebrate Paleontology, v. 22, p. 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu 2003</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D5CDE52FCA44ED8FF23F5EC" blockId="2.[845,1518,1077,1934]" lastBlockId="3.[114,787,1122,1676]" lastPageId="3" lastPageNumber="166" pageId="2" pageNumber="165">
|
||
We prepared a cross-sectional thin section from the middle of
|
||
<materialsCitation id="3B45DD484D5CDE53FA6E4ED8FC3FF6BD" ID-GBIF-Occurrence="3396390334" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="2" pageNumber="165" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
to study the histology of the bone and determine the animal’s age at the time of its death. Because of the condition of the bone (
|
||
<figureCitation id="1316CB904D5CDE53FA6B4E93FA53F69A" box="[1450,1503,1510,1530]" captionStart="FIG. 1" captionStartId="2.[417,443,1015,1031]" captionTargetBox="[144,1515,176,986]" captionTargetId="figure@2.[144,1515,176,986]" captionTargetPageId="2" captionText="FIG. 1.—A Tyrannosaurus rex metatarsal IV (SWAU HRS13997) in four views. Scale bar = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.3749643" httpUri="https://zenodo.org/record/3749643/files/figure.png" pageId="2" pageNumber="165">Fig. 1</figureCitation>
|
||
), we choose to sample from a region with a complete record of the cortex and as close to the mid-diaphysis as possible. Although metatarsals are not commonly sampled, comparative histological sampling among different elements of the same individual of
|
||
<taxonomicName id="4C2DAC964D5CDE53FB3C4D2CFA62F50D" baseAuthorityName="Daudin" baseAuthorityYear="1802" box="[1277,1518,1625,1645]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="2" pageNumber="165" phylum="Chordata" rank="species" species="mississippiensis">
|
||
<emphasis id="B9590B074D5CDE53FB3C4D2CFA62F50D" box="[1277,1518,1625,1645]" italics="true" pageId="2" pageNumber="165">Alligator mississippiensis</emphasis>
|
||
</taxonomicName>
|
||
has shown that the mid-diaphysis of the metatarsals preserve a large amount of skeletochronological information (
|
||
<bibRefCitation id="EFBCAAE44D5CDE53FB2D4DE6FAEDF5C7" author="GARCIA, B. J." bookContentInfo="43 p." box="[1260,1377,1683,1703]" journalOrPublisher="Florida State University, Tallahassee" pageId="2" pageNumber="165" refId="ref8735" refString="GARCIA, B. J., 2011, Skeletochronology of the American Alligator (Alligator mississippiensis): examination of the utility of elements for histological study: Unpublished M. S. thesis, Florida State University, Tallahassee, 43 p." title="Skeletochronology of the American Alligator (Alligator mississippiensis): examination of the utility of elements for histological study" type="book" year="2011">Garcia 2011</bibRefCitation>
|
||
), and so we are confident that the sectioned tyrannosaurid metatarsal records an adequate record of growth. The proximal-most 40 mm of the bone was embedded in AeroMarine (www.aeromarineproducts.com) epoxy (#300) and hardener (#21) following the methods outlined by
|
||
<bibRefCitation id="EFBCAAE44D5CDE53FAF94C72FA31F47A" author="LAMM, E. - T." box="[1336,1469,1798,1818]" editor="K. Padian & E. - T. Lamm" journalOrPublisher="University of California Press, Berkeley, CA" pageId="2" pageNumber="165" pagination="55 - 160" refId="ref9446" refString="LAMM, E. - T., 2013, Preparation and sectioning of specimens, in K. Padian and E. - T. Lamm (eds.), Bone Histology of Fossil Tetrapods: Advancing Methods, Analysis, and Interpretation: University of California Press, Berkeley, CA, p. 55 - 160." title="Preparation and sectioning of specimens" type="book chapter" volumeTitle="Bone Histology of Fossil Tetrapods: Advancing Methods, Analysis, and Interpretation" year="2013">Lamm (2013)</bibRefCitation>
|
||
. The embedded sample was cut using a Covington 5047HG rock saw and ground with Covington diamond lapidary discs (NB1210, NB1220, NB1227, NB1232, and RB1260) using a series of diamond embedded wheels ranging from 100–600 grit. Once the embedded sample was polished and dried, the section was affixed to the frosted side of a 3
|
||
<emphasis id="B9590B074D5DDE52FD274F17FD70F719" box="[742,764,1122,1145]" italics="true" pageId="3" pageNumber="166">
|
||
<superScript id="7C587A5D4D5DDE52FD274F17FD60F710" attach="right" box="[742,748,1122,1136]" fontSize="6" pageId="3" pageNumber="166">′′</superScript>
|
||
×
|
||
</emphasis>
|
||
4
|
||
<emphasis id="B9590B074D5DDE52FCCB4F17FC9CF710" box="[778,784,1122,1136]" italics="true" pageId="3" pageNumber="166">
|
||
<superScript id="7C587A5D4D5DDE52FCCB4F17FC9CF710" attach="none" box="[778,784,1122,1136]" fontSize="6" pageId="3" pageNumber="166">′′</superScript>
|
||
</emphasis>
|
||
glass slide using two-ton epoxy (Devcon GLU-735.90), and allowed to cure overnight. The sample + slide were cut on a rock saw so that the glass slide retained a 1-inch section of embedded bone. The glass slide was then ground using a series of progressively finer grit wheels ending when the sample was evenly transparent (see
|
||
<bibRefCitation id="EFBCAAE44D5DDE52FE054F8FFDC8F66D" author="LAMM, E. - T." box="[452,580,1273,1293]" editor="K. Padian & E. - T. Lamm" journalOrPublisher="University of California Press, Berkeley, CA" pageId="3" pageNumber="166" pagination="55 - 160" refId="ref9446" refString="LAMM, E. - T., 2013, Preparation and sectioning of specimens, in K. Padian and E. - T. Lamm (eds.), Bone Histology of Fossil Tetrapods: Advancing Methods, Analysis, and Interpretation: University of California Press, Berkeley, CA, p. 55 - 160." title="Preparation and sectioning of specimens" type="book chapter" volumeTitle="Bone Histology of Fossil Tetrapods: Advancing Methods, Analysis, and Interpretation" year="2013">Lamm (2013)</bibRefCitation>
|
||
for more detail). The sample was finally hand-ground with 600 grit tungsten carbide powder until the desired opacity was obtained (
|
||
<emphasis id="B9590B074D5DDE52FE174E46FE6BF628" bold="true" box="[470,487,1331,1352]" pageId="3" pageNumber="166">~</emphasis>
|
||
70–100 microns thick). The thin section was viewed with a Nikon Eclipse LV100ND transmitted and reflected light microscope and imaged with a Nikon DS-Fi2 camera and Digital sight DS-U3 interface. High resolution images of the entire slide were captured with this microscope and camera using a Prior ProScan III automated microscope state and digitally assembled using Nikon NIS- Element Basic Research v. 4.40.00 (Build 1084). Images were captured in four light regimes: plane-polarized light, cross-polarized light, crosspolarized light with a gypsum plate (125 nm), and reflected light. Large high-resolution whole-slide images have been uploaded onto the online repository Morphobank (http://www.morphobank.org/, Project Number 2584).
|
||
</paragraph>
|
||
<caption id="DF52879D4D5DDE52FF4B4F7AFF68F758" ID-DOI="http://doi.org/10.5281/zenodo.3749645" ID-Zenodo-Dep="3749645" httpUri="https://zenodo.org/record/3749645/files/figure.png" pageId="3" pageNumber="166" startId="3.[138,164,1039,1055]" targetBox="[117,1489,176,1010]" targetPageId="3">
|
||
<paragraph id="8B92D7154D5DDE52FF4B4F7AFF68F758" blockId="3.[114,1490,1038,1081]" pageId="3" pageNumber="166">
|
||
FIG. 2.—Tooth traces on
|
||
<materialsCitation id="3B45DD484D5DDE52FE8E4F7AFE6BF740" ID-GBIF-Occurrence="3396390345" box="[335,487,1039,1056]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="3" pageNumber="166" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
. The two sets of
|
||
<taxonomicName id="4C2DAC964D5DDE52FDAD4F7BFCABF740" box="[620,807,1038,1056]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="166" phylum="Chordata" rank="species" species="paralleum">
|
||
<emphasis id="B9590B074D5DDE52FDAD4F7BFCABF740" box="[620,807,1038,1056]" italics="true" pageId="3" pageNumber="166">Knethichnus paralleum</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C2DAC964D5DDE52FC8F4F7BFC6FF77F" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[846,995,1038,1056]" class="Reptilia" genus="Linichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="166" phylum="Chordata" rank="species" species="serratus">
|
||
<emphasis id="B9590B074D5DDE52FC8F4F7BFC6FF77F" box="[846,995,1038,1056]" italics="true" pageId="3" pageNumber="166">Linichnus serratus</emphasis>
|
||
</taxonomicName>
|
||
are indicated, with the more prominent pair on the right. Scale bar = 20 mm.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8B92D7154D5DDE52FE5B4DCCFE60F5A9" blockId="3.[410,492,1721,1737]" box="[410,492,1721,1737]" pageId="3" pageNumber="166">
|
||
<heading id="D0DA60794D5DDE52FE5B4DCCFE60F5A9" allCaps="true" bold="true" box="[410,492,1721,1737]" centered="true" fontSize="7" level="3" pageId="3" pageNumber="166" reason="2">
|
||
<emphasis id="B9590B074D5DDE52FE5B4DCCFE60F5A9" bold="true" box="[410,492,1721,1737]" pageId="3" pageNumber="166">RESULTS</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D5DDE52FF4B4D92FD93F4EE" blockId="3.[114,787,1767,1934]" pageId="3" pageNumber="166">
|
||
<materialsCitation id="3B45DD484D5DDE52FF4B4D92FEBFF59A" ID-GBIF-Occurrence="3396390342" box="[138,307,1767,1786]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="3" pageNumber="166" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
shows no signs of weathering, although it does show stage 1 abrasion in the form of slight rounding to broken edges of the element, following the definition from
|
||
<bibRefCitation id="EFBCAAE44D5DDE52FDC84C57FD43F455" author="RYAN, M. J. & RUSSELL, A. P. & EBERTH, D. A. & CURRIE, P. J." box="[521,719,1825,1845]" journalOrPublisher="PALAIOS" pageId="3" pageNumber="166" pagination="482 - 506" part="16" refId="ref10459" refString="RYAN, M. J., RUSSELL, A. P., EBERTH, D. A., AND CURRIE, P. J., 2001, Taphonomy of a Centrosaurus (Ornithischia, Ceratopsidae) bone bed from the Dinosaur Park Formation (upper Campanian), Alberta, Canada, with comments on cranial ontogeny: PALAIOS, v. 16, p. 482 - 506." title="Taphonomy of a Centrosaurus (Ornithischia, Ceratopsidae) bone bed from the Dinosaur Park Formation (upper Campanian), Alberta, Canada, with comments on cranial ontogeny" type="journal article" year="2001">Ryan et al. (2001)</bibRefCitation>
|
||
. From comparison with a similar-sized
|
||
<taxonomicName id="4C2DAC964D5DDE52FE654C35FDA0F432" authorityName="Osborn" authorityYear="1905" box="[420,556,1856,1874]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="166" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D5DDE52FE654C35FDA0F432" box="[420,556,1856,1874]" italics="true" pageId="3" pageNumber="166">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
metatarsal on display at the Los Angeles County Museum (
|
||
<materialsCitation id="3B45DD484D5DDE52FE6C4C28FDA3F410" ID-GBIF-Occurrence="3396390339" box="[429,559,1884,1904]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/ContactSheet.php?QueryName=DetailedQuery&StartAt=1&QueryPage=%2Fdinosaur-institute%2Findex.php&Restriction=%28exists+%28SecDepartment_tab+where+%28SecDepartment+contains+%27Dinosaur+Institute%27%29%29%29&col_SpeSpecimenNoInteger=23845&LimitPerPage=20&Search=Search& " pageId="3" pageNumber="166" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
) we interpret the original length of
|
||
<materialsCitation id="3B45DD484D5DDE52FF0D4C0FFEFAF4ED" ID-GBIF-Occurrence="3396390310" box="[204,374,1914,1933]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="3" pageNumber="166" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
to be
|
||
<emphasis id="B9590B074D5DDE52FE724C0CFE48F4EE" bold="true" box="[435,452,1913,1934]" pageId="3" pageNumber="166">~</emphasis>
|
||
450 mm.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D5DDE52FB854F13FB4EF71A" blockId="3.[1092,1218,1126,1146]" box="[1092,1218,1126,1146]" pageId="3" pageNumber="166">
|
||
<heading id="D0DA60794D5DDE52FB854F13FB4EF71A" bold="true" box="[1092,1218,1126,1146]" centered="true" fontSize="8" level="2" pageId="3" pageNumber="166" reason="2">
|
||
<emphasis id="B9590B074D5DDE52FB854F13FB4EF71A" bold="true" box="[1092,1218,1126,1146]" italics="true" pageId="3" pageNumber="166">Tooth Traces</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D5DDE52FC8B4FE0FA48F558" blockId="3.[818,1491,1173,1934]" pageId="3" pageNumber="166">
|
||
The scratches on the proximal end of
|
||
<materialsCitation id="3B45DD484D5DDE52FB1D4FE3FA02F7C9" ID-GBIF-Occurrence="3396390340" box="[1244,1422,1174,1193]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="3" pageNumber="166" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
match descriptions of tooth scores found in the literature, having length to width ratios of greater than 3:1 and U or V-shaped cross sections (
|
||
<bibRefCitation id="EFBCAAE44D5DDE52FA924FBBFA43F782" author="BINFORD, L. R." box="[1363,1487,1230,1250]" journalOrPublisher="Academic Press, Inc., San Diego, CA" pageId="3" pageNumber="166" pagination="35 - 86" refId="ref7405" refString="BINFORD, L. R., 1981, Patterns of bone modifications produced by nonhuman agents, in Bones: Ancient Men and Modern Myths: Academic Press, Inc., San Diego, CA, p. 35 - 86." title="Patterns of bone modifications produced by nonhuman agents" type="book chapter" volumeTitle="Bones: Ancient Men and Modern Myths" year="1981">Binford 1981</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D5DDE52FCF34F9EFBC5F79F" author="NJAU, J. K. & BLUMENSCHINE, R. J." box="[818,1097,1259,1279]" journalOrPublisher="Journal of Human Evolution" pageId="3" pageNumber="166" pagination="142 - 162" part="50" refId="ref9872" refString="NJAU, J. K. AND BLUMENSCHINE, R. J., 2006, A diagnosis of crocodile feeding traces on larger mammal bone, with fossil examples from the Plio-Pleistocene Olduvai Basin, Tanzania: Journal of Human Evolution, v. 50, p. 142 - 162." title="A diagnosis of crocodile feeding traces on larger mammal bone, with fossil examples from the Plio-Pleistocene Olduvai Basin, Tanzania" type="journal article" year="2006">Njau and Blumenschine 2006</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D5DDE52FB964F9EFA82F79E" author="POBINER, B. L. & DESILVA, J. & SANDERS, W. J. & MITANI, J. C." box="[1111,1294,1259,1279]" journalOrPublisher="Journal of Human Evolution" pageId="3" pageNumber="166" pagination="614 - 636" part="52" refId="ref10320" refString="POBINER, B. L., DESILVA, J., SANDERS, W. J., AND MITANI, J. C., 2007, Taphonomic analysis of skeletal remains from chimpanzee hunts at Ngogo, Kibale National Park, Uganda: Journal of Human Evolution, v. 52, p. 614 - 636." title="Taphonomic analysis of skeletal remains from chimpanzee hunts at Ngogo, Kibale National Park, Uganda" type="journal article" year="2007">Pobiner et al. 2007</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D5DDE52FADD4F9EFA16F79F" author="POBINER, B." box="[1308,1434,1259,1279]" journalOrPublisher="Journal of Taphonomy" pageId="3" pageNumber="166" pagination="373 - 397" part="6" refId="ref10292" refString="POBINER, B., 2008, Paleoecological information in predator tooth marks: Journal of Taphonomy, v. 6, p. 373 - 397." title="Paleoecological information in predator tooth marks" type="journal article" year="2008">Pobiner 2008</bibRefCitation>
|
||
). Two examples each of the ichnotaxa
|
||
<taxonomicName id="4C2DAC964D5DDE52FBA84E72FC35F658" authority="(Jacobsen and Bromley 2009)" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="166" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5DDE52FBA84E72FAC9F67B" box="[1129,1349,1287,1307]" italics="true" pageId="3" pageNumber="166">Knethichnus parallelum</emphasis>
|
||
(
|
||
<bibRefCitation id="EFBCAAE44D5DDE52FA974E72FC3FF658" author="JACOBSEN, A. R. & BROMLEY, R. G." journalOrPublisher="Geological Quarterly" pageId="3" pageNumber="166" pagination="373 - 382" part="53" refId="ref9350" refString="JACOBSEN, A. R. AND BROMLEY, R. G., 2009, New ichnotaxa based on tooth impressions on dinosaur and whale bones: Geological Quarterly, v. 53, p. 373 - 382" title="New ichnotaxa based on tooth impressions on dinosaur and whale bones" type="journal article" year="2009">Jacobsen and Bromley 2009</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C2DAC964D5DDE52FC274E51FA10F658" authority="(Jacobsen and Bromley 2009)" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[998,1436,1316,1336]" class="Reptilia" genus="Linichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="166" phylum="Chordata" rank="species" species="serratus">
|
||
<emphasis id="B9590B074D5DDE52FC274E51FB07F657" box="[998,1163,1316,1336]" italics="true" pageId="3" pageNumber="166">Linichnus serratus</emphasis>
|
||
(
|
||
<bibRefCitation id="EFBCAAE44D5DDE52FB594E51FA1AF658" author="JACOBSEN, A. R. & BROMLEY, R. G." box="[1176,1430,1316,1336]" journalOrPublisher="Geological Quarterly" pageId="3" pageNumber="166" pagination="373 - 382" part="53" refId="ref9350" refString="JACOBSEN, A. R. AND BROMLEY, R. G., 2009, New ichnotaxa based on tooth impressions on dinosaur and whale bones: Geological Quarterly, v. 53, p. 373 - 382" title="New ichnotaxa based on tooth impressions on dinosaur and whale bones" type="journal article" year="2009">Jacobsen and Bromley 2009</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
occur in close proximity to one another, with one pair more prominent than the other (
|
||
<figureCitation id="1316CB904D5DDE52FCB34E28FC3AF610" box="[882,950,1373,1393]" captionStart="FIG. 2" captionStartId="3.[138,164,1039,1055]" captionTargetBox="[117,1489,176,1010]" captionTargetId="figure@3.[117,1489,176,1010]" captionTargetPageId="3" captionText="FIG. 2.—Tooth traces on SWAU HRS13997. The two sets of Knethichnus paralleum and Linichnus serratus are indicated, with the more prominent pair on the right. Scale bar = 20 mm." figureDoi="http://doi.org/10.5281/zenodo.3749645" httpUri="https://zenodo.org/record/3749645/files/figure.png" pageId="3" pageNumber="166">Figs. 2</figureCitation>
|
||
,
|
||
<figureCitation id="1316CB904D5DDE52FC044E28FC5DF611" box="[965,977,1373,1393]" captionStart="FIG. 3" captionStartId="4.[165,191,1014,1030]" captionTargetBox="[143,812,176,985]" captionTargetId="figure@4.[143,812,176,985]" captionTargetPageId="4" captionText="FIG. 3.—Details of Knethichnus parallelum and Linichnus serratus traces on SWAU HRS13997. A) Most prominent K. parallelum (center) and L. serratus (bottom) examples. B) Most prominent L. serratus in more detail and a different lighting angle. Small portion of the most prominent K. parallelum is also visible in the lower right corner. C) Detailed view of the other, less prominent L. serratus (left) and K. parallelum (right) traces. All scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3749647" httpUri="https://zenodo.org/record/3749647/files/figure.png" pageId="3" pageNumber="166">3</figureCitation>
|
||
). The
|
||
<taxonomicName id="4C2DAC964D5DDE52FBD64E28FB7FF611" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[1047,1267,1373,1393]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="166" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5DDE52FBD64E28FB7FF611" box="[1047,1267,1373,1393]" italics="true" pageId="3" pageNumber="166">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
trace is caused by the denticles on a serrated tooth dragging along the surface of a bone (
|
||
<bibRefCitation id="EFBCAAE44D5DDE52FA434E0CFC56F6C9" author="JACOBSEN, A. R. & BROMLEY, R. G." journalOrPublisher="Geological Quarterly" pageId="3" pageNumber="166" pagination="373 - 382" part="53" refId="ref9350" refString="JACOBSEN, A. R. AND BROMLEY, R. G., 2009, New ichnotaxa based on tooth impressions on dinosaur and whale bones: Geological Quarterly, v. 53, p. 373 - 382" title="New ichnotaxa based on tooth impressions on dinosaur and whale bones" type="journal article" year="2009">Jacobsen and Bromley 2009</bibRefCitation>
|
||
) as demonstrated by actualistic experiments on Komodo dragon (
|
||
<taxonomicName id="4C2DAC964D5DDE52FC414EC6FBCBF6A6" box="[896,1095,1458,1478]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="3" pageNumber="166" phylum="Chordata" rank="species" species="komodoensis">
|
||
<emphasis id="B9590B074D5DDE52FC414EC6FBCBF6A6" box="[896,1095,1458,1478]" italics="true" pageId="3" pageNumber="166">Varanus komodoensis</emphasis>
|
||
</taxonomicName>
|
||
) feeding behavior (
|
||
