476 lines
74 KiB
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476 lines
74 KiB
XML
<document id="85C563605B0D31BE5D569FCAB3BA9594" ID-DOI="10.1073/pnas.1216534110" ID-GBIF-Dataset="2beffdbf-ef4c-4992-b131-97fba3dd3a8f" ID-PMC="PMC3732924" ID-PubMed="23858435" ID-Zenodo-Dep="3748449" IM.taxonomicNames_approvedBy="jeremy" checkinTime="1586604899271" checkinUser="jeremy" docAuthor="Robert A. DePalma II, David A. Burnham, Larry D. Martin, Bruce M. Rothschild & Peter L. Larson" docDate="2013" docId="03B887AEA72CF054FF88FC4CBFD1FC86" docLanguage="en" docName="dePalmaetal2013TrexPredator.pdf.imf" docOrigin="Proceedings of the National Academy of Sciences 110 (31)" docStyle="DocumentStyle{}" docTitle="Tyrannosaurus rex Osborn 1905" docType="treatment" docVersion="12" lastPageNumber="12563" masterDocId="FF81FFD6A72DF057FFF6FFD7BA56FFAA" masterDocTitle="Physical evidence of predatory behavior in Tyrannosaurus rex" masterLastPageNumber="12564" masterPageNumber="12560" pageNumber="12561" updateTime="1689602936628" updateUser="jeremy">
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<mods:title id="AD9F13A3B1C9CC60F57585CC373936CD">Physical evidence of predatory behavior in Tyrannosaurus rex</mods:title>
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<mods:roleTerm id="2DFB0275CFCCE63F41A5487BBB2944A5">Author</mods:roleTerm>
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<mods:namePart id="103AC3BF2394E9D976DAB78F22652364">Robert A. DePalma II</mods:namePart>
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<mods:affiliation id="C734155287EA41A7742C22EBE6A0D284">Department of Paleontology, Palm Beach Museum of Natural History, Ft. Lauderdale, FL 33306</mods:affiliation>
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<mods:namePart id="8134A7A8742BE6077CB1E3FF903A2DA1">David A. Burnham</mods:namePart>
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<mods:affiliation id="C325DE9A574E86ECD47B3AF369D93CF6">Division of Vertebrate Paleontology, University of Kansas Biodiversity Institute, Lawrence, KS 66045</mods:affiliation>
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<mods:nameIdentifier id="90AEC8E90B1F73140A47741DEF413CEC" type="email">dinosaur@ku.edu</mods:nameIdentifier>
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<mods:namePart id="7037B13B9A59D1C02FCEDB6A22F6299C">Larry D. Martin</mods:namePart>
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<mods:affiliation id="EE25406E795E447B17ACF92D5CAF5724">Division of Vertebrate Paleontology, University of Kansas Biodiversity Institute, Lawrence, KS 66045</mods:affiliation>
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<mods:namePart id="A2DD3D53EB4615DF167E715EFB9BB631">Bruce M. Rothschild</mods:namePart>
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<mods:affiliation id="5A70710C64A2B12FBFD54A68D3D57992">Division of Vertebrate Paleontology, University of Kansas Biodiversity Institute, Lawrence, KS 66045</mods:affiliation>
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<mods:namePart id="A696A73F63ACB3CE32E671C63F408ACA">Peter L. Larson</mods:namePart>
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<mods:affiliation id="5C210268FA1F7EFA703B178B04093500">Black Hills Institute of Geological Research, Inc., Hill City, SD 57745</mods:affiliation>
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<mods:title id="65CC09F2BFA9A1AB7B0129D458842951">Proceedings of the National Academy of Sciences</mods:title>
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<mods:part id="025D9659F1FA7B4204E8FD61DCE77A1C">
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<mods:date id="1B41A85036F55605BA721D568EE073FA">2013</mods:date>
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<mods:number id="0638B6E0F32258213C02B90827104B96">2013-07-30</mods:number>