<bibRefCitation id="EFBCAAE44D5DDE52FB3F4EC6FC2DF682" author="D'AMORE, D. C. & BLUMENSCHINE, R. J." journalOrPublisher="Paleobiology" pageId="3" pageNumber="166" pagination="79 - 100" part="38" refId="ref8105" refString="D'AMORE, D. C. AND BLUMENSCHINE, R. J., 2012, Using striated tooth marks on bone to predict body size in theropod dinosaurs: a model based on feeding observations of Varanus komodoensis, the Komodo monitor: Paleobiology, v. 38, p. 79 - 100." title="Using striated tooth marks on bone to predict body size in theropod dinosaurs: a model based on feeding observations of Varanus komodoensis, the Komodo monitor" type="journal article" year="2012">D’Amore and Blumenschine 2012</bibRefCitation>
|
||
). If the fauna of the formation in question is well known, as is the case with the heavily studied Lance Formation, then the denticle widths on the teeth from the present carnivorous taxa can be compared to the striation widths of the
|
||
<taxonomicName id="4C2DAC964D5DDE52FC3E4D51FB5AF558" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[1023,1238,1572,1592]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="166" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5DDE52FC3E4D51FB5AF558" box="[1023,1238,1572,1592]" italics="true" pageId="3" pageNumber="166">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
trace to look for a match.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D5DDE52FC8B4D35FBBAF458" blockId="3.[818,1491,1173,1934]" pageId="3" pageNumber="166">
|
||
Tooth scores are located on the distal end of the bone. The scores have a U-shaped cross-section and the grooves vary in width from 0.6–6 mm. The proximal-most trace is made of three separate scores which merge into a single score, the deepest trace on the bone. Distal to this trace are the prominent examples of
|
||
<taxonomicName id="4C2DAC964D5DDE52FBD04DC7FB31F5A5" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[1041,1213,1714,1734]" class="Reptilia" genus="Linichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="166" phylum="Chordata" rank="species" species="serratus">
|
||
<emphasis id="B9590B074D5DDE52FBD04DC7FB31F5A5" box="[1041,1213,1714,1734]" italics="true" pageId="3" pageNumber="166">Linichnus serratus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C2DAC964D5DDE52FB2E4DC7FA41F5A6" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[1263,1485,1714,1734]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="166" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5DDE52FB2E4DC7FA41F5A6" box="[1263,1485,1714,1734]" italics="true" pageId="3" pageNumber="166">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
. This
|
||
<taxonomicName id="4C2DAC964D5DDE52FCA54DBAFBCDF583" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[868,1089,1743,1763]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="166" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5DDE52FCA54DBAFBCDF583" box="[868,1089,1743,1763]" italics="true" pageId="3" pageNumber="166">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
trace cuts across an earlier score (
|
||
<figureCitation id="1316CB904D5DDE52FA4C4DBAFA4AF583" box="[1421,1478,1743,1763]" captionStart="FIG. 3" captionStartId="4.[165,191,1014,1030]" captionTargetBox="[143,812,176,985]" captionTargetId="figure@4.[143,812,176,985]" captionTargetPageId="4" captionText="FIG. 3.—Details of Knethichnus parallelum and Linichnus serratus traces on SWAU HRS13997. A) Most prominent K. parallelum (center) and L. serratus (bottom) examples. B) Most prominent L. serratus in more detail and a different lighting angle. Small portion of the most prominent K. parallelum is also visible in the lower right corner. C) Detailed view of the other, less prominent L. serratus (left) and K. parallelum (right) traces. All scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.3749647" httpUri="https://zenodo.org/record/3749647/files/figure.png" pageId="3" pageNumber="166">Fig. 3</figureCitation>
|
||
). Posterior to these traces is an assortment of small, shallow scores. The striation width (SW) of the
|
||
<taxonomicName id="4C2DAC964D5DDE52FBED4C7DFA8AF47C" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[1068,1286,1800,1820]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="166" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5DDE52FBED4C7DFA8AF47C" box="[1068,1286,1800,1820]" italics="true" pageId="3" pageNumber="166">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
on
|
||
<materialsCitation id="3B45DD484D5DDE52FAEB4C7DFA5FF47B" ID-GBIF-Occurrence="3396390319" box="[1322,1491,1800,1819]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="3" pageNumber="166" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
is three striations per 2 mm.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D5DDE55FC8B4C37FDDAF6D7" blockId="3.[818,1491,1173,1934]" lastBlockId="4.[141,814,1202,1464]" lastPageId="4" lastPageNumber="167" pageId="3" pageNumber="166">
|
||
Measurements of average denticle width (DW) on the mesial and distal carinae (MDW and DDW, respectively) on theropod teeth from the Lance Formation (
|
||
<tableCitation id="C6AFE2AE4D5DDE52FC614C0FFC64F4EE" box="[928,1000,1914,1934]" captionStart="TABLE 1" captionStartId="4.[141,195,1652,1671]" captionTargetBox="[141,1517,1751,1916]" captionTargetPageId="4" captionText="TABLE 1.—Denticle measurements for theropod taxa known from the Lance Formation. Abbreviations: DD/2 mm = distal denticles per two millimeters; DDW = distal denticle width; MD/2 mm = mesial denticles per two millimeters; MDW = mesial denticle width." httpUri="http://table.plazi.org/id/DF52879D4D5ADE55FF4C4D01FAC8F5C3" pageId="3" pageNumber="166" tableUuid="DF52879D4D5ADE55FF4C4D01FAC8F5C3">Table 1</tableCitation>
|
||
) compared with the striation width (SW) of three striations per 2 mm on the
|
||
<taxonomicName id="4C2DAC964D5ADE55FE664FC7FD0BF7A6" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[423,647,1202,1222]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5ADE55FE664FC7FD0BF7A6" box="[423,647,1202,1222]" italics="true" pageId="4" pageNumber="167">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
trace on
|
||
<materialsCitation id="3B45DD484D5ADE55FD2E4FC6FF7EF784" ID-GBIF-Occurrence="3396390321" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="4" pageNumber="167" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
demonstrate that the DW of most Lance Formation theropods are too small to have created the SW on
|
||
<materialsCitation id="3B45DD484D5ADE55FDC74F9AFD3FF662" ID-GBIF-Occurrence="3396390318" box="[518,691,1263,1282]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="4" pageNumber="167" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
. In fact, only the DW on teeth of
|
||
<taxonomicName id="4C2DAC964D5ADE55FE884E7BFE7AF640" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[329,502,1294,1312]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D5ADE55FE884E7BFE7AF640" box="[329,502,1294,1312]" italics="true" pageId="4" pageNumber="167">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
was large enough to produce the observed SW. As noted by
|
||
<bibRefCitation id="EFBCAAE44D5ADE55FE4D4E59FD54F65F" author="D'AMORE, D. C. & BLUMENSCHINE, R. J." box="[396,728,1323,1343]" journalOrPublisher="Paleobiology" pageId="4" pageNumber="167" pagination="79 - 100" part="38" refId="ref8105" refString="D'AMORE, D. C. AND BLUMENSCHINE, R. J., 2012, Using striated tooth marks on bone to predict body size in theropod dinosaurs: a model based on feeding observations of Varanus komodoensis, the Komodo monitor: Paleobiology, v. 38, p. 79 - 100." title="Using striated tooth marks on bone to predict body size in theropod dinosaurs: a model based on feeding observations of Varanus komodoensis, the Komodo monitor" type="journal article" year="2012">D’Amore and Blumenschine (2012)</bibRefCitation>
|
||
, striation widths on a
|
||
<taxonomicName id="4C2DAC964D5ADE55FF3F4E3CFE56F63D" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[254,474,1353,1373]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5ADE55FF3F4E3CFE56F63D" box="[254,474,1353,1373]" italics="true" pageId="4" pageNumber="167">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
trace can be equal to or smaller than the denticle width of the biter’s tooth, but never larger. Thus, it is not possible that the
|
||
<taxonomicName id="4C2DAC964D5ADE55FEE64EF0FD8DF6F9" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[295,513,1413,1433]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5ADE55FEE64EF0FD8DF6F9" box="[295,513,1413,1433]" italics="true" pageId="4" pageNumber="167">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
could have been made by any of the taxa in
|
||
<tableCitation id="C6AFE2AE4D5ADE55FF374ED1FEB5F6D7" box="[246,313,1444,1464]" captionStart="TABLE 1" captionStartId="4.[141,195,1652,1671]" captionTargetBox="[141,1517,1751,1916]" captionTargetPageId="4" captionText="TABLE 1.—Denticle measurements for theropod taxa known from the Lance Formation. Abbreviations: DD/2 mm = distal denticles per two millimeters; DDW = distal denticle width; MD/2 mm = mesial denticles per two millimeters; MDW = mesial denticle width." httpUri="http://table.plazi.org/id/DF52879D4D5ADE55FF4C4D01FAC8F5C3" pageId="4" pageNumber="167" tableUuid="DF52879D4D5ADE55FF4C4D01FAC8F5C3">Table 1</tableCitation>
|
||
except for
|
||
<taxonomicName id="4C2DAC964D5ADE55FE654ED0FDDCF6D7" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[420,592,1445,1463]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D5ADE55FE654ED0FDDCF6D7" box="[420,592,1445,1463]" italics="true" pageId="4" pageNumber="167">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<caption id="DF52879D4D5ADE55FF644883FDDBF7E4" ID-DOI="http://doi.org/10.5281/zenodo.3749647" ID-Zenodo-Dep="3749647" httpUri="https://zenodo.org/record/3749647/files/figure.png" pageId="4" pageNumber="167" startId="4.[165,191,1014,1030]" targetBox="[143,812,176,985]" targetPageId="4">
|
||
<paragraph id="8B92D7154D5ADE55FF644883FDDBF7E4" blockId="4.[141,814,1013,1157]" pageId="4" pageNumber="167">
|
||
FIG. 3.—Details of
|
||
<taxonomicName id="4C2DAC964D5ADE55FE8A4880FD9CF767" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[331,528,1013,1031]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5ADE55FE8A4880FD9CF767" box="[331,528,1013,1031]" italics="true" pageId="4" pageNumber="167">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C2DAC964D5ADE55FD814880FD55F766" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[576,729,1013,1031]" class="Reptilia" genus="Linichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="serratus">
|
||
<emphasis id="B9590B074D5ADE55FD814880FD55F766" box="[576,729,1013,1031]" italics="true" pageId="4" pageNumber="167">Linichnus serratus</emphasis>
|
||
</taxonomicName>
|
||
traces on
|
||
<materialsCitation id="3B45DD484D5ADE55FF4C4F7AFEA6F740" ID-GBIF-Occurrence="3396390322" box="[141,298,1039,1056]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="4" pageNumber="167" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
.
|
||
<emphasis id="B9590B074D5ADE55FEF64F7AFEC9F741" bold="true" box="[311,325,1039,1057]" pageId="4" pageNumber="167">A</emphasis>
|
||
) Most prominent
|
||
<taxonomicName id="4C2DAC964D5ADE55FE244F65FDD5F741" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[485,601,1039,1057]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5ADE55FE244F65FDD5F741" box="[485,601,1039,1057]" italics="true" pageId="4" pageNumber="167">K. parallelum</emphasis>
|
||
</taxonomicName>
|
||
(center) and
|
||
<taxonomicName id="4C2DAC964D5ADE55FD134F65FCA1F740" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[722,813,1040,1056]" class="Reptilia" genus="Linichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="serratus">
|
||
<emphasis id="B9590B074D5ADE55FD134F65FCA1F740" box="[722,813,1040,1056]" italics="true" pageId="4" pageNumber="167">L. serratus</emphasis>
|
||
</taxonomicName>
|
||
(bottom) examples.
|
||
<emphasis id="B9590B074D5ADE55FEF34F5DFEB3F75A" bold="true" box="[306,319,1064,1082]" pageId="4" pageNumber="167">B</emphasis>
|
||
) Most prominent
|
||
<taxonomicName id="4C2DAC964D5ADE55FE194F5CFDBDF759" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[472,561,1065,1081]" class="Reptilia" genus="Linichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="serratus">
|
||
<emphasis id="B9590B074D5ADE55FE194F5CFDBDF759" box="[472,561,1065,1081]" italics="true" pageId="4" pageNumber="167">L. serratus</emphasis>
|
||
</taxonomicName>
|
||
in more detail and a different lighting angle. Small portion of the most prominent
|
||
<taxonomicName id="4C2DAC964D5ADE55FDF54F37FD29F733" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[564,677,1089,1107]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5ADE55FDF54F37FD29F733" box="[564,677,1089,1107]" italics="true" pageId="4" pageNumber="167">K. parallelum</emphasis>
|
||
</taxonomicName>
|
||
is also visible in the lower right corner.
|
||
<emphasis id="B9590B074D5ADE55FE834F2FFEDCF70C" bold="true" box="[322,336,1114,1132]" pageId="4" pageNumber="167">C</emphasis>
|
||
) Detailed view of the other, less prominent
|
||
<taxonomicName id="4C2DAC964D5ADE55FD6A4F2EFC8DF70B" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[683,769,1115,1131]" class="Reptilia" genus="Linichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="serratus">
|
||
<emphasis id="B9590B074D5ADE55FD6A4F2EFC8DF70B" box="[683,769,1115,1131]" italics="true" pageId="4" pageNumber="167">L. serratus</emphasis>
|
||
</taxonomicName>
|
||
(left) and
|
||
<taxonomicName id="4C2DAC964D5ADE55FF704F00FEAEF7E5" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[177,290,1139,1157]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="4" pageNumber="167" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D5ADE55FF704F00FEAEF7E5" box="[177,290,1139,1157]" italics="true" pageId="4" pageNumber="167">K. parallelum</emphasis>
|
||
</taxonomicName>
|
||
(right) traces. All scale bars = 1 mm.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8B92D7154D5ADE55FE704E96FD86F697" blockId="4.[433,522,1507,1527]" box="[433,522,1507,1527]" pageId="4" pageNumber="167">
|
||
<heading id="D0DA60794D5ADE55FE704E96FD86F697" bold="true" box="[433,522,1507,1527]" centered="true" fontSize="8" level="2" pageId="4" pageNumber="167" reason="2">
|
||
<emphasis id="B9590B074D5ADE55FE704E96FD86F697" bold="true" box="[433,522,1507,1527]" italics="true" pageId="4" pageNumber="167">Histology</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D5ADE55FF644D60FBABF1A1" blockId="4.[141,814,1557,1607]" lastBlockId="4.[845,1519,175,1471]" pageId="4" pageNumber="167">
|
||
This histological description uses the terminology of Francillon-Vieillot et al. (1990), as described and expanded on by
|
||
<bibRefCitation id="EFBCAAE44D5ADE55FDA74D46FD71F527" author="WERNING, S. A." bookContentInfo="445 p." box="[614,765,1587,1607]" journalOrPublisher="University of California, Berkeley" pageId="4" pageNumber="167" refId="ref10961" refString="WERNING, S. A., 2013, Evolution of bone histological characters in amniotes, and the implications for the evolution of growth and metabolism: Unpublished Ph. D. dissertation, University of California, Berkeley, 445 p." title="Evolution of bone histological characters in amniotes, and the implications for the evolution of growth and metabolism" type="book" year="2013">Werning (2013)</bibRefCitation>
|
||
. The histological thin section of
|
||
<materialsCitation id="3B45DD484D5ADE55FB874BC5FB63F3A3" ID-GBIF-Occurrence="3396390350" box="[1094,1263,176,195]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="4" pageNumber="167" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
is narrow dorsally, where it forms a rounded apex, with a wide, round ventral surface. The medullary cavity is hollow and the endosteal surface is jagged (
|
||
<figureCitation id="1316CB904D5ADE55FA874B9DFA1EF39C" box="[1350,1426,232,252]" captionStart="FIG. 4" captionStartId="6.[141,167,1865,1881]" captionTargetId="figure@5.[148,1457,176,1930]" captionTargetPageId="5" captionText="FIG. 4.—Histology of a Tyrannosaurus rex metatarsal (SWAU HRS13997) in cross section under different light regimes. A) Section of cortical bone from the medullary cavity (left) to the subperiosteal surface (right) under regularly transmitted light. Boxes show where areas of this bone are shown in higher detail under regularly transmitted light (B, F, J), cross-polarized light (C, G, K), cross-polarized light with a waveplate (D, H, L), and reflected light (E, I, M). Scale bar for A = 5 mm, all other scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.4023994" httpUri="https://zenodo.org/record/4023994/files/figure.png" pageId="4" pageNumber="167" targetBox="[156,1437,176,1928]" targetPageId="5">Fig. 4A</figureCitation>
|
||
), with no evidence of endosteal lamellae or medullary bone. The inner cortex is composed of highly woven-fibered, unremodeled primary bone (
|
||
<figureCitation id="1316CB904D5ADE55FA5D4A55FA5FF254" box="[1436,1491,288,308]" captionStart="FIG. 4" captionStartId="6.[141,167,1865,1881]" captionTargetId="figure@5.[148,1457,176,1930]" captionTargetPageId="5" captionText="FIG. 4.—Histology of a Tyrannosaurus rex metatarsal (SWAU HRS13997) in cross section under different light regimes. A) Section of cortical bone from the medullary cavity (left) to the subperiosteal surface (right) under regularly transmitted light. Boxes show where areas of this bone are shown in higher detail under regularly transmitted light (B, F, J), cross-polarized light (C, G, K), cross-polarized light with a waveplate (D, H, L), and reflected light (E, I, M). Scale bar for A = 5 mm, all other scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.4023994" httpUri="https://zenodo.org/record/4023994/files/figure.png" pageId="4" pageNumber="167" targetBox="[156,1437,176,1928]" targetPageId="5">Fig. 4</figureCitation>
|
||
B– 4E). Both longitudinal simple vascular canals and primary osteons are common, with primary Haversian cavities present but less common, and most vascular cavities anastomose in no preferred orientation. There is no evidence of secondary remodeling, either by the presence of erosion rooms or secondary osteons. Osteocyte lacunae possess no preferred orientation in this region except for some of the lacunae in the lamellar bone of the primary osteons, which are usually oriented circumferentially around the vascular cavity of the osteon. Although the innermost portion (1.25–2 mm) of the cortex, including the endosteal border, is a deep reddish-brown, the next most external cortical bone is light tan, forming a tan-colored circumferential band 1–2 mm thick; the outermost 2–5 mm of the cortex is also light tan in color (
|
||
<figureCitation id="1316CB904D5ADE55FBF74900FB08F1E9" box="[1078,1156,629,649]" captionStart="FIG. 4" captionStartId="6.[141,167,1865,1881]" captionTargetId="figure@5.[148,1457,176,1930]" captionTargetPageId="5" captionText="FIG. 4.—Histology of a Tyrannosaurus rex metatarsal (SWAU HRS13997) in cross section under different light regimes. A) Section of cortical bone from the medullary cavity (left) to the subperiosteal surface (right) under regularly transmitted light. Boxes show where areas of this bone are shown in higher detail under regularly transmitted light (B, F, J), cross-polarized light (C, G, K), cross-polarized light with a waveplate (D, H, L), and reflected light (E, I, M). Scale bar for A = 5 mm, all other scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.4023994" httpUri="https://zenodo.org/record/4023994/files/figure.png" pageId="4" pageNumber="167" targetBox="[156,1437,176,1928]" targetPageId="5">Fig. 4A</figureCitation>
|
||
). This difference in coloration is not correlated with bone tissue or texture, indicating these colored bands are not biological in origin.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D5ADE55FCA449BFFAD9F6DF" blockId="4.[845,1519,175,1471]" pageId="4" pageNumber="167">
|
||
Roughly one-fourth to one-half the distance from the endosteal surface to the subperiosteal surface, the vascular style shifts from being predominantly longitudinal canals to circumferential canals, and anastomosing canals become rare in the external-most 2–3 mm of the cortex. The
|
||
<emphasis id="B9590B074D5ADE55FA46486BFA14F053" bold="true" box="[1415,1432,798,819]" pageId="4" pageNumber="167">~</emphasis>
|
||