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<treatment id="03B887AEA72CF054FF88FC4CBFD1FC86" ID-DOI="http://doi.org/10.5281/zenodo.3809064" ID-GBIF-Taxon="184097720" ID-Zenodo-Dep="3809064" LSID="urn:lsid:plazi:treatment:03B887AEA72CF054FF88FC4CBFD1FC86" httpUri="http://treatment.plazi.org/id/03B887AEA72CF054FF88FC4CBFD1FC86" lastPageId="3" lastPageNumber="12563" pageId="1" pageNumber="12561">
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<subSubSection id="C30B6533A72CF056FF88FC4CBA94FBDB" pageId="1" pageNumber="12561" type="discussion">
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<paragraph id="8BAE36B8A72CF056FF88FC4CBA94FBDB" blockId="1.[102,756,922,1496]" pageId="1" pageNumber="12561">
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Two coalesced hadrosaur (compare with
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<emphasis id="B965EAAAA72CF056FDA4FC4DBA9AFC66" italics="true" pageId="1" pageNumber="12561">Edmontosaurus annectens</emphasis>
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) caudal vertebrae were discovered in the Hell Creek Formation of Harding County, South Dakota (
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<bibRefCitation id="EF804B49A72CF056FDA1FC06B822FC4D" author="DePalma RA" box="[599,628,977,999]" journalOrPublisher="Univ of Kansas, Lawrence, KS" pageId="1" pageNumber="12561" refId="ref5259" refString="40. DePalma RA (2010) Geologyι Taphonomyι and Paleoecology of a Unique Upper cretaceous Bonebed Near the cretaceous-Tertiary Boundary in South Dakota., Master' s thesis (Univ of Kansas, Lawrence, KS)." title="Geologyι Taphonomyι and Paleoecology of a Unique Upper cretaceous Bonebed Near the cretaceous-Tertiary Boundary in South Dakota" type="book" year="2010">40</bibRefCitation>
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). Archosaur fauna identified in this site include crocodiles, dinosaurs, and birds (
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<bibRefCitation id="EF804B49A72CF056FF51FBDFBA95FBB4" author="White PD & Fastovsky DE & Sheehan PM" box="[167,195,1032,1054]" journalOrPublisher="Palaios" pageId="1" pageNumber="12561" pagination="41 - 51" part="13" refId="ref5298" refString="41. White PD, Fastovsky DE, Sheehan PM (1998) Taphonomy and suggested structure of the dinosaurian assemblage of the Hell Creek Formation (Maastrichtian), eastern Montana and western North Dakota. Palaios 13 (1): 41 - 51." title="Taphonomy and suggested structure of the dinosaurian assemblage of the Hell Creek Formation (Maastrichtian), eastern Montana and western North Dakota" type="journal article" year="1998">41</bibRefCitation>
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). Physical evidence of dental penetration and extensive infection (osteomylitis) of the fused vertebral centra and healing (bone overgrowth) document an unsuccessful attack by a large predator.
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</paragraph>
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</subSubSection>
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<subSubSection id="C30B6533A72CF056FF3DFB8CB870FB26" pageId="1" pageNumber="12561" type="nomenclature">
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<paragraph id="8BAE36B8A72CF056FF3DFB8CB870FB26" blockId="1.[102,756,922,1496]" pageId="1" pageNumber="12561">
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A tooth crown was discovered within the wound, permitting identification of the predator as
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<taxonomicName id="4C114D3BA72CF056FE1AFBAFB876FB27" authorityName="Osborn" authorityYear="1905" box="[492,544,1144,1165]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B965EAAAA72CF056FE1AFBAFB876FB27" box="[492,544,1144,1165]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
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</taxonomicName>
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.
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</paragraph>
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</subSubSection>
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<subSubSection id="C30B6533A72CF056FDDDFBA0BAA6FA72" pageId="1" pageNumber="12561" type="discussion">
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<paragraph id="8BAE36B8A72CF056FDDDFBA0BAA6FA72" blockId="1.[102,756,922,1496]" pageId="1" pageNumber="12561">
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This is unambiguous evidence that
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<taxonomicName id="4C114D3BA72CF056FF03FB44BB78FB02" authorityName="Osborn" authorityYear="1905" box="[245,302,1171,1192]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B965EAAAA72CF056FF03FB44BB78FB02" box="[245,302,1171,1192]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
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</taxonomicName>
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was an active predator, fulfilling the criteria that Farlow and Holtz (
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<bibRefCitation id="EF804B49A72CF056FEA5FB79BB36FB6E" author="Farlow JO & Holtz TR" box="[339,352,1198,1220]" journalOrPublisher="Paleontological Society Papers" pageId="1" pageNumber="12561" pagination="251 - 266" part="8" refId="ref3943" refString="3. Farlow JO, Holtz TR (2002) The fossil record of predation in dinosaurs. Paleontological Society Papers 8: 251 - 266." title="The fossil record of predation in dinosaurs" type="journal article" year="2002">3</bibRefCitation>
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) advanced. As
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<taxonomicName id="4C114D3BA72CF056FE0DFB78B867FB6E" authorityName="Osborn" authorityYear="1905" box="[507,561,1199,1220]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B965EAAAA72CF056FE0DFB78B867FB6E" box="[507,561,1199,1220]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
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</taxonomicName>
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comprises between 1% and 16% of the Upper Cretaceous dinosaurian fauna in Western North America (
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<bibRefCitation id="EF804B49A72CF056FE9CFB32BBD5FB51" author="White PD & Fastovsky DE & Sheehan PM" box="[362,387,1253,1275]" journalOrPublisher="Palaios" pageId="1" pageNumber="12561" pagination="41 - 51" part="13" refId="ref5298" refString="41. White PD, Fastovsky DE, Sheehan PM (1998) Taphonomy and suggested structure of the dinosaurian assemblage of the Hell Creek Formation (Maastrichtian), eastern Montana and western North Dakota. Palaios 13 (1): 41 - 51." title="Taphonomy and suggested structure of the dinosaurian assemblage of the Hell Creek Formation (Maastrichtian), eastern Montana and western North Dakota" type="journal article" year="1998">41</bibRefCitation>
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–
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<bibRefCitation id="EF804B49A72CF056FE7BFB32BBF1FB51" author="Eloff FC" box="[397,423,1253,1275]" journalOrPublisher="Koedo" pageId="1" pageNumber="12561" pagination="105 - 113" part="7" refId="ref5486" refString="45. Eloff FC (1964) On the predatory habits of lions and hyaenas. Koedo 7: 105 - 113." title="On the predatory habits of lions and hyaenas" type="journal article" year="1964">45</bibRefCitation>
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), its status as a predator or obligate scavenger is nontrivial and could have significant implications for paleoecological reconstructions of that time period. The present contribution provides unique information demonstrating the ecological role for
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<taxonomicName id="4C114D3BA72CF056FEE9FA83BB02FAC3" authorityName="Osborn" authorityYear="1905" box="[287,340,1364,1385]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B965EAAAA72CF056FEE9FA83BB02FAC3" box="[287,340,1364,1385]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