2–4 mm closest to the subperiosteal surface possesses patches of more organized parallel-fibered bone which become more common in the external-most region of the cortex, and in these regions osteocyte lacunae are preferentially oriented with the collagen fibers. The dorsal-most region of the cortex is the exception to this description, with highly disorganized woven bone and longitudinal canals predominating to the subperiosteal surface. There are no secondary osteons or erosion rooms in the cortical bone. Although there are several circumferential patterns in the cortical bone, close examination reveals these to be neither lines of arrested growth (LAGs), diagnosed by continuous cement lines (which are apparent in reflected light but absent in this specimen;
|
||
<figureCitation id="1316CB904D5ADE55FC1A4F22FBF8F70B" box="[987,1140,1111,1131]" captionStart="FIG. 4" captionStartId="6.[141,167,1865,1881]" captionTargetId="figure@5.[148,1457,176,1930]" captionTargetPageId="5" captionText="FIG. 4.—Histology of a Tyrannosaurus rex metatarsal (SWAU HRS13997) in cross section under different light regimes. A) Section of cortical bone from the medullary cavity (left) to the subperiosteal surface (right) under regularly transmitted light. Boxes show where areas of this bone are shown in higher detail under regularly transmitted light (B, F, J), cross-polarized light (C, G, K), cross-polarized light with a waveplate (D, H, L), and reflected light (E, I, M). Scale bar for A = 5 mm, all other scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.4023994" httpUri="https://zenodo.org/record/4023994/files/figure.png" pageId="4" pageNumber="167" targetBox="[156,1437,176,1928]" targetPageId="5">Fig. 4E, 4I, 4M</figureCitation>
|
||
), or annuli, diagnosed by thin bands of continuous avascular parallel-fibered or lamellar bone (especially apparent in cross-polarized light, but absent in this specimen;
|
||
<figureCitation id="1316CB904D5ADE55FA824FFAFA68F7C3" box="[1347,1508,1167,1187]" captionStart="FIG. 4" captionStartId="6.[141,167,1865,1881]" captionTargetId="figure@5.[148,1457,176,1930]" captionTargetPageId="5" captionText="FIG. 4.—Histology of a Tyrannosaurus rex metatarsal (SWAU HRS13997) in cross section under different light regimes. A) Section of cortical bone from the medullary cavity (left) to the subperiosteal surface (right) under regularly transmitted light. Boxes show where areas of this bone are shown in higher detail under regularly transmitted light (B, F, J), cross-polarized light (C, G, K), cross-polarized light with a waveplate (D, H, L), and reflected light (E, I, M). Scale bar for A = 5 mm, all other scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.4023994" httpUri="https://zenodo.org/record/4023994/files/figure.png" pageId="4" pageNumber="167" targetBox="[156,1437,176,1928]" targetPageId="5">Fig. 4C, 4G, 4K</figureCitation>
|
||
). These circumferential patterns are patterns in coloration, or in a few instances represent bands of increased osteocyte density, but are not composed of cement lines or avascular regions of lamellar bone. Therefore, growth marks are absent, and the bone contains anastomosing and interconnected vascular cavities up to the subperiosteal surface. Although the overall vascular density does not noticeably decrease from the medullary cavity to the subperiosteal surface (remaining well to very well vascularized throughout, sensu
|
||
<bibRefCitation id="EFBCAAE44D5ADE55FC3F4E07FB0AF6E6" author="WERNING, S. A." bookContentInfo="445 p." box="[1022,1158,1394,1414]" journalOrPublisher="University of California, Berkeley" pageId="4" pageNumber="167" refId="ref10961" refString="WERNING, S. A., 2013, Evolution of bone histological characters in amniotes, and the implications for the evolution of growth and metabolism: Unpublished Ph. D. dissertation, University of California, Berkeley, 445 p." title="Evolution of bone histological characters in amniotes, and the implications for the evolution of growth and metabolism" type="book" year="2013">Werning 2013</bibRefCitation>
|
||
;
|
||
<figureCitation id="1316CB904D5ADE55FB574E07FB5EF6E6" box="[1174,1234,1394,1414]" captionStart="FIG. 4" captionStartId="6.[141,167,1865,1881]" captionTargetId="figure@5.[148,1457,176,1930]" captionTargetPageId="5" captionText="FIG. 4.—Histology of a Tyrannosaurus rex metatarsal (SWAU HRS13997) in cross section under different light regimes. A) Section of cortical bone from the medullary cavity (left) to the subperiosteal surface (right) under regularly transmitted light. Boxes show where areas of this bone are shown in higher detail under regularly transmitted light (B, F, J), cross-polarized light (C, G, K), cross-polarized light with a waveplate (D, H, L), and reflected light (E, I, M). Scale bar for A = 5 mm, all other scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.4023994" httpUri="https://zenodo.org/record/4023994/files/figure.png" pageId="4" pageNumber="167" targetBox="[156,1437,176,1928]" targetPageId="5">Fig. 4</figureCitation>
|
||
), there is a shift in vascular orientation in the cortical bone closer to the subperiosteal surface, so that the inner half of the cortex possesses larger vascular cavities.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C337849E4D5ADE59FBA64E9FFA16F78C" lastPageId="8" lastPageNumber="171" pageId="4" pageNumber="167" type="discussion">
|
||
<paragraph id="8B92D7154D5ADE55FBA64E9FFB58F69A" blockId="4.[1127,1236,1514,1530]" box="[1127,1236,1514,1530]" pageId="4" pageNumber="167">
|
||
<heading id="D0DA60794D5ADE55FBA64E9FFB58F69A" allCaps="true" bold="true" box="[1127,1236,1514,1530]" centered="true" fontSize="7" level="3" pageId="4" pageNumber="167" reason="2">
|
||
<emphasis id="B9590B074D5ADE55FBA64E9FFB58F69A" bold="true" box="[1127,1236,1514,1530]" pageId="4" pageNumber="167">DISCUSSION</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D5ADE56FCA44D63FECDF39C" blockId="4.[845,1518,1558,1607]" lastBlockId="7.[114,788,175,1934]" lastPageId="7" lastPageNumber="170" pageId="4" pageNumber="167">
|
||
Although
|
||
<materialsCitation id="3B45DD484D5ADE55FC034D62FBE0F54A" ID-GBIF-Occurrence="3396390348" box="[962,1132,1559,1578]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="4" pageNumber="167" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
is fragmentary, we are confident that this bone is a tyrannosaurid metatarsal, specifically, metatarsal IV, because of similarities between it and the metatarsals of the mounted ‘‘subadult’’
|
||
<taxonomicName id="4C2DAC964D58DE57FF4C4E22FE99F609" authorityName="Osborn" authorityYear="1905" box="[141,277,1367,1385]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="169" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D58DE57FF4C4E22FE99F609" box="[141,277,1367,1385]" italics="true" pageId="6" pageNumber="169">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
on display at the LACM, which is based on material from
|
||
<materialsCitation id="3B45DD484D58DE57FF4C4E01FE82F6E7" ID-GBIF-Occurrence="3396390352" box="[141,270,1395,1415]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/ContactSheet.php?QueryName=DetailedQuery&StartAt=1&QueryPage=%2Fdinosaur-institute%2Findex.php&Restriction=%28exists+%28SecDepartment_tab+where+%28SecDepartment+contains+%27Dinosaur+Institute%27%29%29%29&col_SpeSpecimenNoInteger=23845&LimitPerPage=20&Search=Search& " pageId="6" pageNumber="169" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
(
|
||
<figureCitation id="1316CB904D58DE57FEDF4E06FE03F6E7" box="[286,399,1395,1415]" captionStart="FIG. 5" captionStartId="6.[165,191,1174,1190]" captionTargetBox="[145,1514,176,1144]" captionTargetId="figure@6.[144,1514,176,1144]" captionTargetPageId="6" captionText="FIG. 5.—Comparison between four specimens of tyrannosaurid metatarsals: A) Tyrannosaurus rex metatarsal IV (SWAU HRS13997). B) Tyrannosaurus rex metatarsals II-IV (LACM 23845, with IV sculpted). C) Tyrannosauridae indet. metatarsal IV (IGM 6130; modified from Peecook (2014, fig. 3). D) Tyrannosaurus rex metatarsals II-IV (FMNH PR2081; modified from Brochu (2003, figs. 99, 100, 101). SWAU HRS13997 and IGM 6130 are metatarsal IV. All elements are from the right pes except IGM 6130, which is a left element, although the image has been flipped to make it match the others. Note that FMNH PR2081 is shown in a more medial view than the other metatarsal specimens. Scale bar = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.3749649" httpUri="https://zenodo.org/record/3749649/files/figure.png" pageId="6" pageNumber="169">Fig. 5A, 5B</figureCitation>
|
||
,
|
||
<httpUri id="B5AC09BF4D58DE57FE5D4E06FD30F6E7" box="[412,700,1395,1415]" httpUri="https://www.sepm.org/supplemental-materials/165/view" pageId="6" pageNumber="169">online Supplemental File fig. 1</httpUri>
|
||
). The fossil specimen
|
||
<materialsCitation id="3B45DD484D58DE57FF2A4EE4FEE0F6C4" ID-GBIF-Occurrence="3396390311" box="[235,364,1424,1444]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/ContactSheet.php?QueryName=DetailedQuery&StartAt=1&QueryPage=%2Fdinosaur-institute%2Findex.php&Restriction=%28exists+%28SecDepartment_tab+where+%28SecDepartment+contains+%27Dinosaur+Institute%27%29%29%29&col_SpeSpecimenNoInteger=23845&LimitPerPage=20&Search=Search& " pageId="6" pageNumber="169" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
includes an almost complete right foot missing metatarsals I and IV and the phalanges from the first toe (
|
||
<bibRefCitation id="EFBCAAE44D58DE57FD6F4ED8FCA6F6A1" author="MOLNAR, R. E." box="[686,810,1453,1473]" journalOrPublisher="Journal of Paleontology" pageId="6" pageNumber="169" pagination="102 - 108" part="54" refId="ref9732" refString="MOLNAR, R. E., 1980, An albertosaur from the Hell Creek Formation of Montana: Journal of Paleontology, v. 54, p. 102 - 108." title="An albertosaur from the Hell Creek Formation of Montana" type="journal article" year="1980">Molnar 1980</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D58DE57FF4C4EB9FE8AF6BF" author="LARSON, N." box="[141,262,1483,1503]" editor="P. Larson & K. Carpenter" journalOrPublisher="Indiana University Press, Bloomington, Indiana" pageId="6" pageNumber="169" pagination="1 - 55" refId="ref9507" refString="LARSON, N., 2008, One hundred years of Tyrannosaurus rex: the skeletons, in P. Larson and K. Carpenter (eds.), Tyrannosaurus rex: The Tyrant King: Indiana University Press, Bloomington, Indiana, p. 1 - 55." title="One hundred years of Tyrannosaurus rex: the skeletons" type="book chapter" volumeTitle="Tyrannosaurus rex: The Tyrant King" year="2008">Larson 2008</bibRefCitation>
|
||
). Thus, the metatarsal IV on display is not original but is sculpted based on metatarsals II and III and references to other specimens (P. Fraley, personal communication). Nevertheless, the sculpted metatarsal IV is a relatively accurate model, as evidenced by its similarity to known material from skeletally immature tyrannosaurids, such as ‘‘Jane’’ and IGM 6130 (
|
||
<bibRefCitation id="EFBCAAE44D58DE57FF094D28FEF3F511" author="PEECOOK, B. R. & WILSON, J. A. & HERNANDEZ- RIVERA, R. & MONTELLANO- BALLESTEROS, M. & WILSON, G. P." box="[200,383,1629,1649]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="6" pageNumber="169" pagination="71 - 80" part="59" refId="ref10144" refString="PEECOOK, B. R., WILSON, J. A., HERNANDEZ- RIVERA, R., MONTELLANO- BALLESTEROS, M., AND WILSON, G. P., 2014, First tyrannosaurid remains from the Upper Cretaceous '' El Gallo '' Formation of Baja California, Mexico: Acta Palaeontologica Polonica, v. 59, p. 71 - 80." title="First tyrannosaurid remains from the Upper Cretaceous '' El Gallo '' Formation of Baja California, Mexico" type="journal article" year="2014">Peecook et al. 2014</bibRefCitation>
|
||
) (
|
||
<figureCitation id="1316CB904D58DE57FE524D28FE57F511" box="[403,475,1629,1649]" captionStart="FIG. 5" captionStartId="6.[165,191,1174,1190]" captionTargetBox="[145,1514,176,1144]" captionTargetId="figure@6.[144,1514,176,1144]" captionTargetPageId="6" captionText="FIG. 5.—Comparison between four specimens of tyrannosaurid metatarsals: A) Tyrannosaurus rex metatarsal IV (SWAU HRS13997). B) Tyrannosaurus rex metatarsals II-IV (LACM 23845, with IV sculpted). C) Tyrannosauridae indet. metatarsal IV (IGM 6130; modified from Peecook (2014, fig. 3). D) Tyrannosaurus rex metatarsals II-IV (FMNH PR2081; modified from Brochu (2003, figs. 99, 100, 101). SWAU HRS13997 and IGM 6130 are metatarsal IV. All elements are from the right pes except IGM 6130, which is a left element, although the image has been flipped to make it match the others. Note that FMNH PR2081 is shown in a more medial view than the other metatarsal specimens. Scale bar = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.3749649" httpUri="https://zenodo.org/record/3749649/files/figure.png" pageId="6" pageNumber="169">Fig. 5C</figureCitation>
|
||
).
|
||
<materialsCitation id="3B45DD484D58DE57FE2F4D2BFD14F511" ID-GBIF-Occurrence="3396390328" box="[494,664,1630,1649]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="6" pageNumber="169" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
shares a similar size, diameter, angle of lateral deflection toward the distal end, and the nearly identical flattened medial surfaces for articulation with metatarsal III with the LACM metatarsal IV (online Supplemental File fig. 1), as well as IGM 6130. Initially, we compared
|
||
<materialsCitation id="3B45DD484D58DE57FE154DA6FD0CF586" ID-GBIF-Occurrence="3396390324" box="[468,640,1747,1766]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="6" pageNumber="169" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
to photographs of bones of the
|
||
<taxonomicName id="4C2DAC964D58DE57FEC44D85FE3DF462" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[261,433,1776,1794]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="169" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D58DE57FEC44D85FE3DF462" box="[261,433,1776,1794]" italics="true" pageId="6" pageNumber="169">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
specimen ‘‘Sue’’ (FMNH PR2081) found in
|
||
<figureCitation id="1316CB904D58DE57FCA74E4CFBCCF62D" box="[870,1088,1337,1357]" captionStart="FIGURE 101" captionText="Brochu 2003: FIGURE 101. FMNH PR2081, Tyrannosaurus rex. Right fourth metatarsal (MTIV) in medial (A), lateral (B), dorsal (C), and ventral (D) view. Scale =10 cm. Photographs by J. Weinstein. " figureDoi="http://doi.org/10.5281/zenodo.3740185" httpUri="https://zenodo.org/record/3740185/files/figure.png" pageId="6" pageNumber="169" targetBox="[338,2156,288,1822]" targetPageId="119">
|
||
<bibRefCitation id="EFBCAAE44D58DE57FCA74E4CFC6AF62C" author="BROCHU, C. A." box="[870,998,1337,1357]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="6" pageNumber="169" pagination="1 - 138" part="22" refId="ref7504" refString="BROCHU, C. A., 2003, Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull: Journal of Vertebrate Paleontology, v. 22, p. 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu (2003</bibRefCitation>
|
||
, Fig. 101
|
||
</figureCitation>
|
||
). However, after observing bones from various ontogenetic stages of
|
||
<taxonomicName id="4C2DAC964D58DE57FBDA4E22FB2FF609" authorityName="Osborn" authorityYear="1905" box="[1051,1187,1367,1385]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="169" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D58DE57FBDA4E22FB2FF609" box="[1051,1187,1367,1385]" italics="true" pageId="6" pageNumber="169">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
at the Natural History Museum of Los Angeles County, we realized that
|
||
<taxonomicName id="4C2DAC964D58DE57FB614E01FAA4F6E6" authorityName="Osborn" authorityYear="1905" box="[1184,1320,1396,1414]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="169" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D58DE57FB614E01FAA4F6E6" box="[1184,1320,1396,1414]" italics="true" pageId="6" pageNumber="169">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
metatarsals change in morphology during ontogeny (
|
||
<figureCitation id="1316CB904D58DE57FB444EE5FB4AF6C4" box="[1157,1222,1424,1444]" captionStart="FIG. 5" captionStartId="6.[165,191,1174,1190]" captionTargetBox="[145,1514,176,1144]" captionTargetId="figure@6.[144,1514,176,1144]" captionTargetPageId="6" captionText="FIG. 5.—Comparison between four specimens of tyrannosaurid metatarsals: A) Tyrannosaurus rex metatarsal IV (SWAU HRS13997). B) Tyrannosaurus rex metatarsals II-IV (LACM 23845, with IV sculpted). C) Tyrannosauridae indet. metatarsal IV (IGM 6130; modified from Peecook (2014, fig. 3). D) Tyrannosaurus rex metatarsals II-IV (FMNH PR2081; modified from Brochu (2003, figs. 99, 100, 101). SWAU HRS13997 and IGM 6130 are metatarsal IV. All elements are from the right pes except IGM 6130, which is a left element, although the image has been flipped to make it match the others. Note that FMNH PR2081 is shown in a more medial view than the other metatarsal specimens. Scale bar = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.3749649" httpUri="https://zenodo.org/record/3749649/files/figure.png" pageId="6" pageNumber="169">Fig. 5</figureCitation>
|
||
). The large FMNH PR2081 metatarsals (
|
||
<figureCitation id="1316CB904D58DE57FC7E4ED8FB89F6A1" box="[959,1029,1453,1473]" captionStart="FIG. 5" captionStartId="6.[165,191,1174,1190]" captionTargetBox="[145,1514,176,1144]" captionTargetId="figure@6.[144,1514,176,1144]" captionTargetPageId="6" captionText="FIG. 5.—Comparison between four specimens of tyrannosaurid metatarsals: A) Tyrannosaurus rex metatarsal IV (SWAU HRS13997). B) Tyrannosaurus rex metatarsals II-IV (LACM 23845, with IV sculpted). C) Tyrannosauridae indet. metatarsal IV (IGM 6130; modified from Peecook (2014, fig. 3). D) Tyrannosaurus rex metatarsals II-IV (FMNH PR2081; modified from Brochu (2003, figs. 99, 100, 101). SWAU HRS13997 and IGM 6130 are metatarsal IV. All elements are from the right pes except IGM 6130, which is a left element, although the image has been flipped to make it match the others. Note that FMNH PR2081 is shown in a more medial view than the other metatarsal specimens. Scale bar = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.3749649" httpUri="https://zenodo.org/record/3749649/files/figure.png" pageId="6" pageNumber="169">Fig. 5D</figureCitation>
|
||
) are noticeably less slender, more rugose, and overall more robust than metatarsals from skeletally immature individuals (
|
||
<figureCitation id="1316CB904D58DE57FA0B4EBEFC1FF69C" captionStart="FIG. 5" captionStartId="6.[165,191,1174,1190]" captionTargetBox="[145,1514,176,1144]" captionTargetId="figure@6.[144,1514,176,1144]" captionTargetPageId="6" captionText="FIG. 5.—Comparison between four specimens of tyrannosaurid metatarsals: A) Tyrannosaurus rex metatarsal IV (SWAU HRS13997). B) Tyrannosaurus rex metatarsals II-IV (LACM 23845, with IV sculpted). C) Tyrannosauridae indet. metatarsal IV (IGM 6130; modified from Peecook (2014, fig. 3). D) Tyrannosaurus rex metatarsals II-IV (FMNH PR2081; modified from Brochu (2003, figs. 99, 100, 101). SWAU HRS13997 and IGM 6130 are metatarsal IV. All elements are from the right pes except IGM 6130, which is a left element, although the image has been flipped to make it match the others. Note that FMNH PR2081 is shown in a more medial view than the other metatarsal specimens. Scale bar = 100 mm." figureDoi="http://doi.org/10.5281/zenodo.3749649" httpUri="https://zenodo.org/record/3749649/files/figure.png" pageId="6" pageNumber="169">Fig. 5A–5C</figureCitation>
|
||
), which is consistent with the reasonable assumption that
|
||
<taxonomicName id="4C2DAC964D58DE57FC8C4D72FC59F579" authorityName="Osborn" authorityYear="1905" box="[845,981,1543,1561]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="169" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D58DE57FC8C4D72FC59F579" box="[845,981,1543,1561]" italics="true" pageId="6" pageNumber="169">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
feet would have become wider and more robust as the entire animal increased in mass, length, and height through ontogeny. Using an estimated length of 450 mm and the tyrannosaurid long bone regression equations of
|
||
<bibRefCitation id="EFBCAAE44D58DE57FBED4D28FB3BF511" author="CURRIE, P. J." box="[1068,1207,1629,1649]" journalOrPublisher="Canadian Journal of Earth Sciences" pageId="6" pageNumber="169" pagination="651 - 665" part="40" refId="ref8022" 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, v. 40, p. 651 - 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>