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</taxonomicName>
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as that of an active predator. Despite this documentation of predatory behavior by
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<taxonomicName id="4C114D3BA72CF056FE02FAA7B87EFA2F" authorityName="Osborn" authorityYear="1905" box="[500,552,1392,1413]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B965EAAAA72CF056FE02FAA7B87EFA2F" box="[500,552,1392,1413]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
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</taxonomicName>
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, we do not make the argument that
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<taxonomicName id="4C114D3BA72CF056FF0CFA5BBB66FA0B" authorityName="Osborn" authorityYear="1905" box="[250,304,1420,1441]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B965EAAAA72CF056FF0CFA5BBB66FA0B" box="[250,304,1420,1441]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
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</taxonomicName>
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was an obligate predator. Like most modern large predators (
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<bibRefCitation id="EF804B49A72CF056FEFBFA71BB71FA16" author="Stuart CT & Stuart MD" box="[269,295,1446,1468]" journalOrPublisher="Struik Publishers, Cape Town, South Africa" pageId="1" pageNumber="12561" refId="ref4793" refString="27. Stuart CT, Stuart MD (1997) The Larger Mammals of Africa (Struik Publishers, Cape Town, South Africa)." title="The Larger Mammals of Africa" type="book" year="1997">27</bibRefCitation>
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,
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<bibRefCitation id="EF804B49A72CF056FEC5FA71BB1AFA16" author="Eloff FC" box="[307,332,1446,1468]" journalOrPublisher="Koedo" pageId="1" pageNumber="12561" pagination="105 - 113" part="7" refId="ref5486" refString="45. Eloff FC (1964) On the predatory habits of lions and hyaenas. Koedo 7: 105 - 113." title="On the predatory habits of lions and hyaenas" type="journal article" year="1964">45</bibRefCitation>
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) it almost certainly did also scavenge carcasses (
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<bibRefCitation id="EF804B49A72CF056FF47FA15BAE9FA72" author="Currie PJ" box="[177,191,1474,1496]" editor="Farlow JO & Brett-Surman MK" journalOrPublisher="Indiana Univ Press, Bloomington, IN" pageId="1" pageNumber="12561" pagination="216 - 233" refId="ref4130" refString="9. Currie PJ (1997) The complete Dinosaur, eds Farlow JO, Brett-Surman MK (Indiana Univ Press, Bloomington, IN), pp 216 - 233." title="The complete Dinosaur" type="book chapter" year="1997">9</bibRefCitation>
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,
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<bibRefCitation id="EF804B49A72CF056FF3CFA15BAB2FA72" author="Ryan MJ & Vickaryous MK" box="[202,228,1474,1496]" editor="Currie PJ & Padian K" journalOrPublisher="Academic, San Diego, CA" pageId="1" pageNumber="12561" pagination="169 - 174" refId="ref4411" refString="16. Ryan MJ, Vickaryous MK (1997) Encyclopedia of Dinosaurs, eds Currie PJ, Padian K (Academic, San Diego, CA), pp 169 - 174." title="Encyclopedia of Dinosaurs" type="book chapter" year="1997">16</bibRefCitation>
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).
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</paragraph>
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</subSubSection>
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<subSubSection id="C30B6533A72CF056FCEEFF5BB999FA72" pageId="1" pageNumber="12561" type="description">
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<paragraph id="8BAE36B8A72CF056FCEEFF5BB935FF08" blockId="1.[792,1446,140,1496]" box="[792,867,140,162]" pageId="1" pageNumber="12561">
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<heading id="D0E681D4A72CF056FCEEFF5BB935FF08" bold="true" box="[792,867,140,162]" fontSize="9" level="2" pageId="1" pageNumber="12561" reason="0">
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<emphasis id="B965EAAAA72CF056FCEEFF5BB935FF08" bold="true" box="[792,867,140,162]" pageId="1" pageNumber="12561">Results</emphasis>
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</heading>
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</paragraph>
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<paragraph id="8BAE36B8A72CF056FCEEFF7BBF13FD53" blockId="1.[792,1446,140,1496]" pageId="1" pageNumber="12561">
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The fused vertebrae are an articulated pair from the midcaudal series, estimated to have been in positions between caudals 24 and 31, of an adult hadrosaur, most likely
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<taxonomicName id="4C114D3BA72CF056FB73FF31BEABFF51" baseAuthorityName="Marsh" baseAuthorityYear="1892" box="[1157,1277,230,251]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="annectens">
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<emphasis id="B965EAAAA72CF056FB73FF31BEABFF51" box="[1157,1277,230,251]" italics="true" pageId="1" pageNumber="12561">E. annectens</emphasis>
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</taxonomicName>
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(
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<figureCitation id="132A2A3DA72CF056FAFCFF32BFC1FF51" box="[1290,1431,229,251]" captionStart="Fig. 1" captionStartId="1.[102,130,803,820]" captionTargetBox="[104,753,143,777]" captionTargetId="figure@1.[103,755,141,779]" captionTargetPageId="1" captionText="Fig. 1. Depiction of a hadrosaur skeleton showing the position in the tail of the fused vertebrae (A) and a lateral view of the affected vertebrae with the circle indicating the location of the theropod tooth (B)." figureDoi="http://doi.org/10.5281/zenodo.3748451" httpUri="https://zenodo.org/record/3748451/files/figure.png" pageId="1" pageNumber="12561">
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Fig. 1
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<emphasis id="B965EAAAA72CF056FABEFF31BF0EFF51" box="[1352,1368,230,251]" italics="true" pageId="1" pageNumber="12561">A</emphasis>
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and
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<emphasis id="B965EAAAA72CF056FA7EFF31BFC1FF51" box="[1416,1431,230,251]" italics="true" pageId="1" pageNumber="12561">B</emphasis>
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</figureCitation>
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||
). The dimensions of both vertebrae are nearly identical, with a dorsal process 15.5 cm in length and a centrum length of 5.5 cm. Although the specimen shows some abrasion caused by fluvial transport, there does not appear to have been an appreciable loss of exterior bone and it is well preserved overall. Fusion of the vertebrae was accomplished through extensive rugose syndesmophyte-like bone growth that covers the majority of the external surface of the centra (
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<figureCitation id="132A2A3DA72CF056FCE9FE1FB9C5FE77" box="[799,915,456,478]" captionStart="Fig. 2" captionStartId="1.[102,130,1852,1869]" captionTargetBox="[345,1205,1567,1827]" captionTargetId="figure@1.[343,1206,1565,1828]" captionTargetPageId="1" captionText="Fig. 2. CT scans of the fused hadrosaur vertebrae, showing embedded theropod tooth and reactive bone (A–c). Longitudinal slice through the fused vertebrae shows the substantial overgrowth of reactive bone on the outside of the centra, while the articular surfaces remain largely unaffected (A). Two oblique slices through the vertebrae show the embedded theropod tooth in cross-section and the reactive bone that partially surrounds it (B and c). (Scale bars: A, 20 mm and B and c, 10 mm.)" figureDoi="http://doi.org/10.5281/zenodo.3748453" httpUri="https://zenodo.org/record/3748453/files/figure.png" pageId="1" pageNumber="12561">