|
||
, we estimated the femoral length (813 mm) and femoral circumference (309 mm) for this individual. Then, using the
|
||
<bibRefCitation id="EFBCAAE44D58DE57FC704DEDFB02F5CC" author="ANDERSON, J. F. & HALL- MARTIN, A. & RUSSELL, D. A." box="[945,1166,1688,1708]" journalOrPublisher="Journal of Zoology" pageId="6" pageNumber="169" pagination="53 - 61" part="207" refId="ref7193" refString="ANDERSON, J. F., HALL- MARTIN, A., AND RUSSELL, D. A., 1985, Long bone circumference and weight in mammals, birds, and dinosaurs: Journal of Zoology, v. 207, p. 53 - 61." title="Long bone circumference and weight in mammals, birds, and dinosaurs" type="journal article" year="1985">Anderson et al. (1985)</bibRefCitation>
|
||
equation for estimating mass of an organism from femur circumference, we arrived at an estimated mass of 1,000 kg. Inserting this estimated mass into the
|
||
<bibRefCitation id="EFBCAAE44D58DE57FADF4DA7FA61F586" author="ERICKSON, G. M. & MAKOVICKY, P. J. & CURRIE, P. J. & NORELL, M. A. & YERBY, S. A. & BROCHU, C. A." box="[1310,1517,1746,1766]" journalOrPublisher="Nature" pageId="6" pageNumber="169" pagination="772 - 775" part="430" refId="ref8409" refString="ERICKSON, G. M., MAKOVICKY, P. J., CURRIE, P. J., NORELL, M. A., YERBY, S. A., AND BROCHU, C. A., 2004, Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs: Nature, v. 430, p. 772 - 775." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">Erickson et al. (2004)</bibRefCitation>
|
||
growth curve equation based on mass, we arrived at an estimated age of thirteen years old. Despite these admittedly tentative estimates, the results are similar to
|
||
<materialsCitation id="3B45DD484D59DE56FF314BB9FEFEF380" ID-GBIF-Occurrence="3396390341" box="[240,370,204,224]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/ContactSheet.php?QueryName=DetailedQuery&StartAt=1&QueryPage=%2Fdinosaur-institute%2Findex.php&Restriction=%28exists+%28SecDepartment_tab+where+%28SecDepartment+contains+%27Dinosaur+Institute%27%29%29%29&col_SpeSpecimenNoInteger=23845&LimitPerPage=20&Search=Search& " pageId="7" pageNumber="170" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
, which was estimated at fourteen years old by
|
||
<bibRefCitation id="EFBCAAE44D59DE56FFB34B9DFEB1F39C" author="ERICKSON, G. M. & MAKOVICKY, P. J. & CURRIE, P. J. & NORELL, M. A. & YERBY, S. A. & BROCHU, C. A." box="[114,317,232,252]" journalOrPublisher="Nature" pageId="7" pageNumber="170" pagination="772 - 775" part="430" refId="ref8409" refString="ERICKSON, G. M., MAKOVICKY, P. J., CURRIE, P. J., NORELL, M. A., YERBY, S. A., AND BROCHU, C. A., 2004, Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs: Nature, v. 430, p. 772 - 775." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">Erickson et al. (2004)</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
<caption id="DF52879D4D5ADE55FF4C4D01FAC8F5C3" ID-Table-UUID="DF52879D4D5ADE55FF4C4D01FAC8F5C3" httpUri="http://table.plazi.org/id/DF52879D4D5ADE55FF4C4D01FAC8F5C3" pageId="4" pageNumber="167" startId="4.[141,195,1652,1671]" subCaptionStartIDs="4.[938,1068,1651,1671]" subCaptionStarts="Abbr" targetBox="[141,1517,1751,1916]" targetIsTable="true" targetPageId="4">
|
||
<paragraph id="8B92D7154D5ADE55FF4C4D01FAC8F5C3" blockId="4.[141,1518,1651,1699]" pageId="4" pageNumber="167">
|
||
TABLE 1.—
|
||
<emphasis id="B9590B074D5ADE55FF314D06FB02F5E6" box="[240,1166,1651,1671]" italics="true" pageId="4" pageNumber="167">Denticle measurements for theropod taxa known from the Lance Formation. Abbreviations: DD/2 mm</emphasis>
|
||
=
|
||
<emphasis id="B9590B074D5ADE55FB664D06FEE6F5C2" italics="true" pageId="4" pageNumber="167">distal denticles per two millimeters; DDW</emphasis>
|
||
=
|
||
<emphasis id="B9590B074D5ADE55FE464DFAFD22F5C2" box="[391,686,1679,1699]" italics="true" pageId="4" pageNumber="167">distal denticle width; MD/2 mm</emphasis>
|
||
=
|
||
<emphasis id="B9590B074D5ADE55FD0B4DFAFBD2F5C2" box="[714,1118,1679,1699]" italics="true" pageId="4" pageNumber="167">mesial denticles per two millimeters; MDW</emphasis>
|
||
=
|
||
<emphasis id="B9590B074D5ADE55FBB84DFAFAB2F5C3" box="[1145,1342,1679,1699]" italics="true" pageId="4" pageNumber="167">mesial denticle width</emphasis>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8B92D7154D5ADE55FF4C4DADFB11F41C" pageId="4" pageNumber="167">
|
||
<table id="F92D25B54D5A21AEFF4C4DA2FA61F41C" box="[141,1517,1751,1916]" gridcols="7" gridrows="6" pageId="4" pageNumber="167">
|
||
<tr id="351DD5574D5A21AEFF4C4DA2FA61F589" box="[141,1517,1751,1769]" gridrow="0" pageId="4" pageNumber="167">
|
||
<th id="76CCBC2B4D5A21AEFF4C4DA2FE89F589" box="[141,261,1751,1769]" gridcol="0" gridrow="0" pageId="4" pageNumber="167">Taxon</th>
|
||
<th id="76CCBC2B4D5A21AEFE8E4DA2FE45F589" box="[335,457,1751,1769]" gridcol="1" gridrow="0" pageId="4" pageNumber="167"># of specimens</th>
|
||
<th id="76CCBC2B4D5A21AEFDD34DA2FD28F589" box="[530,676,1751,1769]" gridcol="2" gridrow="0" pageId="4" pageNumber="167">AVG MDW (mm)</th>
|
||
<th id="76CCBC2B4D5A21AEFD2C4DA2FCCEF589" box="[749,834,1751,1769]" gridcol="3" gridrow="0" pageId="4" pageNumber="167">MD/2 mm</th>
|
||
<th id="76CCBC2B4D5A21AEFC4A4DA2FB95F589" box="[907,1049,1751,1769]" gridcol="4" gridrow="0" pageId="4" pageNumber="167">AVG DDW (mm)</th>
|
||
<th id="76CCBC2B4D5A21AEFBA24DA2FB39F589" box="[1123,1205,1751,1769]" gridcol="5" gridrow="0" pageId="4" pageNumber="167">DD/2 mm</th>
|
||
<th id="76CCBC2B4D5A21AEFB3F4DA2FA61F589" box="[1278,1517,1751,1769]" gridcol="6" gridrow="0" pageId="4" pageNumber="167">Literature D/2 mm (MD/DD)</th>
|
||
</tr>
|
||
<tr id="351DD5574D5A21AEFF4C4C70FA61F477" box="[141,1517,1797,1815]" gridrow="1" pageId="4" pageNumber="167" rowspan-6="1">
|
||
<th id="76CCBC2B4D5A21AEFF4C4C70FE89F477" box="[141,261,1797,1815]" gridcol="0" gridrow="1" pageId="4" pageNumber="167">
|
||
<taxonomicName id="4C2DAC964D5ADE55FF4C4C70FF75F477" box="[141,249,1797,1815]" class="Reptilia" family="Dromaeosauridae" genus="Acheroraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="167" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D5ADE55FF4C4C70FF75F477" box="[141,249,1797,1815]" italics="true" pageId="4" pageNumber="167">Acheroraptor</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="76CCBC2B4D5A21AEFE8E4C70FE45F477" box="[335,457,1797,1815]" gridcol="1" gridrow="1" pageId="4" pageNumber="167">15</td>
|
||
<td id="76CCBC2B4D5A21AEFDD34C70FD28F477" box="[530,676,1797,1815]" gridcol="2" gridrow="1" pageId="4" pageNumber="167">0.327</td>
|
||
<td id="76CCBC2B4D5A21AEFD2C4C70FCCEF477" box="[749,834,1797,1815]" gridcol="3" gridrow="1" pageId="4" pageNumber="167">6.12</td>
|
||
<td id="76CCBC2B4D5A21AEFC4A4C70FB95F477" box="[907,1049,1797,1815]" gridcol="4" gridrow="1" pageId="4" pageNumber="167">0.334</td>
|
||
<td id="76CCBC2B4D5A21AEFBA24C70FB39F477" box="[1123,1205,1797,1815]" gridcol="5" gridrow="1" pageId="4" pageNumber="167">5.99</td>
|
||
</tr>
|
||
<tr id="351DD5574D5A21AEFF4C4C6BFA61F450" box="[141,1517,1822,1840]" gridrow="2" pageId="4" pageNumber="167">
|
||
<th id="76CCBC2B4D5A21AEFF4C4C6BFE89F450" box="[141,261,1822,1840]" gridcol="0" gridrow="2" pageId="4" pageNumber="167">
|
||
<taxonomicName id="4C2DAC964D5ADE55FF4C4C6BFF77F450" box="[141,251,1822,1840]" class="Reptilia" family="Dromaeosauridae" genus="Dakotaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="167" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D5ADE55FF4C4C6BFF77F450" box="[141,251,1822,1840]" italics="true" pageId="4" pageNumber="167">Dakotaraptor</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="76CCBC2B4D5A21AEFE8E4C6BFE45F450" box="[335,457,1822,1840]" gridcol="1" gridrow="2" pageId="4" pageNumber="167">-</td>
|
||
<td id="76CCBC2B4D5A21AEFDD34C6BFD28F450" box="[530,676,1822,1840]" gridcol="2" gridrow="2" pageId="4" pageNumber="167">-</td>
|
||
<td id="76CCBC2B4D5A21AEFD2C4C6BFCCEF450" box="[749,834,1822,1840]" gridcol="3" gridrow="2" pageId="4" pageNumber="167">-</td>
|
||
<td id="76CCBC2B4D5A21AEFC4A4C6BFB95F450" box="[907,1049,1822,1840]" gridcol="4" gridrow="2" pageId="4" pageNumber="167">-</td>
|
||
<td id="76CCBC2B4D5A21AEFBA24C6BFB39F450" box="[1123,1205,1822,1840]" gridcol="5" gridrow="2" pageId="4" pageNumber="167">-</td>
|
||
<td id="76CCBC2B4D5A21AEFB3F4C6BFA61F450" box="[1278,1517,1822,1840]" gridcol="6" gridrow="2" pageId="4" pageNumber="167">
|
||
9.2/6.8 (
|
||
<bibRefCitation id="EFBCAAE44D5ADE55FA804C6BFA64F450" author="DEPALMA, R. A. & BURNHAM, D. A. & MARTIN, L. D. & LARSON, P. L. & BAKKER, R. T." box="[1345,1512,1822,1840]" journalOrPublisher="Paleontological Contributions" pageId="4" pageNumber="167" pagination="1 - 16" part="14" refId="ref8343" refString="DEPALMA, R. A., BURNHAM, D. A., MARTIN, L. D., LARSON, P. L., AND BAKKER, R. T., 2015, The first giant raptor (Theropoda: Dromaeosauridae) from the Hell Creek Formation: Paleontological Contributions, v. 14, p. 1 - 16." title="The first giant raptor (Theropoda: Dromaeosauridae) from the Hell Creek Formation" type="journal article" year="2015">DePalma et al. 2015</bibRefCitation>
|
||
)
|
||
</td>
|
||
</tr>
|
||
<tr id="351DD5574D5A21AEFF4C4C4DFA61F42A" box="[141,1517,1848,1866]" gridrow="3" pageId="4" pageNumber="167" rowspan-6="1">
|
||
<th id="76CCBC2B4D5A21AEFF4C4C4DFE89F42A" box="[141,261,1848,1866]" gridcol="0" gridrow="3" pageId="4" pageNumber="167">
|
||
<taxonomicName id="4C2DAC964D5ADE55FF4C4C4CFF72F429" box="[141,254,1849,1865]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="167" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D5ADE55FF4C4C4CFF72F429" box="[141,254,1849,1865]" italics="true" pageId="4" pageNumber="167">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="76CCBC2B4D5A21AEFE8E4C4DFE45F42A" box="[335,457,1848,1866]" gridcol="1" gridrow="3" pageId="4" pageNumber="167">16</td>
|
||
<td id="76CCBC2B4D5A21AEFDD34C4DFD28F42A" box="[530,676,1848,1866]" gridcol="2" gridrow="3" pageId="4" pageNumber="167">0.333</td>
|
||
<td id="76CCBC2B4D5A21AEFD2C4C4DFCCEF42A" box="[749,834,1848,1866]" gridcol="3" gridrow="3" pageId="4" pageNumber="167">6.01</td>
|
||
<td id="76CCBC2B4D5A21AEFC4A4C4DFB95F42A" box="[907,1049,1848,1866]" gridcol="4" gridrow="3" pageId="4" pageNumber="167">0.394</td>
|
||
<td id="76CCBC2B4D5A21AEFBA24C4DFB39F42A" box="[1123,1205,1848,1866]" gridcol="5" gridrow="3" pageId="4" pageNumber="167">5.08</td>
|
||
</tr>
|
||
<tr id="351DD5574D5A21AEFF4C4C24FA61F403" box="[141,1517,1873,1891]" gridrow="4" pageId="4" pageNumber="167">
|
||
<th id="76CCBC2B4D5A21AEFF4C4C24FE89F403" box="[141,261,1873,1891]" gridcol="0" gridrow="4" pageId="4" pageNumber="167">
|
||
<taxonomicName id="4C2DAC964D5ADE55FF4C4C27FE89F402" authorityName="Osborn" authorityYear="1905" box="[141,261,1874,1890]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="167" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D5ADE55FF4C4C27FE89F402" box="[141,261,1874,1890]" italics="true" pageId="4" pageNumber="167">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="76CCBC2B4D5A21AEFE8E4C24FE45F403" box="[335,457,1873,1891]" gridcol="1" gridrow="4" pageId="4" pageNumber="167">14</td>
|
||
<td id="76CCBC2B4D5A21AEFDD34C24FD28F403" box="[530,676,1873,1891]" gridcol="2" gridrow="4" pageId="4" pageNumber="167">0.622</td>
|
||
<td id="76CCBC2B4D5A21AEFD2C4C24FCCEF403" box="[749,834,1873,1891]" gridcol="3" gridrow="4" pageId="4" pageNumber="167">3.22</td>
|
||
<td id="76CCBC2B4D5A21AEFC4A4C24FB95F403" box="[907,1049,1873,1891]" gridcol="4" gridrow="4" pageId="4" pageNumber="167">0.528</td>
|
||
<td id="76CCBC2B4D5A21AEFBA24C24FB39F403" box="[1123,1205,1873,1891]" gridcol="5" gridrow="4" pageId="4" pageNumber="167">3.79</td>
|
||
<td id="76CCBC2B4D5A21AEFB3F4C24FA61F403" box="[1278,1517,1873,1891]" gridcol="6" gridrow="4" pageId="4" pageNumber="167">
|
||
3.7/3.8 (
|
||
<bibRefCitation id="EFBCAAE44D5ADE55FA804C24FA43F403" author="SMITH, J. B. & VANN, D. R. & DODSON, P." box="[1345,1487,1873,1891]" journalOrPublisher="The Anatomical Record Part A" pageId="4" pageNumber="167" pagination="699 - 736" part="285" refId="ref10751" refString="SMITH, J. B., VANN, D. R., AND DODSON, P., 2005, Dental morphology and variation in theropod dinosaurs: implications for the taxonomic identification of isolated teeth: The Anatomical Record Part A, v. 285, p. 699 - 736." title="Dental morphology and variation in theropod dinosaurs: implications for the taxonomic identification of isolated teeth" type="journal article" year="2005">Smith et al. 2005</bibRefCitation>
|
||
)
|
||
</td>
|
||
</tr>
|
||
<tr id="351DD5574D5A21AEFF4C4C1FFA61F41C" box="[141,1517,1898,1916]" gridrow="5" pageId="4" pageNumber="167" rowspan-6="1">
|
||
<th id="76CCBC2B4D5A21AEFF4C4C1FFE89F41C" box="[141,261,1898,1916]" gridcol="0" gridrow="5" pageId="4" pageNumber="167">
|
||
<taxonomicName id="4C2DAC964D5ADE55FF4C4C1FFF5DF41C" authorityName="Leidy" authorityYear="1856" box="[141,209,1898,1916]" class="Reptilia" family="Troodontidae" genus="Troodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="167" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D5ADE55FF4C4C1FFF5DF41C" box="[141,209,1898,1916]" italics="true" pageId="4" pageNumber="167">Troodon</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="76CCBC2B4D5A21AEFE8E4C1FFE45F41C" box="[335,457,1898,1916]" gridcol="1" gridrow="5" pageId="4" pageNumber="167">15</td>
|
||
<td id="76CCBC2B4D5A21AEFDD34C1FFD28F41C" box="[530,676,1898,1916]" gridcol="2" gridrow="5" pageId="4" pageNumber="167">-</td>
|
||
<td id="76CCBC2B4D5A21AEFD2C4C1FFCCEF41C" box="[749,834,1898,1916]" gridcol="3" gridrow="5" pageId="4" pageNumber="167">-</td>
|
||
<td id="76CCBC2B4D5A21AEFC4A4C1FFB95F41C" box="[907,1049,1898,1916]" gridcol="4" gridrow="5" pageId="4" pageNumber="167">0.385</td>
|
||
<td id="76CCBC2B4D5A21AEFBA24C1FFB39F41C" box="[1123,1205,1898,1916]" gridcol="5" gridrow="5" pageId="4" pageNumber="167">5.19</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<caption id="DF52879D4D58DE57FF644FE3FE14F66B" ID-DOI="http://doi.org/10.5281/zenodo.3749649" ID-Zenodo-Dep="3749649" httpUri="https://zenodo.org/record/3749649/files/figure.png" pageId="6" pageNumber="169" startId="6.[165,191,1174,1190]" subCaptionStartIDs="6.[1141,1165,1198,1216] 6.[523,555,1223,1241]" subCaptionStarts="fig. 3 & figs. 99" targetBox="[145,1514,176,1144]" targetPageId="6">
|
||
<paragraph id="8B92D7154D58DE57FF644FE3FE14F66B" blockId="6.[141,1518,1173,1292]" pageId="6" pageNumber="169">
|
||
FIG. 5.—Comparison between four specimens of tyrannosaurid metatarsals:
|
||
<emphasis id="B9590B074D58DE57FCC74FE0FC98F7C7" bold="true" box="[774,788,1173,1191]" pageId="6" pageNumber="169">A</emphasis>
|
||
)
|
||
<taxonomicName id="4C2DAC964D58DE57FCE04FE3FC34F7C6" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[801,952,1174,1190]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="169" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D58DE57FCE04FE3FC34F7C6" box="[801,952,1174,1190]" italics="true" pageId="6" pageNumber="169">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
metatarsal IV (
|
||
<materialsCitation id="3B45DD484D58DE57FBF44FE0FB43F7C7" ID-GBIF-Occurrence="3396390313" box="[1077,1231,1173,1191]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="6" pageNumber="169" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
).
|
||
<emphasis id="B9590B074D58DE57FB1F4FE0FB67F7C7" bold="true" box="[1246,1259,1173,1191]" pageId="6" pageNumber="169">B</emphasis>
|
||
)
|
||
<taxonomicName id="4C2DAC964D58DE57FB364FE3FA03F7C6" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[1271,1423,1174,1190]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="169" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D58DE57FB364FE3FA03F7C6" box="[1271,1423,1174,1190]" italics="true" pageId="6" pageNumber="169">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
metatarsals II-IV (
|
||
<materialsCitation id="3B45DD484D58DE57FF014FDAFEB8F7A0" ID-GBIF-Occurrence="3396390320" box="[192,308,1198,1216]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/ContactSheet.php?QueryName=DetailedQuery&StartAt=1&QueryPage=%2Fdinosaur-institute%2Findex.php&Restriction=%28exists+%28SecDepartment_tab+where+%28SecDepartment+contains+%27Dinosaur+Institute%27%29%29%29&col_SpeSpecimenNoInteger=23845&LimitPerPage=20&Search=Search& " pageId="6" pageNumber="169" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
, with IV sculpted).
|
||
<emphasis id="B9590B074D58DE57FE134FDBFE6CF7A0" bold="true" box="[466,480,1198,1216]" pageId="6" pageNumber="169">C</emphasis>
|
||
)
|
||
<taxonomicName id="4C2DAC964D58DE57FE2D4FDBFDE2F7A0" authorityName="Osborn" authorityYear="1906" box="[492,622,1198,1216]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="169" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
indet. metatarsal IV (IGM 6130; modified from
|
||
<bibRefCitation id="EFBCAAE44D58DE57FC354FDBFBE1F7A0" author="PEECOOK, B. R. & WILSON, J. A. & HERNANDEZ- RIVERA, R. & MONTELLANO- BALLESTEROS, M. & WILSON, G. P." box="[1012,1133,1198,1216]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="6" pageNumber="169" pagination="71 - 80" part="59" refId="ref10144" refString="PEECOOK, B. R., WILSON, J. A., HERNANDEZ- RIVERA, R., MONTELLANO- BALLESTEROS, M., AND WILSON, G. P., 2014, First tyrannosaurid remains from the Upper Cretaceous '' El Gallo '' Formation of Baja California, Mexico: Acta Palaeontologica Polonica, v. 59, p. 71 - 80." title="First tyrannosaurid remains from the Upper Cretaceous '' El Gallo '' Formation of Baja California, Mexico" type="journal article" year="2014">Peecook (2014</bibRefCitation>
|
||
, fig. 3).
|
||
<emphasis id="B9590B074D58DE57FB714FDBFB32F7A0" bold="true" box="[1200,1214,1198,1216]" pageId="6" pageNumber="169">D</emphasis>
|
||
)
|
||
<taxonomicName id="4C2DAC964D58DE57FB0A4FDAFAEEF7DF" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[1227,1378,1199,1215]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="169" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D58DE57FB0A4FDAFAEEF7DF" box="[1227,1378,1199,1215]" italics="true" pageId="6" pageNumber="169">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
metatarsals II-IV (FMNH PR2081; modified from
|
||
<bibRefCitation id="EFBCAAE44D58DE57FE554FB2FD8FF7B9" author="BROCHU, C. A." box="[404,515,1223,1241]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="6" pageNumber="169" pagination="1 - 138" part="22" refId="ref7504" refString="BROCHU, C. A., 2003, Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull: Journal of Vertebrate Paleontology, v. 22, p. 1 - 138." title="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">Brochu (2003</bibRefCitation>
|
||
, figs. 99, 100, 101).
|
||
<materialsCitation id="3B45DD484D58DE57FD664FBDFCB7F7B9" ID-GBIF-Occurrence="3396390333" box="[679,827,1224,1241]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="6" pageNumber="169" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
and IGM 6130 are metatarsal IV. All elements are from the right pes except IGM 6130, which is a left element, although the image has been flipped to make it match the others. Note that FMNH PR2081 is shown in a more medial view than the other metatarsal specimens. Scale bar = 100 mm.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF52879D4D58DE57FF4C4C3CFA63F4EC" ID-DOI="http://doi.org/10.5281/zenodo.4023994" ID-Zenodo-Dep="4023994" httpUri="https://zenodo.org/record/4023994/files/figure.png" pageId="6" pageNumber="169" targetBox="[156,1437,176,1928]" targetPageId="5">
|
||
<paragraph id="8B92D7154D58DE57FF4C4C3CFA63F4EC" blockId="6.[141,1519,1840,1933]" pageId="6" pageNumber="169">
|
||
FIG. 4.—Histology of a
|
||
<taxonomicName id="4C2DAC964D58DE57FE894C3CFE53F439" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[328,479,1865,1881]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="169" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D58DE57FE894C3CFE53F439" box="[328,479,1865,1881]" italics="true" pageId="6" pageNumber="169">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
metatarsal (
|
||
<materialsCitation id="3B45DD484D58DE57FDFF4C3CFD59F43A" ID-GBIF-Occurrence="3396390343" box="[574,725,1864,1882]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="6" pageNumber="169" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
) in cross section under different light regimes.