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||
Fig. 2
|
||
<emphasis id="B965EAAAA72CF056FC9EFE1FB9C5FE77" box="[872,915,456,478]" italics="true" pageId="1" pageNumber="12561">A–C</emphasis>
|
||
</figureCitation>
|
||
). This reactive bone massively overgrew the entire external surface of the two vertebrae. The articular surfaces of the centra appear to have been spared and are completely free of reactive bone. Postmortem breakage on the ventral surface of the centra indicates that the chevron could also have been fused to the centra, but had been subsequently broken off during fluvial transport. A tooth of a large theropod was situated ventrolaterally on the left side, lodged between the two vertebral centra. The tooth consists of the crown only, with no root material present. The rugose bone growth on the centra extends up to and around the embedded tooth, partly enclosing it (
|
||
<figureCitation id="132A2A3DA72CF056FB44FD34BEA0FD53" box="[1202,1270,739,761]" captionStart="Fig. 1" captionStartId="1.[102,130,803,820]" captionTargetBox="[104,753,143,777]" captionTargetId="figure@1.[103,755,141,779]" captionTargetPageId="1" captionText="Fig. 1. Depiction of a hadrosaur skeleton showing the position in the tail of the fused vertebrae (A) and a lateral view of the affected vertebrae with the circle indicating the location of the theropod tooth (B)." figureDoi="http://doi.org/10.5281/zenodo.3748451" httpUri="https://zenodo.org/record/3748451/files/figure.png" pageId="1" pageNumber="12561">Figs. 1</figureCitation>
|
||
and
|
||
<figureCitation id="132A2A3DA72CF056FADCFD34BF6EFD53" box="[1322,1336,739,761]" captionStart="Fig. 2" captionStartId="1.[102,130,1852,1869]" captionTargetBox="[345,1205,1567,1827]" captionTargetId="figure@1.[343,1206,1565,1828]" captionTargetPageId="1" captionText="Fig. 2. CT scans of the fused hadrosaur vertebrae, showing embedded theropod tooth and reactive bone (A–c). Longitudinal slice through the fused vertebrae shows the substantial overgrowth of reactive bone on the outside of the centra, while the articular surfaces remain largely unaffected (A). Two oblique slices through the vertebrae show the embedded theropod tooth in cross-section and the reactive bone that partially surrounds it (B and c). (Scale bars: A, 20 mm and B and c, 10 mm.)" figureDoi="http://doi.org/10.5281/zenodo.3748453" httpUri="https://zenodo.org/record/3748453/files/figure.png" pageId="1" pageNumber="12561">2</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BAE36B8A72CF056FCC6FD29B999FA72" blockId="1.[792,1446,140,1496]" pageId="1" pageNumber="12561">
|
||
The theropod tooth crown is well preserved, with the broken basal portion of the tooth partially exposed (
|
||
<figureCitation id="132A2A3DA72CF056FB3DFCCDBF52FC9A" box="[1227,1284,794,816]" captionStart="Fig. 1" captionStartId="1.[102,130,803,820]" captionTargetBox="[104,753,143,777]" captionTargetId="figure@1.[103,755,141,779]" captionTargetPageId="1" captionText="Fig. 1. Depiction of a hadrosaur skeleton showing the position in the tail of the fused vertebrae (A) and a lateral view of the affected vertebrae with the circle indicating the location of the theropod tooth (B)." figureDoi="http://doi.org/10.5281/zenodo.3748451" httpUri="https://zenodo.org/record/3748451/files/figure.png" pageId="1" pageNumber="12561">Fig. 1</figureCitation>
|
||
). Computerized tomographic (CT) scans revealed the crown height to be 3.75 cm, and visual inspection revealed the crown base length to be 2.35 cm and the crown base width 1.65 cm. Denticles are well preserved on the mesial and distal carinae (
|
||
<figureCitation id="132A2A3DA72CF056FBB4FC5CBE2AFC0B" box="[1090,1148,907,929]" captionStart="Fig. 3" captionStartId="2.[114,142,388,405]" captionTargetBox="[116,767,191,364]" captionTargetId="figure@2.[116,767,141,364]" captionTargetPageId="2" captionText="Fig. 3. Lateral view of embedded tooth crown showing well-developed blood grooves oriented obliquely toward the base and large, chisel-shaped mesial denticles typical of T. rex (60); (Scale bar: 5 mm.)" figureDoi="http://doi.org/10.5281/zenodo.3748455" httpUri="https://zenodo.org/record/3748455/files/figure.png" pageId="1" pageNumber="12561">Fig. 3</figureCitation>
|
||
), and the distal basal denticle (DB) and mesial basal denticle (MB) densities are 16 per cm and 13 per cm, respectively. Comparison of the embedded tooth’ s dimensions and morphometric relationships with the data from the Smith et al. (
|
||
<bibRefCitation id="EF804B49A72CF056FC3DFC2AB9B0FBB9" author="Smith JB & Vann DR & Dodson P" box="[971,998,1021,1043]" journalOrPublisher="Anat Rec A Discov Mol cell Evol Biol" pageId="1" pageNumber="12561" pagination="699 - 736" part="285" refId="ref5509" refString="46. Smith JB, Vann DR, Dodson P (2005) Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth. Anat Rec A Discov Mol cell Evol Biol 285 (2): 699 - 736." title="Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth" type="journal article" year="2005">46</bibRefCitation>
|
||
) study reveals a strong alliance with
|
||
<taxonomicName id="4C114D3BA72CF056FA9AFC29BFF0FBB9" authorityName="Osborn" authorityYear="1905" box="[1388,1446,1022,1043]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72CF056FA9AFC29BFF0FBB9" box="[1388,1446,1022,1043]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="132A2A3DA72CF056FCE9FBCEB908FB85" box="[799,862,1049,1071]" captionStart="Fig. 4" captionStartId="2.[114,142,1827,1844]" captionTargetBox="[356,1216,1024,1801]" captionTargetId="figure@2.[355,1218,1023,1803]" captionTargetPageId="2" captionText="Fig. 4. Graph of tooth dimensions for T. rex, Albertosaurus, and Nanotyrannus, compared with that of the embedded theropod tooth (A and B). When the ICL is plotted against the DCT (A), the three examined taxa are clearly separated, with a large gap existing between T. rex and Albertosaurus, and minor overlap between Albertosaurus and Nanotyrannus. The embedded theropod tooth falls only within the region occupied by T. rex (bold black line in A) and does not overlap with the other two taxa.Graphed ranges of tooth denticles per cm (B) indicates a similar trend, in which the embedded theropod tooth only overlaps T. rex and does not match either Albertosaurus or Nanotyrannus." figureDoi="http://doi.org/10.5281/zenodo.3748457" httpUri="https://zenodo.org/record/3748457/files/figure.png" pageId="1" pageNumber="12561">Fig. 4</figureCitation>
|
||
). The tooth is indistinguishable in morphology, size, and denticle character from known
|
||
<taxonomicName id="4C114D3BA72CF056FBA7FBE1BEDEFBE1" authorityName="Osborn" authorityYear="1905" box="[1105,1160,1078,1099]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72CF056FBA7FBE1BEDEFBE1" box="[1105,1160,1078,1099]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
|
||
</taxonomicName>
|
||
subadults (e.g.,
|
||
<materialsCitation id="3B793CE5A72CF056FADFFBE1BE5CFBC2" ID-GBIF-Occurrence="3396414301" 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="1" pageNumber="12561" specimenCode="LACM 23845">Los Angeles County Museum–23845</materialsCitation>
|
||
and
|
||
<materialsCitation id="3B793CE5A72CF056FBB6FB85BF06FBC2" ID-GBIF-Occurrence="3333873303" box="[1088,1360,1106,1128]" collectionCode="BHI" pageId="1" pageNumber="12561" specimenCode="BHI 6439">Black Hills Institute–6439</materialsCitation>
|
||
). An independent comparison of the ratio of the distance from crown tip (DCT) to the incremental crown length (ICL) for
|
||
<taxonomicName id="4C114D3BA72CF056FAB6FB5CBF2EFB0A" authorityName="Osborn" authorityYear="1905" box="[1344,1400,1163,1184]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72CF056FAB6FB5CBF2EFB0A" box="[1344,1400,1163,1184]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C114D3BA72CF056FCEEFB7FB9F4FB17" authority="Bakker, Currie & Williams, 1988" box="[792,930,1192,1213]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72CF056FCEEFB7FB9F4FB17" box="[792,930,1192,1213]" italics="true" pageId="1" pageNumber="12561">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