|
||
<emphasis id="B9590B074D58DE57FB864C3DFBD9F43A" bold="true" box="[1095,1109,1864,1882]" pageId="6" pageNumber="169">A</emphasis>
|
||
) Section of cortical bone from the medullary cavity (left) to the subperiosteal surface (right) under regularly transmitted light. Boxes show where areas of this bone are shown in higher detail under regularly transmitted light (
|
||
<emphasis id="B9590B074D58DE57FA1A4C17FA64F414" bold="true" box="[1499,1512,1890,1908]" pageId="6" pageNumber="169">B</emphasis>
|
||
,
|
||
<emphasis id="B9590B074D58DE57FF4C4C0EFF15F4ED" bold="true" box="[141,153,1915,1933]" pageId="6" pageNumber="169">F</emphasis>
|
||
,
|
||
<emphasis id="B9590B074D58DE57FF644C0EFF23F4ED" bold="true" box="[165,175,1915,1933]" pageId="6" pageNumber="169">J</emphasis>
|
||
), cross-polarized light (
|
||
<emphasis id="B9590B074D58DE57FEAE4C0EFEF1F4ED" bold="true" box="[367,381,1915,1933]" pageId="6" pageNumber="169">C</emphasis>
|
||
,
|
||
<emphasis id="B9590B074D58DE57FE484C0EFE15F4ED" bold="true" box="[393,409,1915,1933]" pageId="6" pageNumber="169">G</emphasis>
|
||
,
|
||
<emphasis id="B9590B074D58DE57FE624C0EFE3FF4ED" bold="true" box="[419,435,1915,1933]" pageId="6" pageNumber="169">K</emphasis>
|
||
), cross-polarized light with a waveplate (
|
||
<emphasis id="B9590B074D58DE57FCC14C0EFC82F4ED" bold="true" box="[768,782,1915,1933]" pageId="6" pageNumber="169">D</emphasis>
|
||
,
|
||
<emphasis id="B9590B074D58DE57FCD84C0EFCA5F4ED" bold="true" box="[793,809,1915,1933]" pageId="6" pageNumber="169">H</emphasis>
|
||
,
|
||
<emphasis id="B9590B074D58DE57FCF54C0EFCCDF4ED" bold="true" box="[820,833,1915,1933]" pageId="6" pageNumber="169">L</emphasis>
|
||
), and reflected light (
|
||
<emphasis id="B9590B074D58DE57FC314C0EFC71F4ED" bold="true" box="[1008,1021,1915,1933]" pageId="6" pageNumber="169">E</emphasis>
|
||
,
|
||
<emphasis id="B9590B074D58DE57FBC94C0EFB9CF4ED" bold="true" box="[1032,1040,1915,1933]" pageId="6" pageNumber="169">I</emphasis>
|
||
,
|
||
<emphasis id="B9590B074D58DE57FBDA4C0EFBA2F4ED" bold="true" box="[1051,1070,1915,1933]" pageId="6" pageNumber="169">M</emphasis>
|
||
). Scale bar for A = 5 mm, all other scale bars = 1 mm.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8B92D7154D59DE56FF4B4A73FE7FF00F" blockId="7.[114,788,175,1934]" pageId="7" pageNumber="170">
|
||
We interpret the long grooves on
|
||
<materialsCitation id="3B45DD484D59DE56FE0E4A73FDF0F279" ID-GBIF-Occurrence="3396390325" box="[463,636,262,281]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="7" pageNumber="170" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
as tooth scores based on their greater than 3:1 length to width ratios and U-shaped cross sections (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FF074A4BFECFF232" author="BINFORD, L. R." box="[198,323,318,338]" journalOrPublisher="Academic Press, Inc., San Diego, CA" pageId="7" pageNumber="170" pagination="35 - 86" refId="ref7405" refString="BINFORD, L. R., 1981, Patterns of bone modifications produced by nonhuman agents, in Bones: Ancient Men and Modern Myths: Academic Press, Inc., San Diego, CA, p. 35 - 86." title="Patterns of bone modifications produced by nonhuman agents" type="book chapter" volumeTitle="Bones: Ancient Men and Modern Myths" year="1981">Binford 1981</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FE8D4A4BFDD6F232" author="NJAU, J. K. & BLUMENSCHINE, R. J." box="[332,602,318,338]" journalOrPublisher="Journal of Human Evolution" pageId="7" pageNumber="170" pagination="142 - 162" part="50" refId="ref9872" refString="NJAU, J. K. AND BLUMENSCHINE, R. J., 2006, A diagnosis of crocodile feeding traces on larger mammal bone, with fossil examples from the Plio-Pleistocene Olduvai Basin, Tanzania: Journal of Human Evolution, v. 50, p. 142 - 162." title="A diagnosis of crocodile feeding traces on larger mammal bone, with fossil examples from the Plio-Pleistocene Olduvai Basin, Tanzania" type="journal article" year="2006">Njau and Blumenschine 2006</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FDA24A4BFC83F232" author="POBINER, B. L. & DESILVA, J. & SANDERS, W. J. & MITANI, J. C." box="[611,783,318,338]" journalOrPublisher="Journal of Human Evolution" pageId="7" pageNumber="170" pagination="614 - 636" part="52" refId="ref10320" refString="POBINER, B. L., DESILVA, J., SANDERS, W. J., AND MITANI, J. C., 2007, Taphonomic analysis of skeletal remains from chimpanzee hunts at Ngogo, Kibale National Park, Uganda: Journal of Human Evolution, v. 52, p. 614 - 636." title="Taphonomic analysis of skeletal remains from chimpanzee hunts at Ngogo, Kibale National Park, Uganda" type="journal article" year="2007">Pobiner et al. 2007</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FFB34A2FFF7CF20E" author="POBINER, B." box="[114,240,346,366]" journalOrPublisher="Journal of Taphonomy" pageId="7" pageNumber="170" pagination="373 - 397" part="6" refId="ref10292" refString="POBINER, B., 2008, Paleoecological information in predator tooth marks: Journal of Taphonomy, v. 6, p. 373 - 397." title="Paleoecological information in predator tooth marks" type="journal article" year="2008">Pobiner 2008</bibRefCitation>
|
||
). The presence of both
|
||
<taxonomicName id="4C2DAC964D59DE56FE094A2FFDFFF20E" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[456,627,346,366]" class="Reptilia" genus="Linichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="170" phylum="Chordata" rank="species" species="serratus">
|
||
<emphasis id="B9590B074D59DE56FE094A2FFDFFF20E" box="[456,627,346,366]" italics="true" pageId="7" pageNumber="170">Linichnus serratus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C2DAC964D59DE56FD634A2FFF59F2EB" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="170" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D59DE56FD634A2FFF59F2EB" italics="true" pageId="7" pageNumber="170">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
confirm that at least some of these marks were made by serrated teeth. Even though the SW on the
|
||
<taxonomicName id="4C2DAC964D59DE56FDC24AE6FD53F2C7" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[515,735,403,423]" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="7" pageNumber="170" phylum="Chordata" rank="species" species="parallelum">
|
||
<emphasis id="B9590B074D59DE56FDC24AE6FD53F2C7" box="[515,735,403,423]" italics="true" pageId="7" pageNumber="170">Knethichnus parallelum</emphasis>
|
||
</taxonomicName>
|
||
most closely matches the DW of
|
||
<taxonomicName id="4C2DAC964D59DE56FEA44AC4FD9CF2A3" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[357,528,433,451]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D59DE56FEA44AC4FD9CF2A3" box="[357,528,433,451]" italics="true" pageId="7" pageNumber="170">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
teeth, it is possible that there is another carnivorous varanid or theropod with large denticles in the Lance Formation. Indeed, a large dromaeosaur was recently discovered in the Hell Creek Formation (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FEAF4970FDA3F179" author="DEPALMA, R. A. & BURNHAM, D. A. & MARTIN, L. D. & LARSON, P. L. & BAKKER, R. T." box="[366,559,517,537]" journalOrPublisher="Paleontological Contributions" pageId="7" pageNumber="170" pagination="1 - 16" part="14" refId="ref8343" refString="DEPALMA, R. A., BURNHAM, D. A., MARTIN, L. D., LARSON, P. L., AND BAKKER, R. T., 2015, The first giant raptor (Theropoda: Dromaeosauridae) from the Hell Creek Formation: Paleontological Contributions, v. 14, p. 1 - 16." title="The first giant raptor (Theropoda: Dromaeosauridae) from the Hell Creek Formation" type="journal article" year="2015">DePalma et al. 2015</bibRefCitation>
|
||
), although those authors concluded that
|
||
<taxonomicName id="4C2DAC964D59DE56FED44957FE1BF156" box="[277,407,546,566]" class="Reptilia" family="Dromaeosauridae" genus="Dakotaraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D59DE56FED44957FE1BF156" box="[277,407,546,566]" italics="true" pageId="7" pageNumber="170">Dakotaraptor</emphasis>
|
||
</taxonomicName>
|
||
had a higher denticle density than
|
||
<taxonomicName id="4C2DAC964D59DE56FFB34935FF76F132" authorityName="Osborn" authorityYear="1905" box="[114,250,576,594]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D59DE56FFB34935FF76F132" box="[114,250,576,594]" italics="true" pageId="7" pageNumber="170">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
, because of its smaller DW (
|
||
<tableCitation id="C6AFE2AE4D59DE56FDE3494BFDE7F132" box="[546,619,574,594]" captionStart="TABLE 1" captionStartId="4.[141,195,1652,1671]" captionTargetBox="[141,1517,1751,1916]" captionTargetPageId="4" captionText="TABLE 1.—Denticle measurements for theropod taxa known from the Lance Formation. Abbreviations: DD/2 mm = distal denticles per two millimeters; DDW = distal denticle width; MD/2 mm = mesial denticles per two millimeters; MDW = mesial denticle width." httpUri="http://table.plazi.org/id/DF52879D4D5ADE55FF4C4D01FAC8F5C3" pageId="7" pageNumber="170" tableUuid="DF52879D4D5ADE55FF4C4D01FAC8F5C3">Table 1</tableCitation>
|
||
). Although these marks may have conceivably been made by another unknown contemporaneous carnivorous taxon, the simplest explanation is that these tooth traces were left by
|
||
<taxonomicName id="4C2DAC964D59DE56FEDD49E0FE49F1C7" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[284,453,661,679]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D59DE56FEDD49E0FE49F1C7" box="[284,453,661,679]" italics="true" pageId="7" pageNumber="170">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
. Since 1965, over 42
|
||
<taxonomicName id="4C2DAC964D59DE56FD4B49E0FF03F1A4" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D59DE56FD4B49E0FF03F1A4" italics="true" pageId="7" pageNumber="170">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
skeletons (5% to 80% complete by bone count) have been collected from western North America (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FEBE49B8FE78F181" author="LARSON, N." box="[383,500,717,737]" editor="P. Larson & K. Carpenter" journalOrPublisher="Indiana University Press, Bloomington, Indiana" pageId="7" pageNumber="170" pagination="1 - 55" refId="ref9507" refString="LARSON, N., 2008, One hundred years of Tyrannosaurus rex: the skeletons, in P. Larson and K. Carpenter (eds.), Tyrannosaurus rex: The Tyrant King: Indiana University Press, Bloomington, Indiana, p. 1 - 55." title="One hundred years of Tyrannosaurus rex: the skeletons" type="book chapter" volumeTitle="Tyrannosaurus rex: The Tyrant King" year="2008">Larson 2008</bibRefCitation>
|
||
), and
|
||
<taxonomicName id="4C2DAC964D59DE56FDE849BBFD58F180" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[553,724,718,736]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D59DE56FDE849BBFD58F180" box="[553,724,718,736]" italics="true" pageId="7" pageNumber="170">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
fossils have been found from Alberta to New Mexico (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FD82499FFF2FF07A" author="SAMPSON, S. D. & LOEWEN, M. A." journalOrPublisher="Journal of Vertebrate Paleontology" pageId="7" pageNumber="170" pagination="469 - 472" part="25" refId="ref10533" refString="SAMPSON, S. D. AND LOEWEN, M. A., 2005, Tyrannosaurus rex from the Upper Cretaceous (Maastrichtian) North Horn Formation of Utah: biogeographic and paleoecologic implications: Journal of Vertebrate Paleontology, v. 25, p. 469 - 472." title="Tyrannosaurus rex from the Upper Cretaceous (Maastrichtian) North Horn Formation of Utah: biogeographic and paleoecologic implications" type="journal article" year="2005">Sampson and Loewen 2005</bibRefCitation>
|
||
). The plethora of
|
||
<taxonomicName id="4C2DAC964D59DE56FE804872FE45F079" authorityName="Osborn" authorityYear="1905" box="[321,457,775,793]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D59DE56FE804872FE45F079" box="[321,457,775,793]" italics="true" pageId="7" pageNumber="170">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
fossils that have been found, and the absence of any other large carnivores with appropriately-sized DWs, makes it likely that this trace made by a large theropod is from
|
||
<taxonomicName id="4C2DAC964D59DE56FDA54835FD61F032" authorityName="Osborn" authorityYear="1905" box="[612,749,832,850]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D59DE56FDA54835FD61F032" box="[612,749,832,850]" italics="true" pageId="7" pageNumber="170">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
and not some other large, undiscovered taxon.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D59DE56FF4B480CFB3BF3BF" blockId="7.[114,788,175,1934]" lastBlockId="7.[818,1492,175,1934]" pageId="7" pageNumber="170">
|
||
Because
|
||
<materialsCitation id="3B45DD484D59DE56FF1C480CFE04F0EC" ID-GBIF-Occurrence="3396390335" box="[221,392,889,908]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="7" pageNumber="170" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
is from a small Maastrichtian tyrannosaur from western North America, this element could possibly be assigned to the contentious taxon
|
||
<taxonomicName id="4C2DAC964D59DE56FE9048C7FC9CF0A5" authority="(Bakker et al. 1988)" baseAuthorityName="Bakker" baseAuthorityYear="1988" box="[337,784,945,965]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="species" species="lancensis">
|
||
<emphasis id="B9590B074D59DE56FE9048C7FDBFF0A5" box="[337,563,945,965]" italics="true" pageId="7" pageNumber="170">Nanotyrannus lancensis</emphasis>
|
||
(
|
||
<bibRefCitation id="EFBCAAE44D59DE56FD8648C4FC85F0A5" author="BAKKER, R. T. & WILLIAMS, M. & CURRIE, P. J." box="[583,777,945,965]" journalOrPublisher="Hunteria" pageId="7" pageNumber="170" pagination="1 - 30" part="1" refId="ref7243" refString="BAKKER, R. T., WILLIAMS, M., AND CURRIE, P. J., 1988, Nanotyrannus, a new genus of pygmy tyrannosaur, from the Latest Cretaceous of Montana: Hunteria, v. 1, p. 1 - 30." title="Nanotyrannus, a new genus of pygmy tyrannosaur, from the Latest Cretaceous of Montana" type="journal article" year="1988">Bakker et al. 1988</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
. Although some researchers consider
|
||
<taxonomicName id="4C2DAC964D59DE56FE0048BAFD10F081" box="[449,668,973,993]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="species" species="lancensis">
|
||
<emphasis id="B9590B074D59DE56FE0048BAFD10F081" box="[449,668,973,993]" italics="true" pageId="7" pageNumber="170">Nanotyrannus lancensis</emphasis>
|
||
</taxonomicName>
|
||
to be a valid species (e.g.,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FF2F489FFEE5F09E" author="CURRIE, P. J." box="[238,361,1002,1022]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="7" pageNumber="170" pagination="191 - 226" part="48" refId="ref8067" refString="CURRIE, P. J., 2003 b, Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada: Acta Palaeontologica Polonica, v. 48, p. 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">Currie 2003b</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FEB2489EFE64F09E" author="LARSON, P." box="[371,488,1002,1022]" journalOrPublisher="Society of Vertebrate Paleontology Abstracts with Programs, Los Angeles, CA" pageId="7" pageNumber="170" pagination="159" refId="ref9558" refString="LARSON, P., 2013, The validity of Nanotyrannus lancensis (Theropoda, Lancian-Upper Maastrichtian of North America): Society of Vertebrate Paleontology Abstracts with Programs, Los Angeles, CA, p. 159." title="The validity of Nanotyrannus lancensis (Theropoda, Lancian-Upper Maastrichtian of North America)" type="proceedings paper" year="2013">Larson 2013</bibRefCitation>
|
||
), other paleontologists view it as a skeletally immature
|
||
<taxonomicName id="4C2DAC964D59DE56FE854F7DFE78F77A" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[324,500,1032,1050]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D59DE56FE854F7DFE78F77A" box="[324,500,1032,1050]" italics="true" pageId="7" pageNumber="170">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
(e.g.,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FDF44F72FD1BF77A" author="CARR, T. D." box="[565,663,1031,1050]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="7" pageNumber="170" pagination="497 - 520" part="19" refId="ref7641" refString="CARR, T. D., 1999, Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria): Journal of Vertebrate Paleontology, v. 19, p. 497 - 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">Carr 1999</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FD654F73FC83F77A" author="HOLTZ, T. R." box="[676,783,1030,1050]" editor="D. H. Tanke & K. Carpenter" journalOrPublisher="Indiana University Press, Bloomington, Indiana" pageId="7" pageNumber="170" pagination="64 - 83" refId="ref8828" refString="HOLTZ, T. R., 2001, The phylogeny and taxonomy of the Tyrannosauridae, in D. H. Tanke and K. Carpenter (eds.), Mesozoic Vertebrate Life: Indiana University Press, Bloomington, Indiana, p. 64 - 83." title="The phylogeny and taxonomy of the Tyrannosauridae" type="book chapter" volumeTitle="Mesozoic Vertebrate Life" year="2001">Holtz 2001</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FFB34F51FF2EF757" author="HOLTZ, T. R." box="[114,162,1060,1079]" editor="D. B. Weishampel & P. Dodson & H. Osmolska" journalOrPublisher="University of California Press, Berkeley, CA" pageId="7" pageNumber="170" pagination="111 - 136" refId="ref8878" refString="HOLTZ, T. R., 2004, Tyrannosauroidea, in D. B. Weishampel, P. Dodson, and H. Osmolska (eds.), The Dinosauria: University of California Press, Berkeley, CA, p. 111 - 136." title="Tyrannosauroidea" type="book chapter" volumeTitle="The Dinosauria" year="2004">2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FF6D4F51FE11F757" author="CARR, T. D. & WILLIAMSON, T. E." box="[172,413,1059,1079]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="7" pageNumber="170" pagination="479 - 523" part="142" refId="ref7675" refString="CARR, T. D. AND WILLIAMSON, T. E., 2004, Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America: Zoological Journal of the Linnean Society, v. 142, p. 479 - 523." title="Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America" type="book chapter" year="2004">Carr and Williamson 2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FE674F51FDD5F757" author="BRUSATTE, S. L. & NORELL, M. A. & CARR, T. D. & ERICKSON, G. M. & HUTCHINSON, J. R. & BALANOFF, A. M. & BEVER, G. S. & COINIERE, J. N. & MAKOVICKY, P. J. & XU, X." box="[422,601,1059,1079]" journalOrPublisher="Science" pageId="7" pageNumber="170" pagination="1481 - 1485" part="329" refId="ref7548" refString="BRUSATTE, S. L., NORELL, M. A., CARR, T. D., ERICKSON, G. M., HUTCHINSON, J. R., BALANOFF, A. M., BEVER, G. S., COINIERE, J. N., MAKOVICKY, P. J., AND XU, X., 2010, Tyrannosaur paleobiology: new research on ancient exemplar organisms: Science, v. 329, p. 1481 - 1485." title="Tyrannosaur paleobiology: new research on ancient exemplar organisms" type="journal article" year="2010">Brusatte et al. 2010</bibRefCitation>
|
||
). The osteohistology of
|
||
<materialsCitation id="3B45DD484D59DE56FF4E4F35FEB7F733" ID-GBIF-Occurrence="3396390330" box="[143,315,1088,1107]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="7" pageNumber="170" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
possesses features that are indicative of a young, skeletally immature individual that is still undergoing rapid growth. The high level of vascularization extending to the subperiosteal surface on the outer portion of the histological section suggests that the growth had not ceased in this individual because high vascular density and connectivity is strongly correlated with growth rate (e.g.,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FE344FBAFD2FF782" author="MONTES, L. & CASTANET, J. & CUBO, J." box="[501,675,1230,1250]" journalOrPublisher="Animal Biology" pageId="7" pageNumber="170" pagination="25 - 41" part="60" refId="ref9765" refString="MONTES, L., CASTANET, J., AND CUBO, J., 2010, Relationship between bone growth rate and bone tissue organization in amniotes: first test of Amprino's rule in a phylogenetic context: Animal Biology, v. 60, p. 25 - 41." title="Relationship between bone growth rate and bone tissue organization in amniotes: first test of Amprino's rule in a phylogenetic context" type="journal article" year="2010">Montes et al. 2010</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FD6F4FBBFF2EF79E" author="CUBO, J. & LE ROY, N. & MARTINEZ- MAZA, C. & MONTES, L." journalOrPublisher="Paleobiology" pageId="7" pageNumber="170" pagination="335 - 349" part="38" refId="ref7975" refString="CUBO, J., LE ROY, N., MARTINEZ- MAZA, C., AND MONTES, L., 2012, Paleohistological estimation of bone growth rate in extinct archosaurs: Paleobiology, v. 38, p. 335 - 349." title="Paleohistological estimation of bone growth rate in extinct archosaurs" type="book chapter" year="2012">Cubo et al. 2012</bibRefCitation>
|
||
;
|
||
<figureCitation id="1316CB904D59DE56FF734F9FFEDEF79E" box="[178,338,1258,1278]" captionStart="FIG. 4" captionStartId="6.[141,167,1865,1881]" captionTargetId="figure@5.[148,1457,176,1930]" captionTargetPageId="5" captionText="FIG. 4.—Histology of a Tyrannosaurus rex metatarsal (SWAU HRS13997) in cross section under different light regimes. A) Section of cortical bone from the medullary cavity (left) to the subperiosteal surface (right) under regularly transmitted light. Boxes show where areas of this bone are shown in higher detail under regularly transmitted light (B, F, J), cross-polarized light (C, G, K), cross-polarized light with a waveplate (D, H, L), and reflected light (E, I, M). Scale bar for A = 5 mm, all other scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.4023994" httpUri="https://zenodo.org/record/4023994/files/figure.png" pageId="7" pageNumber="170">Fig. 4A, 4J–4M</figureCitation>
|
||
). Further, this vascular structure suggests that growth had not even begun to slow as is expected at the onset of skeletal maturity in tyrannosaurids (e.g.,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FE664E56FDE7F657" author="ERICKSON, G. M. & MAKOVICKY, P. J. & CURRIE, P. J. & NORELL, M. A. & YERBY, S. A. & BROCHU, C. A." box="[423,619,1315,1335]" journalOrPublisher="Nature" pageId="7" pageNumber="170" pagination="772 - 775" part="430" refId="ref8409" refString="ERICKSON, G. M., MAKOVICKY, P. J., CURRIE, P. J., NORELL, M. A., YERBY, S. A., AND BROCHU, C. A., 2004, Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs: Nature, v. 430, p. 772 - 775." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">Erickson et al. 2004</bibRefCitation>
|
||