, the only two contemporaneous large-bodied and large-toothed theropods (
|
||
<bibRefCitation id="EF804B49A72CF056FBDDFB14BE10FB73" author="Smith JB & Vann DR & Dodson P" box="[1067,1094,1219,1241]" journalOrPublisher="Anat Rec A Discov Mol cell Evol Biol" pageId="1" pageNumber="12561" pagination="699 - 736" part="285" refId="ref5509" refString="46. Smith JB, Vann DR, Dodson P (2005) Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth. Anat Rec A Discov Mol cell Evol Biol 285 (2): 699 - 736." title="Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth" type="journal article" year="2005">46</bibRefCitation>
|
||
–
|
||
<bibRefCitation id="EF804B49A72CF056FBA4FB14BE38FB73" author="Witmer LM & Ridgely RC" box="[1106,1134,1219,1241]" journalOrPublisher="Anat Rec (Hoboken)" pageId="1" pageNumber="12561" pagination="1266 - 1296" part="292" refId="ref5628" refString="49. Witmer LM, Ridgely RC (2009) New insights into the brain, braincase, and ear region of tyrannosaurs (Dinosauria, Theropoda), with implications for sensory organization and behavior. Anat Rec (Hoboken) 292 (9): 1266 - 1296." title="New insights into the brain, braincase, and ear region of tyrannosaurs (Dinosauria, Theropoda), with implications for sensory organization and behavior" type="journal article" year="2009">49</bibRefCitation>
|
||
), with that of the embedded tooth places it firmly within the
|
||
<taxonomicName id="4C114D3BA72CF056FB98FB37BEFFFB5F" authorityName="Osborn" authorityYear="1905" box="[1134,1193,1248,1269]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72CF056FB98FB37BEFFFB5F" box="[1134,1193,1248,1269]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
|
||
</taxonomicName>
|
||
range (
|
||
<figureCitation id="132A2A3DA72CF056FB0AFB08BF6AFB5F" box="[1276,1340,1247,1269]" captionStart="Fig. 3" captionStartId="2.[114,142,388,405]" captionTargetBox="[116,767,191,364]" captionTargetId="figure@2.[116,767,141,364]" captionTargetPageId="2" captionText="Fig. 3. Lateral view of embedded tooth crown showing well-developed blood grooves oriented obliquely toward the base and large, chisel-shaped mesial denticles typical of T. rex (60); (Scale bar: 5 mm.)" figureDoi="http://doi.org/10.5281/zenodo.3748455" httpUri="https://zenodo.org/record/3748455/files/figure.png" pageId="1" pageNumber="12561">Fig. 3</figureCitation>
|
||
). For this study,
|
||
<taxonomicName id="4C114D3BA72CF056FCAFFB2CB989FABB" authorityName="Osborn" authorityYear="1905" box="[857,991,1275,1297]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72CF056FCAFFB2CB989FABB" box="[857,991,1275,1297]" italics="true" pageId="1" pageNumber="12561">Albertosaurus</emphasis>
|
||
</taxonomicName>
|
||
was added as a control. In addition, study of the embedded tooth’ s denticle density indicates that its DB and MB values overlap those of only one animal studied,
|
||
<taxonomicName id="4C114D3BA72CF056FA9FFAE2BFF0FAE0" authorityName="Osborn" authorityYear="1905" box="[1385,1446,1333,1354]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72CF056FA9FFAE2BFF0FAE0" box="[1385,1446,1333,1354]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="132A2A3DA72CF056FCE9FA86B937FACD" box="[799,865,1361,1383]" captionStart="Fig. 3" captionStartId="2.[114,142,388,405]" captionTargetBox="[116,767,191,364]" captionTargetId="figure@2.[116,767,141,364]" captionTargetPageId="2" captionText="Fig. 3. Lateral view of embedded tooth crown showing well-developed blood grooves oriented obliquely toward the base and large, chisel-shaped mesial denticles typical of T. rex (60); (Scale bar: 5 mm.)" figureDoi="http://doi.org/10.5281/zenodo.3748455" httpUri="https://zenodo.org/record/3748455/files/figure.png" pageId="1" pageNumber="12561">Fig. 3</figureCitation>
|
||
). Morphologic and morphometric characters of
|
||
<taxonomicName id="4C114D3BA72CF056FA91FA85B93DFA29" authority="Bakker, Currie & Williams