). Vascular pattern may be somewhat related to relative growth rate (e.g.,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FDAB4E35FD7EF634" author="AMPRINO, R." box="[618,754,1344,1364]" journalOrPublisher="Archives de Biologie" pageId="7" pageNumber="170" pagination="317 - 330" part="58" refId="ref7156" refString="AMPRINO, R., 1947, La structure du tissu osseux envisagee comme expression de differences dans la vitesse de l'accroissement: Archives de Biologie, v. 58, p. 317 - 330." title="La structure du tissu osseux envisagee comme expression de differences dans la vitesse de l'accroissement" type="book chapter" year="1947">Amprino 1947</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FD3C4E35FF67F610" author="DE RICQLES, A." journalOrPublisher="Annales de Paleontologie" pageId="7" pageNumber="170" pagination="51 - 129" part="61" refId="ref8291" refString="DE RICQLES, A., 1975, Recherches paleohistologiques sur les os longs des tetrapodes VII, Sur la classification, la signification fonctionnelle et l'histoire des tissus oseux des tetrapodes, premiere partie, structures: Annales de Paleontologie, v. 61, p. 51 - 129." title="Recherches paleohistologiques sur les os longs des tetrapodes VII, Sur la classification, la signification fonctionnelle et l'histoire des tissus oseux des tetrapodes, premiere partie, structures" type="journal article" year="1975">de Ricqlès 1975</bibRefCitation>
|
||
; Francillon-Vieillot et al. 1990;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FDC84E28FD37F610" author="CASTANET, J. & GRANDIN, A. & ABOURACHID, A. & DE RICQLES, A." box="[521,699,1372,1392]" journalOrPublisher="Comptes Rendus de l'Academie des Sciences de Paris, Serie III" pageId="7" pageNumber="170" pagination="301 - 308" part="319" refId="ref7723" refString="CASTANET, J., GRANDIN, A., ABOURACHID, A., AND DE RICQLES, A., 1996, Expression de la dynamique de croissance dans la structure de l'os periostique chez A nas platyrhynchos: Comptes Rendus de l'Academie des Sciences de Paris, Serie III, v. 319, p. 301 - 308." title="Expression de la dynamique de croissance dans la structure de l'os periostique chez A nas platyrhynchos" type="journal article" year="1996">Castanet et al. 1996</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FD044E28FD79F610" author="CASTANET, J. & ROGERS, K. C. & CUBO, J. & JACQUES- BOISARD, J." box="[709,757,1373,1392]" journalOrPublisher="Comptes Rendus de l'Academie des Sciences, Series III, Sciences de la Vie" pageId="7" pageNumber="170" pagination="543 - 550" part="323" refId="ref7786" refString="CASTANET, J., ROGERS, K. C., CUBO, J., AND JACQUES- BOISARD, J., 2000, Periosteal bone growth rates in extant ratites (ostrich and emu). Implications for assessing growth in dinosaurs: Comptes Rendus de l'Academie des Sciences, Series III, Sciences de la Vie, v. 323, p. 543 - 550." title="Periosteal bone growth rates in extant ratites (ostrich and emu). Implications for assessing growth in dinosaurs" type="journal article" year="2000">2000</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FD3C4E29FEA6F6ED" author="DE MARGERIE, E. & CUBO, J. & CASTANET, J." journalOrPublisher="Comptes Rendus Biologies" pageId="7" pageNumber="170" pagination="221 - 230" part="325" refId="ref8161" refString="DE MARGERIE, E., CUBO, J., AND CASTANET, J., 2002, Bone typology and growth rate: testing and quantifying ' Amprino's rule' in the mallard (Anas platyrhynchos): Comptes Rendus Biologies, v. 325, p. 221 - 230." title="Bone typology and growth rate: testing and quantifying ' Amprino's rule' in the mallard (Anas platyrhynchos)" type="journal article" year="2002">de Margerie et al. 2002</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FEF54E0CFEE8F6ED" author="DE MARGERIE, E. & ROBIN, J. - P. & VERRIER, D. & CUBO, J. & GROSCOLAS, R. & CASTANET, J." box="[308,356,1401,1421]" journalOrPublisher="The Journal of Experimental Biology" pageId="7" pageNumber="170" pagination="869 - 879" part="207" refId="ref8214" refString="DE MARGERIE, E., ROBIN, J. - P., VERRIER, D., CUBO, J., GROSCOLAS, R., AND CASTANET, J., 2004, Assessing a relationship between bone microstructure and growth rate: a fluorescent labelling study in the king penguin chick (Aptenodytes patagonicus): The Journal of Experimental Biology, v. 207, p. 869 - 879." title="Assessing a relationship between bone microstructure and growth rate: a fluorescent labelling study in the king penguin chick (Aptenodytes patagonicus)" type="journal article" year="2004">2004</bibRefCitation>
|
||
), but this relationship may be taxon- or speciesspecific, and the view of a simple relationship between growth and vascular pattern has recently been challenged (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FE334EC7FC83F6A6" author="SIMONS, E. L. R. & O'CONNOR, P. M." box="[498,783,1458,1478]" journalOrPublisher="The Anatomical Record" pageId="7" pageNumber="170" pagination="386 - 396" part="295" refId="ref10661" refString="SIMONS, E. L. R. AND O'CONNOR, P. M., 2012, Bone laminarity in the avian forelimb skeleton and its relationship to flight mode: testing functional interpretations: The Anatomical Record, v. 295, p. 386 - 396." title="Bone laminarity in the avian forelimb skeleton and its relationship to flight mode: testing functional interpretations" type="journal article" year="2012">Simons and O’Connor 2012</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FFB34EBBFF77F682" author="WERNING, S. A." bookContentInfo="445 p." box="[114,251,1486,1506]" journalOrPublisher="University of California, Berkeley" pageId="7" pageNumber="170" refId="ref10961" refString="WERNING, S. A., 2013, Evolution of bone histological characters in amniotes, and the implications for the evolution of growth and metabolism: Unpublished Ph. D. dissertation, University of California, Berkeley, 445 p." title="Evolution of bone histological characters in amniotes, and the implications for the evolution of growth and metabolism" type="book" year="2013">Werning 2013</bibRefCitation>
|
||
), leaving the relationship between vascular pattern and growth rate unclear. Regardless, the predominant vascular patterns (i.e., longitudinal and circumferential) are consistent with rapid relative growth in those studies that find a correlation between vascular pattern and relative growth rate (e.g.,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FED64D34FE5EF534" author="CASTANET, J. & ROGERS, K. C. & CUBO, J. & JACQUES- BOISARD, J." box="[279,466,1600,1620]" journalOrPublisher="Comptes Rendus de l'Academie des Sciences, Series III, Sciences de la Vie" pageId="7" pageNumber="170" pagination="543 - 550" part="323" refId="ref7786" refString="CASTANET, J., ROGERS, K. C., CUBO, J., AND JACQUES- BOISARD, J., 2000, Periosteal bone growth rates in extant ratites (ostrich and emu). Implications for assessing growth in dinosaurs: Comptes Rendus de l'Academie des Sciences, Series III, Sciences de la Vie, v. 323, p. 543 - 550." title="Periosteal bone growth rates in extant ratites (ostrich and emu). Implications for assessing growth in dinosaurs" type="journal article" year="2000">Castanet et al. 2000</bibRefCitation>
|
||
). Although the precise relationship between Haversian remodeling and maturity varies by clade, the number of secondary osteons present in the cortex of a long bone increases with ontogenetic age (e.g.,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FE814DE2FE62F5CA" author="HORNER, J. R. & DE RICQLES, A. & PADIAN, K." box="[320,494,1686,1706]" journalOrPublisher="Paleobiology" pageId="7" pageNumber="170" pagination="295 - 304" part="25" refId="ref9090" refString="HORNER, J. R., DE RICQLES, A., AND PADIAN, K., 1999, Variation in dinosaur skeletochronology indicators: implications for age assessment and physiology: Paleobiology, v. 25, p. 295 - 304." title="Variation in dinosaur skeletochronology indicators: implications for age assessment and physiology" type="journal article" year="1999">Horner et al. 1999</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FE3B4DE2FDA6F5CA" author="HORNER, J. R. & DE RICQLES, A. & PADIAN, K." box="[506,554,1687,1706]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="7" pageNumber="170" pagination="115 - 129" part="20" refId="ref9136" refString="HORNER, J. R., DE RICQLES, A., AND PADIAN, K., 2000, Long bone histology of the hadrosaurid dinosaur Maiasaura peeblesorum: growth dynamics and physiology based on an ontogenetic series of skeletal elements: Journal of Vertebrate Paleontology, v. 20, p. 115 - 129." title="Long bone histology of the hadrosaurid dinosaur Maiasaura peeblesorum: growth dynamics and physiology based on an ontogenetic series of skeletal elements" type="journal article" year="2000">2000</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FDF74DE3FC83F5CA" author="KLEIN, N. & SANDER, M." box="[566,783,1686,1706]" journalOrPublisher="Special Papers in Palaeontology" pageId="7" pageNumber="170" pagination="169 - 206" part="77" refId="ref9391" refString="KLEIN, N. AND SANDER, M., 2007, Bone histology and growth of the prosauropod dinosaur Plateosaurus engelhardti Meyer, 1837 from the Norian bonebeds of Trossingen (Germany) and Frick (Switzerland): Special Papers in Palaeontology, v. 77, p. 169 - 206." title="Bone histology and growth of the prosauropod dinosaur Plateosaurus engelhardti Meyer, 1837 from the Norian bonebeds of Trossingen (Germany) and Frick (Switzerland)" type="journal article" year="2007">Klein and Sander 2007</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FFB34DC7FF7AF5A6" author="WERNING, S. A." box="[114,246,1714,1734]" journalOrPublisher="PLoS ONE" pageId="7" pageNumber="170" pagination="e 33539" part="7" refId="ref10923" refString="WERNING, S. A., 2012, The ontogenetic osteohistology of Tenontosaurus tilletti: PLoS ONE, v. 7, p. e 33539, doi: 10.1371 / journal. pone. 0033539." title="The ontogenetic osteohistology of Tenontosaurus tilletti" type="journal article" year="2012">Werning 2012</bibRefCitation>
|
||
), and extensive Haversian remodeling is present in the ribs of the largest known
|
||
<taxonomicName id="4C2DAC964D59DE56FEF04DA5FE50F582" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[305,476,1744,1762]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D59DE56FEF04DA5FE50F582" box="[305,476,1744,1762]" italics="true" pageId="7" pageNumber="170">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
(FMNH PR 2081;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FD4D4DBAFF2FF59F" author="ERICKSON, G. M. & MAKOVICKY, P. J. & CURRIE, P. J. & NORELL, M. A. & YERBY, S. A. & BROCHU, C. A." journalOrPublisher="Nature" pageId="7" pageNumber="170" pagination="772 - 775" part="430" refId="ref8409" refString="ERICKSON, G. M., MAKOVICKY, P. J., CURRIE, P. J., NORELL, M. A., YERBY, S. A., AND BROCHU, C. A., 2004, Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs: Nature, v. 430, p. 772 - 775." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">Erickson et al., 2004</bibRefCitation>
|
||
). However,
|
||
<materialsCitation id="3B45DD484D59DE56FED14D99FE36F59F" ID-GBIF-Occurrence="3396390351" box="[272,442,1772,1791]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="7" pageNumber="170" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
possesses no secondary osteons (
|
||
<figureCitation id="1316CB904D59DE56FD2E4D9EFFF3F47C" captionStart="FIG. 4" captionStartId="6.[141,167,1865,1881]" captionTargetId="figure@5.[148,1457,176,1930]" captionTargetPageId="5" captionText="FIG. 4.—Histology of a Tyrannosaurus rex metatarsal (SWAU HRS13997) in cross section under different light regimes. A) Section of cortical bone from the medullary cavity (left) to the subperiosteal surface (right) under regularly transmitted light. Boxes show where areas of this bone are shown in higher detail under regularly transmitted light (B, F, J), cross-polarized light (C, G, K), cross-polarized light with a waveplate (D, H, L), and reflected light (E, I, M). Scale bar for A = 5 mm, all other scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.4023994" httpUri="https://zenodo.org/record/4023994/files/figure.png" pageId="7" pageNumber="170">Fig. 4</figureCitation>
|
||
), even though as a smaller, weight-bearing element it is more likely to experience secondary remodeling, and to experience it earlier in ontogeny (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FFB94C34FE92F435" author="PADIAN, K. & WERNING, S. A. & HORNER, J. R." box="[120,286,1857,1877]" journalOrPublisher="Comptes Rendus Palevol" pageId="7" pageNumber="170" pagination="40 - 48" part="15" refId="ref10065" refString="PADIAN, K., WERNING, S. A., AND HORNER, J. R., 2016, A hypothesis of differential secondary bone formation in dinosaurs: Comptes Rendus Palevol, v. 15, p. 40 - 48." title="A hypothesis of differential secondary bone formation in dinosaurs" type="journal article" year="2016">Padian et al. 2016</bibRefCitation>
|
||
). Therefore, the complete lack of secondary remodeling also supports
|
||
<materialsCitation id="3B45DD484D59DE56FF2F4C2BFE1AF411" ID-GBIF-Occurrence="3396390327" box="[238,406,1886,1905]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="7" pageNumber="170" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
as a rapidly growing, immature individual that has not approached full size. We are confident that this element does not belong to an individual of
|
||
<taxonomicName id="4C2DAC964D59DE56FBAB4BC5FB66F3A2" box="[1130,1258,176,194]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D59DE56FBAB4BC5FB66F3A2" box="[1130,1258,176,194]" italics="true" pageId="7" pageNumber="170">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
, but is most likely the metatarsal of an immature
|
||
<taxonomicName id="4C2DAC964D59DE56FBE84BB9FB3DF3BE" authorityName="Osborn" authorityYear="1905" box="[1065,1201,204,222]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D59DE56FBE84BB9FB3DF3BE" box="[1065,1201,204,222]" italics="true" pageId="7" pageNumber="170">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D59DE56FC8B4B92FCE2F1E4" blockId="7.[818,1492,175,1934]" pageId="7" pageNumber="170">
|
||
Tyrannosaurid tooth scores, like those on
|
||
<materialsCitation id="3B45DD484D59DE56FB054B9DFAE2F39B" ID-GBIF-Occurrence="3396390331" box="[1220,1390,232,251]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="7" pageNumber="170" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
, have been found on other fossils (e.g.,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FBEC4A76FB3EF277" author="JACOBSEN, A. R." box="[1069,1202,259,279]" journalOrPublisher="Historical Biology" pageId="7" pageNumber="170" pagination="17 - 26" part="13" refId="ref9262" refString="JACOBSEN, A. R., 1998, Feeding behavior of carnivorous dinosaurs as determined by tooth marks on dinosaur bones: Historical Biology, v. 13, p. 17 - 26." title="Feeding behavior of carnivorous dinosaurs as determined by tooth marks on dinosaur bones" type="journal article" year="1998">Jacobsen 1998</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FB7D4A71FB60F277" author="JACOBSEN, A. R." box="[1212,1260,260,279]" editor="D. H. Tanke & K. Carpenter" journalOrPublisher="Indiana University Press, Bloomington, IN" pageId="7" pageNumber="170" pagination="59 - 63" refId="ref9298" refString="JACOBSEN, A. R., 2001, Tooth-marked small theropod bone: an extremely rare trace, in D. H. Tanke and K. Carpenter (eds.), Mesozoic Vertebrate Life: Indiana University Press, Bloomington, IN, p. 59 - 63." title="Tooth-marked small theropod bone: an extremely rare trace" type="book chapter" volumeTitle="Mesozoic Vertebrate Life" year="2001">2001</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FB354A71FA45F277" author="HONE, D. W. E. & WATABE, M." box="[1268,1481,259,279]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="7" pageNumber="170" pagination="627 - 634" part="55" refId="ref9049" refString="HONE, D. W. E. AND WATABE, M., 2010, New information on scavenging and selective feeding behavior of tyrannosaurids: Acta Palaeontologica Polonica, v. 55, p. 627 - 634." title="New information on scavenging and selective feeding behavior of tyrannosaurids" type="journal article" year="2010">Hone and Watabe 2010</bibRefCitation>
|
||
), as have punctures and furrows (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FBA84A6AFAEDF253" author="ERICKSON, G. M. & OLSON, K. H." box="[1129,1377,287,307]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="7" pageNumber="170" pagination="175 - 178" part="16" refId="ref8475" refString="ERICKSON, G. M. AND OLSON, K. H., 1996, Bite marks attributable to Tyrannosaurus rex: preliminary description and implications: Journal of Vertebrate Paleontology, v. 16, p. 175 - 178." title="Bite marks attributable to Tyrannosaurus rex: preliminary description and implications" type="journal article" year="1996">Erickson and Olson 1996</bibRefCitation>
|
||
). Given the robust jaws and teeth of tyrannosaurids, one might expect tyrannosaurid tooth traces on bones to be deeper than seen on
|
||
<materialsCitation id="3B45DD484D59DE56FB1F4A2DFA05F20B" ID-GBIF-Occurrence="3396390309" box="[1246,1417,344,363]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="7" pageNumber="170" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
. Indeed, it has been demonstrated, both through functional morphology studies and direct fossil evidence, that tyrannosaurids could bite deeply into bones (e.g.,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FCA94AD9FBD8F2A0" author="ERICKSON, G. M. & OLSON, K. H." box="[872,1108,428,448]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="7" pageNumber="170" pagination="175 - 178" part="16" refId="ref8475" refString="ERICKSON, G. M. AND OLSON, K. H., 1996, Bite marks attributable to Tyrannosaurus rex: preliminary description and implications: Journal of Vertebrate Paleontology, v. 16, p. 175 - 178." title="Bite marks attributable to Tyrannosaurus rex: preliminary description and implications" type="journal article" year="1996">Erickson and Olson 1996</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FB9E4AD9FA90F2A0" author="ERICKSON, G. M. & VAN KIRK, S. D. & SU, J. & LEVENSTON, M. E. & CALER, W. E. & CARTER, D. R." box="[1119,1308,428,448]" journalOrPublisher="Nature" pageId="7" pageNumber="170" pagination="706 - 708" part="382" refId="ref8518" refString="ERICKSON, G. M., VAN KIRK, S. D., SU, J., LEVENSTON, M. E., CALER, W. E., AND CARTER, D. R., 1996, Bite-force estimation for Tyrannosaurus rex from tooth-marked bones: Nature, v. 382, p. 706 - 708." title="Bite-force estimation for Tyrannosaurus rex from tooth-marked bones" type="journal article" year="1996">Erickson et al. 1996</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FAE64AD8FA1AF2DF" author="MEERS, M. B." box="[1319,1430,428,448]" journalOrPublisher="Historical Biology" pageId="7" pageNumber="170" pagination="1 - 12" part="16" refId="ref9691" refString="MEERS, M. B., 2002, Maximum bite force and prey size of Tyrannosaurus rex and their relationships to the inference of feeding behavior: Historical Biology, v. 16, p. 1 - 12." title="Maximum bite force and prey size of Tyrannosaurus rex and their relationships to the inference of feeding behavior" type="journal article" year="2002">Meers 2002</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FA604AD8FB82F2BB" author="BATES, K. T. & FALKINGHAM, P. L." journalOrPublisher="Biology Letters" pageId="7" pageNumber="170" pagination="660 - 664" part="8" refId="ref7293" refString="BATES, K. T. AND FALKINGHAM, P. L., 2012, Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics: Biology Letters, v. 8, p. 660 - 664." title="Estimating maximum bite performance in Tyrannosaurus rex using multi-body dynamics" type="journal article" year="2012">Bates and Falkingham 2012</bibRefCitation>
|
||
as predicted by
|
||
<bibRefCitation id="EFBCAAE44D59DE56FB0C4ABDFA1BF2BC" author="BAKKER, R. T. & WILLIAMS, M. & CURRIE, P. J." box="[1229,1431,456,476]" journalOrPublisher="Hunteria" pageId="7" pageNumber="170" pagination="1 - 30" part="1" refId="ref7243" refString="BAKKER, R. T., WILLIAMS, M., AND CURRIE, P. J., 1988, Nanotyrannus, a new genus of pygmy tyrannosaur, from the Latest Cretaceous of Montana: Hunteria, v. 1, p. 1 - 30." title="Nanotyrannus, a new genus of pygmy tyrannosaur, from the Latest Cretaceous of Montana" type="journal article" year="1988">Bakker et al. 1988</bibRefCitation>
|
||
), and tyrannosaurids probably had the strongest skulls (i.e., most resistant to bending) of any theropod (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FBEC4975FB45F174" author="HENDERSON, D. M." box="[1069,1225,512,532]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="7" pageNumber="170" pagination="766 - 778" part="22" refId="ref8780" refString="HENDERSON, D. M., 2002, The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force: Journal of Vertebrate Paleontology, v. 22, p. 766 - 778." title="The eyes have it: the sizes, shapes, and orientations of theropod orbits as indicators of skull strength and bite force" type="journal article" year="2002">Henderson 2002</bibRefCitation>
|
||
). Additionally, tyrannosaurid teeth occasionally show irregular spalled surfaces which are caused by tooth contact with food, probably bones, although it is difficult to determine whether this contact was intentional or not (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FADC4921FCEFF1E4" author="SCHUBERT, B. W. & UNGAR, P. S." journalOrPublisher="Acta Palaeontologica Polonica" pageId="7" pageNumber="170" pagination="93 - 99" part="50" refId="ref10584" refString="SCHUBERT, B. W. AND UNGAR, P. S., 2005, Wear facets and enamel spalling in tyrannosaurid dinosaurs: Acta Palaeontologica Polonica, v. 50, p. 93 - 99." title="Wear facets and enamel spalling in tyrannosaurid dinosaurs" type="journal article" year="2005">Schubert and Ungar 2005</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D59DE56FC8B49F9FC2BF0FC" blockId="7.[818,1492,175,1934]" pageId="7" pageNumber="170">
|
||
Further support for powerful bites in tyrannosaurids is found in their very strong teeth, wider and longer than most other theropod dinosaurs (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FCF949B1FC68F1B8" author="FARLOW, J. O. & BRINKMAN, D. L. & ABLER, W. L. & CURRIE, P. J." box="[824,996,708,728]" journalOrPublisher="Modern Geology" pageId="7" pageNumber="170" pagination="161 - 198" part="16" refId="ref8583" refString="FARLOW, J. O., BRINKMAN, D. L., ABLER, W. L., AND CURRIE, P. J., 1991, Size, shape, and serration density of theropod dinosaur lateral teeth: Modern Geology, v. 16, p. 161 - 198." title="Size, shape, and serration density of theropod dinosaur lateral teeth" type="journal article" year="1991">Farlow et al. 1991</bibRefCitation>
|
||
). Skeletally immature tyrannosaurids possess serrated and labiolingually compressed (ziphodont) teeth as do other theropods (although with crowns still thicker labiolingually than those of similarly sized theropods (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FC11486DFBB7F04C" author="HOLTZ, T. R." box="[976,1083,792,812]" editor="D. B. Weishampel & P. Dodson & H. Osmolska" journalOrPublisher="University of California Press, Berkeley, CA" pageId="7" pageNumber="170" pagination="111 - 136" refId="ref8878" refString="HOLTZ, T. R., 2004, Tyrannosauroidea, in D. B. Weishampel, P. Dodson, and H. Osmolska (eds.), The Dinosauria: University of California Press, Berkeley, CA, p. 111 - 136." title="Tyrannosauroidea" type="book chapter" volumeTitle="The Dinosauria" year="2004">Holtz 2004</bibRefCitation>
|
||
)), but skeletally mature tyrannosaurids have incrassate maxillary and dentary teeth (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FB5E4841FA85F028" author="HOLTZ, T. R." box="[1183,1289,820,840]" editor="D. H. Tanke & K. Carpenter" journalOrPublisher="Indiana University Press, Bloomington, Indiana" pageId="7" pageNumber="170" pagination="64 - 83" refId="ref8828" refString="HOLTZ, T. R., 2001, The phylogeny and taxonomy of the Tyrannosauridae, in D. H. Tanke and K. Carpenter (eds.), Mesozoic Vertebrate Life: Indiana University Press, Bloomington, Indiana, p. 64 - 83." title="The phylogeny and taxonomy of the Tyrannosauridae" type="book chapter" volumeTitle="Mesozoic Vertebrate Life" year="2001">Holtz 2001</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FAD74840FAC9F028" author="HOLTZ, T. R." box="[1302,1349,821,840]" editor="D. B. Weishampel & P. Dodson & H. Osmolska" journalOrPublisher="University of California Press, Berkeley, CA" pageId="7" pageNumber="170" pagination="111 - 136" refId="ref8878" refString="HOLTZ, T. R., 2004, Tyrannosauroidea, in D. B. Weishampel, P. Dodson, and H. Osmolska (eds.), The Dinosauria: University of California Press, Berkeley, CA, p. 111 - 136." title="Tyrannosauroidea" type="book chapter" volumeTitle="The Dinosauria" year="2004">2004</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FA904841FAF3F028" author="HOLTZ, T. R." box="[1361,1407,820,840]" editor="P. Larson & K. Carpenter" journalOrPublisher="Indiana University Press, Bloomington, IN" pageId="7" pageNumber="170" pagination="370 - 396" refId="ref8927" refString="HOLTZ, T. R., 2008, A critical reappraisal of the obligate scavenging hypothesis for Tyrannosaurus rex and other tyrant dinosaurs, in P. Larson and K. Carpenter (eds.), Tyrannosaurus rex: The Tyrant King: Indiana University Press, Bloomington, IN, p. 370 - 396." title="A critical reappraisal of the obligate scavenging hypothesis for Tyrannosaurus rex and other tyrant dinosaurs" type="book chapter" volumeTitle="Tyrannosaurus rex: The Tyrant King" year="2008">2008</bibRefCitation>
|
||
) that are considerably thicker labiolingually such that the labiolingual width is sometimes even thicker than the mesiodistal length (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FADC4819FA43F0E0" author="BAKKER, R. T. & WILLIAMS, M. & CURRIE, P. J." box="[1309,1487,876,896]" journalOrPublisher="Hunteria" pageId="7" pageNumber="170" pagination="1 - 30" part="1" refId="ref7243" refString="BAKKER, R. T., WILLIAMS, M., AND CURRIE, P. J., 1988, Nanotyrannus, a new genus of pygmy tyrannosaur, from the Latest Cretaceous of Montana: Hunteria, v. 1, p. 1 - 30." title="Nanotyrannus, a new genus of pygmy tyrannosaur, from the Latest Cretaceous of Montana" type="journal article" year="1988">Bakker et al. 1988</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FCF348FDFC11F0FC" author="HOLTZ, T. R." box="[818,925,904,924]" editor="D. B. Weishampel & P. Dodson & H. Osmolska" journalOrPublisher="University of California Press, Berkeley, CA" pageId="7" pageNumber="170" pagination="111 - 136" refId="ref8878" refString="HOLTZ, T. R., 2004, Tyrannosauroidea, in D. B. Weishampel, P. Dodson, and H. Osmolska (eds.), The Dinosauria: University of California Press, Berkeley, CA, p. 111 - 136." title="Tyrannosauroidea" type="book chapter" volumeTitle="The Dinosauria" year="2004">Holtz 2004</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D59DE56FC8B48D1FAAAF6D1" blockId="7.[818,1492,175,1934]" pageId="7" pageNumber="170">
|
||
Nevertheless, there are shallow bite traces attributed to tyrannosaurids, such as scrape feeding on a
|
||
<taxonomicName id="4C2DAC964D59DE56FB8D48B5FB32F0B4" authorityName="Brown" authorityYear="1912" box="[1100,1214,960,980]" class="Reptilia" family="Hadrosauridae" genus="Saurolophus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D59DE56FB8D48B5FB32F0B4" box="[1100,1214,960,980]" italics="true" pageId="7" pageNumber="170">Saurolophus</emphasis>
|
||
</taxonomicName>
|
||
humerus (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FAE948B4FCEFF090" author="HONE, D. W. E. & WATABE, M." journalOrPublisher="Acta Palaeontologica Polonica" pageId="7" pageNumber="170" pagination="627 - 634" part="55" refId="ref9049" refString="HONE, D. W. E. AND WATABE, M., 2010, New information on scavenging and selective feeding behavior of tyrannosaurids: Acta Palaeontologica Polonica, v. 55, p. 627 - 634." title="New information on scavenging and selective feeding behavior of tyrannosaurids" type="journal article" year="2010">Hone and Watabe 2010</bibRefCitation>
|
||
). The ability to selectively feed is reflected in the teeth of tyrannosaurids, which have more obvious heterodonty than other large carnivorous theropods (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FBCF4F60FBF0F749" author="SMITH, J. B." box="[1038,1148,1045,1065]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="7" pageNumber="170" pagination="865 - 887" part="25" refId="ref10711" refString="SMITH, J. B., 2005, Heterodonty in Tyrannosaurus rex: implications for the taxonomic and systematic utility of theropod dentitions: Journal of Vertebrate Paleontology, v. 25, p. 865 - 887." title="Heterodonty in Tyrannosaurus rex: implications for the taxonomic and systematic utility of theropod dentitions" type="journal article" year="2005">Smith 2005</bibRefCitation>
|
||
) in that their premaxillary teeth are considerably smaller than their maxillary (or lateral) teeth (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FA814F45FA2AF724" author="HOLTZ, T. R." box="[1344,1446,1072,1092]" editor="D. B. Weishampel & P. Dodson & H. Osmolska" journalOrPublisher="University of California Press, Berkeley, CA" pageId="7" pageNumber="170" pagination="111 - 136" refId="ref8878" refString="HOLTZ, T. R., 2004, Tyrannosauroidea, in D. B. Weishampel, P. Dodson, and H. Osmolska (eds.), The Dinosauria: University of California Press, Berkeley, CA, p. 111 - 136." title="Tyrannosauroidea" type="book chapter" volumeTitle="The Dinosauria" year="2004">Holtz 2004</bibRefCitation>
|
||
) and possess D-shaped cross-sections, as does
|
||
<emphasis id="B9590B074D59DE56FB764F3BFA93F700" box="[1207,1311,1102,1120]" italics="true" pageId="7" pageNumber="170">Eotyrannus</emphasis>
|
||
(
|
||
<bibRefCitation id="EFBCAAE44D59DE56FAEE4F38FA46F701" author="HUTT, S. D. & NAISH, D. & MARTILL, D. M. & BARKER, M. J. & NEWBERRY, P." box="[1327,1482,1101,1121]" journalOrPublisher="Cretaceous Research" pageId="7" pageNumber="170" pagination="227 - 242" part="22" refId="ref9195" refString="HUTT, S. D., NAISH, D., MARTILL, D. M., BARKER, M. J., AND NEWBERRY, P., 2001, A preliminary account of a new tyrannosauroid theropod from the Wessex Formation (Early Cretaceous) of southern England: Cretaceous Research, v. 22, p. 227 - 242." title="A preliminary account of a new tyrannosauroid theropod from the Wessex Formation (Early Cretaceous) of southern England" type="journal article" year="2001">Hutt et al. 2001</bibRefCitation>
|
||
). Although the premaxillary teeth of tyrannosauroids differ significantly in morphology from the rest of their dentition, there does not appear to be any significant difference in the average denticle densities on either the mesial or distal carinae when compared to those of the dentary or maxillary teeth (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FCF94FACFC53F78D" author="SMITH, J. B. & VANN, D. R. & DODSON, P." box="[824,991,1241,1261]" journalOrPublisher="The Anatomical Record Part A" pageId="7" pageNumber="170" pagination="699 - 736" part="285" refId="ref10751" refString="SMITH, J. B., VANN, D. R., AND DODSON, P., 2005, Dental morphology and variation in theropod dinosaurs: implications for the taxonomic identification of isolated teeth: The Anatomical Record Part A, v. 285, p. 699 - 736." title="Dental morphology and variation in theropod dinosaurs: implications for the taxonomic identification of isolated teeth" type="journal article" year="2005">Smith et al. 2005</bibRefCitation>
|
||
). The premaxillary teeth of tyrannosaurids were more closely spaced than the maxillary or dentary teeth, and they were probably used in nipping and scraping flesh off of bones (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FB274E64FA43F645" author="ERICKSON, G. M. & OLSON, K. H." box="[1254,1487,1297,1317]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="7" pageNumber="170" pagination="175 - 178" part="16" refId="ref8475" refString="ERICKSON, G. M. AND OLSON, K. H., 1996, Bite marks attributable to Tyrannosaurus rex: preliminary description and implications: Journal of Vertebrate Paleontology, v. 16, p. 175 - 178." title="Bite marks attributable to Tyrannosaurus rex: preliminary description and implications" type="journal article" year="1996">Erickson and Olson 1996</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FCF34E5BFB9EF620" author="HONE, D. W. E. & WATABE, M." box="[818,1042,1325,1345]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="7" pageNumber="170" pagination="627 - 634" part="55" refId="ref9049" refString="HONE, D. W. E. AND WATABE, M., 2010, New information on scavenging and selective feeding behavior of tyrannosaurids: Acta Palaeontologica Polonica, v. 55, p. 627 - 634." title="New information on scavenging and selective feeding behavior of tyrannosaurids" type="journal article" year="2010">Hone and Watabe 2010</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FBDE4E5BFB7EF621" author="HONE, D. W. E. & TANKE, D. H." box="[1055,1266,1325,1345]" journalOrPublisher="PeerJ" pageId="7" pageNumber="170" pagination="e 885" part="3" refId="ref8986" refString="HONE, D. W. E. AND TANKE, D. H., 2015, Pre- and postmortem tyrannosaurid bite marks on the remains of Daspletosaurus (Tyrannosaurinae: Theropoda) from Dinosaur Provincial Park, Alberta, Canada: PeerJ, v. 3, p. e 885, doi: DOI 10.7717 / peerj. 885." title="Pre- and postmortem tyrannosaurid bite marks on the remains of Daspletosaurus (Tyrannosaurinae: Theropoda) from Dinosaur Provincial Park, Alberta, Canada" type="journal article" year="2015">Hone and Tanke 2015</bibRefCitation>
|
||
), which would result in shallower tooth traces. We suspect that the scores on
|
||
<materialsCitation id="3B45DD484D59DE56FAE64E3FFA5FF63D" ID-GBIF-Occurrence="3396390305" box="[1319,1491,1354,1373]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="7" pageNumber="170" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
may be a combination of
|
||
<taxonomicName id="4C2DAC964D59DE56FBEC4E13FB39F618" authorityName="Osborn" authorityYear="1905" box="[1069,1205,1382,1400]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="170" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D59DE56FBEC4E13FB39F618" box="[1069,1205,1382,1400]" italics="true" pageId="7" pageNumber="170">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
premaxillary teeth and lateral teeth dragging along the surface to strip flesh off of the bone, given the varying depths and orientations of scores on the bone.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D59DE59FC8B4ECCFDD6F150" blockId="7.[818,1492,175,1934]" lastBlockId="8.[141,815,175,1934]" lastPageId="8" lastPageNumber="171" pageId="7" pageNumber="170">
|
||
There are several other examples in the fossil record of tyrannosaurid tooth traces on tyrannosaurid bones.
|
||
<bibRefCitation id="EFBCAAE44D59DE56FB454EA0FA94F689" author="JACOBSEN, A. R." box="[1156,1304,1493,1513]" journalOrPublisher="Historical Biology" pageId="7" pageNumber="170" pagination="17 - 26" part="13" refId="ref9262" refString="JACOBSEN, A. R., 1998, Feeding behavior of carnivorous dinosaurs as determined by tooth marks on dinosaur bones: Historical Biology, v. 13, p. 17 - 26." title="Feeding behavior of carnivorous dinosaurs as determined by tooth marks on dinosaur bones" type="journal article" year="1998">Jacobsen (1998)</bibRefCitation>
|
||
noted tyrannosaurid tooth traces on tyrannosaurid bones from the Dinosaur Park Formation, which he attributed to feeding. Tyrannosaurid face-biting due to intraspecific aggression has been suggested in the literature as a cause for tyrannosaurid skulls possessing tyrannosaurid tooth traces (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FAB24D30FC28F515" author="TANKE, D. H. & CURRIE, P. J." journalOrPublisher="Gaia" pageId="7" pageNumber="170" pagination="167 - 184" part="15" refId="ref10806" refString="TANKE, D. H. AND CURRIE, P. J., 1998, Head-biting behavior in theropod dinosaurs: paleopathological evidence: Gaia, v. 15, p. 167 - 184." title="Head-biting behavior in theropod dinosaurs: paleopathological evidence" type="journal article" year="1998">Tanke and Currie 1998</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D59DE56FC6F4D17FBE4F515" author="PETERSON, J. E. & HENDERSON, M. D. & SCHERER, R. P. & VITTORE, C. P." box="[942,1128,1633,1653]" journalOrPublisher="PALAIOS" pageId="7" pageNumber="170" pagination="780 - 784" part="24" refId="ref10212" refString="PETERSON, J. E., HENDERSON, M. D., SCHERER, R. P., AND VITTORE, C. P., 2009, Face biting on a juvenile tyrannosaurid and behavioral implications: PALAIOS, v. 24, p. 780 - 784." title="Face biting on a juvenile tyrannosaurid and behavioral implications" type="journal article" year="2009">Peterson et al. 2009</bibRefCitation>
|
||
).
|
||
<bibRefCitation id="EFBCAAE44D59DE56FBB84D17FAB1F515" author="PETERSON, J. E. & HENDERSON, M. D. & SCHERER, R. P. & VITTORE, C. P." box="[1145,1341,1633,1653]" journalOrPublisher="PALAIOS" pageId="7" pageNumber="170" pagination="780 - 784" part="24" refId="ref10212" refString="PETERSON, J. E., HENDERSON, M. D., SCHERER, R. P., AND VITTORE, C. P., 2009, Face biting on a juvenile tyrannosaurid and behavioral implications: PALAIOS, v. 24, p. 780 - 784." title="Face biting on a juvenile tyrannosaurid and behavioral implications" type="journal article" year="2009">Peterson et al. (2009)</bibRefCitation>
|
||
used the Extant Phylogenetic Bracket—a technique usually reserved for inferring the morphology of soft tissues in fossil organisms (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FB344DECFAE3F5CD" author="WITMER, L. M." box="[1269,1391,1689,1709]" editor="J. Thomason" journalOrPublisher="Cambridge University Press, Cambridge" pageId="7" pageNumber="170" pagination="19 - 33" refId="ref11005" refString="WITMER, L. M., 1995, The extant phylogenetic bracket and the importance of reconstructing soft tissues in fossils, in J. Thomason (ed.), Functional Morphology in Vertebrate Paleontology: Cambridge University Press, Cambridge, p. 19 - 33." title="The extant phylogenetic bracket and the importance of reconstructing soft tissues in fossils" type="book chapter" volumeTitle="Functional Morphology in Vertebrate Paleontology" year="1995">Witmer 1995</bibRefCitation>
|
||
)—to infer such behaviors could have occurred in tyrannosaurids, since intraspecific face biting can be found in the only two extant archosaur groups: crocodilians (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FC6F4D9BFBE6F461" author="PETERSON, J. E. & HENDERSON, M. D. & SCHERER, R. P. & VITTORE, C. P." box="[942,1130,1773,1793]" journalOrPublisher="PALAIOS" pageId="7" pageNumber="170" pagination="780 - 784" part="24" refId="ref10212" refString="PETERSON, J. E., HENDERSON, M. D., SCHERER, R. P., AND VITTORE, C. P., 2009, Face biting on a juvenile tyrannosaurid and behavioral implications: PALAIOS, v. 24, p. 780 - 784." title="Face biting on a juvenile tyrannosaurid and behavioral implications" type="journal article" year="2009">Peterson et al. 2009</bibRefCitation>
|
||
) and birds (
|
||
<bibRefCitation id="EFBCAAE44D59DE56FB1E4D98FA03F461" author="BLANCO, G. & TRAVERSO, J. M. & MARCHAMALO, J. & MARTINEZ, F." box="[1247,1423,1773,1793]" journalOrPublisher="Journal of Raptor Research" pageId="7" pageNumber="170" pagination="77 - 79" part="31" refId="ref7449" refString="BLANCO, G., TRAVERSO, J. M., MARCHAMALO, J., AND MARTINEZ, F., 1997, Interspecific and intraspecific aggression among Griffon and Cinereous vultures at nesting and foraging sites: Journal of Raptor Research, v. 31, p. 77 - 79." title="Interspecific and intraspecific aggression among Griffon and Cinereous vultures at nesting and foraging sites" type="journal article" year="1997">Blanco et al. 1997</bibRefCitation>
|
||
). Facebiting behavior in tyrannosaurs was also suggested to explain a tyrannosaurid left dentary (TMP 1996.05.13) with a tyrannosaurid tooth lodged in it from the Dinosaur Park Formation; however,
|
||
<bibRefCitation id="EFBCAAE44D59DE56FA844C37FCE2F412" author="BELL, P. R. & CURRIE, P. J." journalOrPublisher="Lethaia" pageId="7" pageNumber="170" pagination="278 - 281" part="43" refId="ref7333" refString="BELL, P. R. AND CURRIE, P. J., 2010, A tyrannosaur jaw bitten by a confamilial: scavenging or fatal agonism?: Lethaia, v. 43, p. 278 - 281." title="A tyrannosaur jaw bitten by a confamilial: scavenging or fatal agonism?" type="journal article" year="2010">Bell and Currie (2010)</bibRefCitation>
|
||
were unable to determine whether the bite occurred pre- or postmortem, nor could they discern whether the biter was the same species as the victim. A skull and mandible from a skeletally immature specimen of the tyrannosaurine
|
||
<emphasis id="B9590B074D56DE59FE974BBEFE73F3BE" box="[342,511,203,223]" italics="true" pageId="8" pageNumber="171">
|
||
<taxonomicName id="4C2DAC964D56DE59FE974BBEFE68F3BF" authorityName="Russell" authorityYear="1970" box="[342,484,203,223]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
sp
|
||
</emphasis>
|
||
. (TMP 1994.143.0001) from the Dinosaur Park Formation of Dinosaur Provincial Park, Alberta, Canada appears to show both healed, premortem tooth traces and postmortem tooth traces on the mandible, all made by another tyrannosaurid (
|
||
<bibRefCitation id="EFBCAAE44D56DE59FD114A55FF70F22F" author="HONE, D. W. E. & TANKE, D. H." journalOrPublisher="PeerJ" pageId="8" pageNumber="171" pagination="e 885" part="3" refId="ref8986" refString="HONE, D. W. E. AND TANKE, D. H., 2015, Pre- and postmortem tyrannosaurid bite marks on the remains of Daspletosaurus (Tyrannosaurinae: Theropoda) from Dinosaur Provincial Park, Alberta, Canada: PeerJ, v. 3, p. e 885, doi: DOI 10.7717 / peerj. 885." title="Pre- and postmortem tyrannosaurid bite marks on the remains of Daspletosaurus (Tyrannosaurinae: Theropoda) from Dinosaur Provincial Park, Alberta, Canada" type="journal article" year="2015">Hone and Tanke 2015</bibRefCitation>
|
||
). The authors attributed the premortem traces to intraspecific combat and the postmortem traces to scavenging. However, the authors were unable to determine if the scavenging was cannibalistic because of the presence of two tyrannosaurid species in the Dinosaur Park Formation, as in the case of TMP 1996.05.13 and the tyrannosaurid bones studied by
|
||
<bibRefCitation id="EFBCAAE44D56DE59FF4C4ABDFEA6F2BC" author="JACOBSEN, A. R." box="[141,298,456,476]" journalOrPublisher="Historical Biology" pageId="8" pageNumber="171" pagination="17 - 26" part="13" refId="ref9262" refString="JACOBSEN, A. R., 1998, Feeding behavior of carnivorous dinosaurs as determined by tooth marks on dinosaur bones: Historical Biology, v. 13, p. 17 - 26." title="Feeding behavior of carnivorous dinosaurs as determined by tooth marks on dinosaur bones" type="journal article" year="1998">Jacobsen (1998)</bibRefCitation>
|
||
. According to
|
||
<bibRefCitation id="EFBCAAE44D56DE59FE054ABCFD3DF2BC" author="HONE, D. W. E. & TANKE, D. H." box="[452,689,456,476]" journalOrPublisher="PeerJ" pageId="8" pageNumber="171" pagination="e 885" part="3" refId="ref8986" refString="HONE, D. W. E. AND TANKE, D. H., 2015, Pre- and postmortem tyrannosaurid bite marks on the remains of Daspletosaurus (Tyrannosaurinae: Theropoda) from Dinosaur Provincial Park, Alberta, Canada: PeerJ, v. 3, p. e 885, doi: DOI 10.7717 / peerj. 885." title="Pre- and postmortem tyrannosaurid bite marks on the remains of Daspletosaurus (Tyrannosaurinae: Theropoda) from Dinosaur Provincial Park, Alberta, Canada" type="journal article" year="2015">Hone and Tanke (2015)</bibRefCitation>
|
||
, the definite postmortem bite trace on the dentary was probably made by a tooth from the maxilla or non-anterior dentary of a tyrannosaurid because tyrannosaurid premaxillary teeth are more closely spaced.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D56DE59FF64494DFF43F669" blockId="8.[141,815,175,1934]" pageId="8" pageNumber="171">
|
||
<bibRefCitation id="EFBCAAE44D56DE59FF64494DFEFCF12C" author="LONGRICH, N. R. & HORNER, J. R. & ERICKSON, G. M. & CURRIE, P. J." box="[165,368,568,588]" journalOrPublisher="PLoS ONE" pageId="8" pageNumber="171" pagination="e 13419" part="5" refId="ref9597" refString="LONGRICH, N. R., HORNER, J. R., ERICKSON, G. M., AND CURRIE, P. J., 2010, Cannibalism in Tyrannosaurus rex: PLoS ONE, v. 5, p. e 13419, doi: doi: 10.1371 / journal. pone. 0013419." title="Cannibalism in Tyrannosaurus rex" type="journal article" year="2010">Longrich et al. (2010)</bibRefCitation>
|
||
described four specimens of
|
||
<taxonomicName id="4C2DAC964D56DE59FD40494CFCA2F12B" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[641,814,569,587]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D56DE59FD40494CFCA2F12B" box="[641,814,569,587]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<materialsCitation id="3B45DD484D56DE59FF524920FEAFF108" ID-GBIF-Occurrence="3396390344" box="[147,291,596,616]" collectionCode="UCMP" httpUri="https://ucmpdb.berkeley.edu/cgi/ucmp_query2?admin=&query_src=ucmp_index&table=ucmp2&spec_id=V137538&one=T" pageId="8" pageNumber="171" specimenCode="UCMP 137538">UCMP 137538</materialsCitation>
|
||
,
|
||
<materialsCitation id="3B45DD484D56DE59FEEF4920FE14F108" ID-GBIF-Occurrence="3396390306" box="[302,408,596,616]" collectionCode="MOR" pageId="8" pageNumber="171" specimenCode="MOR 1126">MOR 1126</materialsCitation>
|
||
,
|
||
<materialsCitation id="3B45DD484D56DE59FE654920FD88F108" ID-GBIF-Occurrence="3396390338" box="[420,516,597,616]" collectionCode="MOR" pageId="8" pageNumber="171" specimenCode="MOR 920">MOR 920</materialsCitation>
|
||
, and
|
||
<materialsCitation id="3B45DD484D56DE59FDF94920FD2EF108" ID-GBIF-Occurrence="3396390353" box="[568,674,596,616]" collectionCode="MOR" pageId="8" pageNumber="171" specimenCode="MOR 1602">MOR 1602</materialsCitation>
|
||
) from the Hell Creek Formation of Montana that possess tooth scores made by a large, predatory animal. Five furrows—called gouges by
|
||
<bibRefCitation id="EFBCAAE44D56DE59FD9249F9FC99F1C0" author="LONGRICH, N. R. & HORNER, J. R. & ERICKSON, G. M. & CURRIE, P. J." box="[595,789,652,672]" journalOrPublisher="PLoS ONE" pageId="8" pageNumber="171" pagination="e 13419" part="5" refId="ref9597" refString="LONGRICH, N. R., HORNER, J. R., ERICKSON, G. M., AND CURRIE, P. J., 2010, Cannibalism in Tyrannosaurus rex: PLoS ONE, v. 5, p. e 13419, doi: doi: 10.1371 / journal. pone. 0013419." title="Cannibalism in Tyrannosaurus rex" type="journal article" year="2010">Longrich et al. (2010)</bibRefCitation>
|
||
— mark the proximal end of
|
||
<materialsCitation id="3B45DD484D56DE59FE5249DCFDA8F1DC" ID-GBIF-Occurrence="3396390316" box="[403,548,680,700]" collectionCode="UCMP" httpUri="https://ucmpdb.berkeley.edu/cgi/ucmp_query2?admin=&query_src=ucmp_index&table=ucmp2&spec_id=V137538&one=T" pageId="8" pageNumber="171" specimenCode="UCMP 137538">UCMP 137538</materialsCitation>
|
||
, an isolated pedal phalanx. Another pedal phalanx,
|
||
<materialsCitation id="3B45DD484D56DE59FEA949B0FE5DF1B8" ID-GBIF-Occurrence="3396390349" box="[360,465,708,728]" collectionCode="MOR" pageId="8" pageNumber="171" specimenCode="MOR 1126">MOR 1126</materialsCitation>
|
||
, this time from a partial skeleton, also possesses tooth traces in the form of furrows and scores with at least one containing denticle striae.
|
||
<materialsCitation id="3B45DD484D56DE59FE404988FE52F070" ID-GBIF-Occurrence="3396390308" box="[385,478,765,784]" collectionCode="MOR" pageId="8" pageNumber="171" specimenCode="MOR 920">MOR 920</materialsCitation>
|
||
is a left humerus that was found as part of an associated, skeletally mature
|
||
<taxonomicName id="4C2DAC964D56DE59FDE0486CFD59F04B" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[545,725,793,811]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D56DE59FDE0486CFD59F04B" box="[545,725,793,811]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
skeleton featuring several scores on its posterior surface. The final specimen they described (
|
||
<materialsCitation id="3B45DD484D56DE59FF324824FED3F004" ID-GBIF-Occurrence="3396390346" box="[243,351,848,868]" collectionCode="MOR" pageId="8" pageNumber="171" specimenCode="MOR 1602">MOR 1602</materialsCitation>
|
||
) was an isolated right metatarsal III missing the proximal half of its shaft. Two scores can be found on the medial surface. All of these tooth traces are attributed by the authors to feeding rather than fighting because the bites would have been difficult to inflict on a live animal, as three of the examples are from the feet. Additionally, at least one of the specimens (
|
||
<materialsCitation id="3B45DD484D56DE59FEF148A8FE14F090" ID-GBIF-Occurrence="3396390323" box="[304,408,988,1008]" collectionCode="MOR" pageId="8" pageNumber="171" specimenCode="MOR 1126">MOR 1126</materialsCitation>
|
||
) was bitten more than once, and the score on
|
||
<materialsCitation id="3B45DD484D56DE59FF4C488CFF79F76C" ID-GBIF-Occurrence="3396390307" box="[141,245,1016,1036]" collectionCode="MOR" pageId="8" pageNumber="171" specimenCode="MOR 1602">MOR 1602</materialsCitation>
|
||
runs across the bone’s articulation with metatarsal II. None of the bites showed any evidence of healing, which confirms that they must have been bitten shortly before, at, or after death. These tooth scores are smaller in width than others previously attributed to
|
||
<taxonomicName id="4C2DAC964D56DE59FD644F3BFE07F71C" authority="(Erickson and Olson 1996)" baseAuthorityName="Erickson and Olson" baseAuthorityYear="1996" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FD644F3BFCA1F700" box="[677,813,1102,1120]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus</emphasis>
|
||
(
|
||
<bibRefCitation id="EFBCAAE44D56DE59FF524F1DFE08F71C" author="ERICKSON, G. M. & OLSON, K. H." box="[147,388,1128,1148]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="8" pageNumber="171" pagination="175 - 178" part="16" refId="ref8475" refString="ERICKSON, G. M. AND OLSON, K. H., 1996, Bite marks attributable to Tyrannosaurus rex: preliminary description and implications: Journal of Vertebrate Paleontology, v. 16, p. 175 - 178." title="Bite marks attributable to Tyrannosaurus rex: preliminary description and implications" type="journal article" year="1996">Erickson and Olson 1996</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, which led
|
||
<bibRefCitation id="EFBCAAE44D56DE59FE3B4F1DFD4BF71C" author="LONGRICH, N. R. & HORNER, J. R. & ERICKSON, G. M. & CURRIE, P. J." box="[506,711,1128,1148]" journalOrPublisher="PLoS ONE" pageId="8" pageNumber="171" pagination="e 13419" part="5" refId="ref9597" refString="LONGRICH, N. R., HORNER, J. R., ERICKSON, G. M., AND CURRIE, P. J., 2010, Cannibalism in Tyrannosaurus rex: PLoS ONE, v. 5, p. e 13419, doi: doi: 10.1371 / journal. pone. 0013419." title="Cannibalism in Tyrannosaurus rex" type="journal article" year="2010">Longrich et al. (2010)</bibRefCitation>
|
||
to suggest they were made by juvenile or sub-adult
|
||
<taxonomicName id="4C2DAC964D56DE59FDC14FF3FD04F7F8" baseAuthorityName="Erickson and Olson" baseAuthorityYear="1996" box="[512,648,1158,1176]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FDC14FF3FD04F7F8" box="[512,648,1158,1176]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
individuals (these authors consider
|
||
<taxonomicName id="4C2DAC964D56DE59FEEB4FD7FE26F7D4" box="[298,426,1186,1204]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FEEB4FD7FE26F7D4" box="[298,426,1186,1204]" italics="true" pageId="8" pageNumber="171">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
to be a juvenile
|
||
<taxonomicName id="4C2DAC964D56DE59FD884FD7FD7AF7D4" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[585,758,1186,1204]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D56DE59FD884FD7FD7AF7D4" box="[585,758,1186,1204]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
). The authors reason that because they only studied a relatively limited sample of tooth-marked bones, cannibalism must have been common in
|
||
<taxonomicName id="4C2DAC964D56DE59FD184FAFFF45F668" baseAuthorityName="Erickson and Olson" baseAuthorityYear="1996" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FD184FAFFF45F668" italics="true" pageId="8" pageNumber="171">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D56DE59FF644E67FCA9F47D" blockId="8.[141,815,175,1934]" pageId="8" pageNumber="171">
|
||
<materialsCitation id="3B45DD484D56DE59FF644E67FEC3F645" ID-GBIF-Occurrence="3396390326" box="[165,335,1298,1317]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="8" pageNumber="171" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
is also suggestive of cannibalism in
|
||
<taxonomicName id="4C2DAC964D56DE59FD644E67FF26F620" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D56DE59FD644E67FF26F620" italics="true" pageId="8" pageNumber="171">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
. That the tooth traces represent feeding rather than some other biting behavior is supported by four lines of reasoning. First, although it is possible that a
|
||
<taxonomicName id="4C2DAC964D56DE59FED64E13FE13F618" baseAuthorityName="Erickson and Olson" baseAuthorityYear="1996" box="[279,415,1382,1400]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FED64E13FE13F618" box="[279,415,1382,1400]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
could bite the foot of another
|
||
<taxonomicName id="4C2DAC964D56DE59FD7B4E13FF26F6F4" baseAuthorityName="Erickson and Olson" baseAuthorityYear="1996" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FD7B4E13FF26F6F4" italics="true" pageId="8" pageNumber="171">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
in intraspecific combat, it seems unlikely, especially since there appear to be multiple bites in the same area. Second, these scores are perpendicular to the long axis of the bone, which would be expected for a feeding trace (
|
||
<bibRefCitation id="EFBCAAE44D56DE59FEDF4EA0FDA3F689" author="NJAU, J. K. & BLUMENSCHINE, R. J." box="[286,559,1493,1513]" journalOrPublisher="Journal of Human Evolution" pageId="8" pageNumber="171" pagination="142 - 162" part="50" refId="ref9872" refString="NJAU, J. K. AND BLUMENSCHINE, R. J., 2006, A diagnosis of crocodile feeding traces on larger mammal bone, with fossil examples from the Plio-Pleistocene Olduvai Basin, Tanzania: Journal of Human Evolution, v. 50, p. 142 - 162." title="A diagnosis of crocodile feeding traces on larger mammal bone, with fossil examples from the Plio-Pleistocene Olduvai Basin, Tanzania" type="journal article" year="2006">Njau and Blumenschine 2006</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D56DE59FDF84EA0FD64F689" author="POBINER, B. L. & DESILVA, J. & SANDERS, W. J. & MITANI, J. C." box="[569,744,1493,1513]" journalOrPublisher="Journal of Human Evolution" pageId="8" pageNumber="171" pagination="614 - 636" part="52" refId="ref10320" refString="POBINER, B. L., DESILVA, J., SANDERS, W. J., AND MITANI, J. C., 2007, Taphonomic analysis of skeletal remains from chimpanzee hunts at Ngogo, Kibale National Park, Uganda: Journal of Human Evolution, v. 52, p. 614 - 636." title="Taphonomic analysis of skeletal remains from chimpanzee hunts at Ngogo, Kibale National Park, Uganda" type="journal article" year="2007">Pobiner et al. 2007</bibRefCitation>
|
||
). Third, all of these marks are examples of scores, where the teeth are dragging along the surface of the bone, rather than pits or punctures. This means that the animal was scraping its jaw along the foot, not just simply biting it as might be expected in a combat scenario. Scores could also occur in a scenario where the feeder is dragging a carcass; however, these marks are shallow and lack pits or punctures, which is consistent with the defleshing of a bone during feeding. Fourth, the three scores in cluster 1 begin as separate traces, but then join together to form a single score (
|
||
<figureCitation id="1316CB904D56DE59FD7D4DC0FD78F5A9" box="[700,756,1717,1737]" captionStart="FIG. 4" captionStartId="6.[141,167,1865,1881]" captionTargetId="figure@5.[148,1457,176,1930]" captionTargetPageId="5" captionText="FIG. 4.—Histology of a Tyrannosaurus rex metatarsal (SWAU HRS13997) in cross section under different light regimes. A) Section of cortical bone from the medullary cavity (left) to the subperiosteal surface (right) under regularly transmitted light. Boxes show where areas of this bone are shown in higher detail under regularly transmitted light (B, F, J), cross-polarized light (C, G, K), cross-polarized light with a waveplate (D, H, L), and reflected light (E, I, M). Scale bar for A = 5 mm, all other scale bars = 1 mm." figureDoi="http://doi.org/10.5281/zenodo.4023994" httpUri="https://zenodo.org/record/4023994/files/figure.png" pageId="8" pageNumber="171">Fig. 4</figureCitation>
|
||
). This suggests that the animal is turning its head as it is biting, probably to scrape off some flesh from the bone. The combination of all of these factors suggests that the biter was indeed feeding on the other individual’s foot.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D56DE59FF644C53FC44F299" blockId="8.[141,815,175,1934]" lastBlockId="8.[845,1518,175,730]" pageId="8" pageNumber="171">
|
||
Whether the larger
|
||
<taxonomicName id="4C2DAC964D56DE59FE984C52FE6DF459" baseAuthorityName="Erickson and Olson" baseAuthorityYear="1996" box="[345,481,1831,1849]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FE984C52FE6DF459" box="[345,481,1831,1849]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
killed the smaller individual cannot be determined from the available evidence. However, considering that it would be difficult for a larger
|
||
<taxonomicName id="4C2DAC964D56DE59FE5F4C2AFDAAF411" baseAuthorityName="Erickson and Olson" baseAuthorityYear="1996" box="[414,550,1887,1905]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FE5F4C2AFDAAF411" box="[414,550,1887,1905]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
to reach its jaws low enough to bite a smaller individual’s foot while the smaller individual was still standing, then these bites were likely made after the smaller individual was already lying on the ground. Additionally, there would not have been a great deal of meat attached to the metatarsals, which suggests this may be a scavenging scenario where the larger tyrannosaurid is attempting to get even the smallest bits of meat off of an already picked-over carcass. A larger
|
||
<taxonomicName id="4C2DAC964D56DE59FC4E4A48FB9BF22F" baseAuthorityName="Erickson and Olson" baseAuthorityYear="1996" box="[911,1047,317,335]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FC4E4A48FB9BF22F" box="[911,1047,317,335]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
feeding on a smaller individual could suggest infanticide, but there is no way to determine the relationship between the two individuals. Furthermore, given the similarity in size between
|
||
<materialsCitation id="3B45DD484D56DE59FA6E4A00FC3EF2C4" ID-GBIF-Occurrence="3396390303" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="8" pageNumber="171" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
and the metatarsal of
|
||
<materialsCitation id="3B45DD484D56DE59FB514AE4FA9AF2C4" ID-GBIF-Occurrence="3396390347" box="[1168,1302,400,420]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/ContactSheet.php?QueryName=DetailedQuery&StartAt=1&QueryPage=%2Fdinosaur-institute%2Findex.php&Restriction=%28exists+%28SecDepartment_tab+where+%28SecDepartment+contains+%27Dinosaur+Institute%27%29%29%29&col_SpeSpecimenNoInteger=23845&LimitPerPage=20&Search=Search& " pageId="8" pageNumber="171" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
, the small tyrannosaur would have probably been a subadult animal (
|
||
<bibRefCitation id="EFBCAAE44D56DE59FB2F4AD8FA6FF2A1" author="CARR, T. D. & WILLIAMSON, T. E." box="[1262,1507,429,449]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="8" pageNumber="171" pagination="479 - 523" part="142" refId="ref7675" refString="CARR, T. D. AND WILLIAMSON, T. E., 2004, Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America: Zoological Journal of the Linnean Society, v. 142, p. 479 - 523." title="Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America" type="book chapter" year="2004">Carr and Williamson 2004</bibRefCitation>
|
||
), which suggests it was likely living on its own without parental care for several years.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D56DE59FCA44977FB55F1BA" blockId="8.[845,1518,175,730]" pageId="8" pageNumber="171">
|
||
Because tyrannosaurids could and did eat bones (e.g.,
|
||
<bibRefCitation id="EFBCAAE44D56DE59FA944974FA66F175" author="CHIN, K. & TOKARYK, T. T. & ERICKSON, G. M. & CALK, L. C." box="[1365,1514,513,533]" journalOrPublisher="Nature" pageId="8" pageNumber="171" pagination="680 - 682" part="393" refId="ref7929" refString="CHIN, K., TOKARYK, T. T., ERICKSON, G. M., AND CALK, L. C., 1998, A king-sized theropod coprolite: Nature, v. 393, p. 680 - 682." title="A king-sized theropod coprolite" type="journal article" year="1998">Chin et al. 1998</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFBCAAE44D56DE59FC8C496BFCF1F151" author="CHIN, K. & EBERTH, D. A. & SCHWEITZER, M. H. & RANDO, T. A. & SLOBODA, W. J. & HORNER, J. R." box="[845,893,542,561]" journalOrPublisher="PALAIOS" pageId="8" pageNumber="171" pagination="286 - 294" part="18" refId="ref7857" refString="CHIN, K., EBERTH, D. A., SCHWEITZER, M. H., RANDO, T. A., SLOBODA, W. J., AND HORNER, J. R., 2003, Remarkable preservation of undigested muscle tissue within a Late Cretaceous tyrannosaurid coprolite from Alberta, Canada: PALAIOS, v. 18, p. 286 - 294." title="Remarkable preservation of undigested muscle tissue within a Late Cretaceous tyrannosaurid coprolite from Alberta, Canada" type="journal article" year="2003">2003</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFBCAAE44D56DE59FC494968FB90F151" author="VARRICCHIO, D. J." box="[904,1052,541,561]" journalOrPublisher="Journal of Paleontology" pageId="8" pageNumber="171" pagination="401 - 406" part="75" refId="ref10843" refString="VARRICCHIO, D. J., 2001, Gut contents from a Cretaceous tyrannosaurid: implications for theropod dinosaur digestive tracts: Journal of Paleontology, v. 75, p. 401 - 406." title="Gut contents from a Cretaceous tyrannosaurid: implications for theropod dinosaur digestive tracts" type="journal article" year="2001">Varricchio 2001</bibRefCitation>
|
||
), it is unclear why the larger tyrannosaurid did not simply swallow the smaller individual’s foot or at least this metatarsal whole. There does not appear to be any evidence of etching by digestive processes on
|
||
<materialsCitation id="3B45DD484D56DE59FC124906FB0FF1E6" ID-GBIF-Occurrence="3396390337" box="[979,1155,627,646]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="8" pageNumber="171" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
as has been noticed in other bones ingested by tyrannosaurids (
|
||
<bibRefCitation id="EFBCAAE44D56DE59FB9D49FBFB74F1C2" author="VARRICCHIO, D. J." box="[1116,1272,654,674]" journalOrPublisher="Journal of Paleontology" pageId="8" pageNumber="171" pagination="401 - 406" part="75" refId="ref10843" refString="VARRICCHIO, D. J., 2001, Gut contents from a Cretaceous tyrannosaurid: implications for theropod dinosaur digestive tracts: Journal of Paleontology, v. 75, p. 401 - 406." title="Gut contents from a Cretaceous tyrannosaurid: implications for theropod dinosaur digestive tracts" type="journal article" year="2001">Varricchio 2001</bibRefCitation>
|
||
). For whatever reason, it appears that this
|
||
<taxonomicName id="4C2DAC964D56DE59FC2749DEFBE2F1DD" baseAuthorityName="Erickson and Olson" baseAuthorityYear="1996" box="[998,1134,683,701]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FC2749DEFBE2F1DD" box="[998,1134,683,701]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
individual preferred to nip flesh off of the metatarsal rather than swallowing it whole.
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D56DE59FB9D4873FB53F076" blockId="8.[1116,1247,774,790]" box="[1116,1247,774,790]" pageId="8" pageNumber="171">
|
||
<heading id="D0DA60794D56DE59FB9D4873FB53F076" allCaps="true" bold="true" box="[1116,1247,774,790]" centered="true" fontSize="7" level="3" pageId="8" pageNumber="171" reason="2">
|
||
<emphasis id="B9590B074D56DE59FB9D4873FB53F076" bold="true" box="[1116,1247,774,790]" pageId="8" pageNumber="171">CONCLUSIONS</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8B92D7154D56DE59FCA44844FA16F78C" blockId="8.[845,1518,817,1260]" pageId="8" pageNumber="171">
|
||
The discovery of
|
||
<materialsCitation id="3B45DD484D56DE59FBCB4847FB3AF025" ID-GBIF-Occurrence="3396390315" box="[1034,1206,818,837]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="8" pageNumber="171" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
, a fourth metatarsal from a small tyrannosaurid, in the Lance Formation of Wyoming, possessing
|
||
<taxonomicName id="4C2DAC964D56DE59FA58483AFC05F01D" baseAuthorityName="Erickson and Olson" baseAuthorityYear="1996" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FA58483AFC05F01D" italics="true" pageId="8" pageNumber="171">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
tooth scores on its surface, is best interpreted to be further evidence for cannibalistic behavior in this large theropod. Although cannibalism is a likely explanation for these tooth scores, it is also possible that a
|
||
<taxonomicName id="4C2DAC964D56DE59FC8C48CAFC73F0B1" baseAuthorityName="Carr and Williamson" baseAuthorityYear="2004" box="[845,1023,959,977]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B9590B074D56DE59FC8C48CAFC73F0B1" box="[845,1023,959,977]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
was feeding upon an immature individual from a currently unknown different species of tyrannosaurid. Even if this bone does pertain to a separate species, this particular fossil presents us with insights into tyrannosaur feeding behavior. It appears that on this occasion, a
|
||
<taxonomicName id="4C2DAC964D56DE59FCA14F45FC64F722" baseAuthorityName="Erickson and Olson" baseAuthorityYear="1996" box="[864,1000,1072,1090]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="171" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9590B074D56DE59FCA14F45FC64F722" box="[864,1000,1072,1090]" italics="true" pageId="8" pageNumber="171">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
stripped flesh off of a bone rather than swallowing the bone whole, even when very little flesh was present. This find represents the first evidence for tyrannosaurids feeding on tyrannosaurids in the Lance Formation, as well as the first instance of
|
||
<taxonomicName id="4C2DAC964D56DE59FAED4FF6FC1DF7D4" class="Reptilia" genus="Knethichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="171" phylum="Chordata" rank="species" species="subparallelum">
|
||
<emphasis id="B9590B074D56DE59FAED4FF6FC1DF7D4" italics="true" pageId="8" pageNumber="171">Knethichnus subparallelum</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C2DAC964D56DE59FC7F4FD5FBE4F7D3" baseAuthorityName="Jacobsen and Bromley" baseAuthorityYear="2009" box="[958,1128,1184,1204]" class="Reptilia" genus="Linichnus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="8" pageNumber="171" phylum="Chordata" rank="species" species="serratus">
|
||
<emphasis id="B9590B074D56DE59FC7F4FD5FBE4F7D3" box="[958,1128,1184,1204]" italics="true" pageId="8" pageNumber="171">Linichnus serratus</emphasis>
|
||
</taxonomicName>
|
||
on a tyrannosaurid bone. It is possible that further excavations at the location of discovery of
|
||
<materialsCitation id="3B45DD484D56DE59FAD64FC9FA4DF7AF" ID-GBIF-Occurrence="3396390312" box="[1303,1473,1212,1231]" collectingCountry="United States" collectingRegion="Wyoming" collectionCode="SWAU" county="Niobrara" formation="Lance Formation" pageId="8" pageNumber="171" preparations="partial right metatarsal IV" specimenCode="SWAU HRS13997">SWAU HRS13997</materialsCitation>
|
||
may uncover more material from this small scavenged tyrannosaurid.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |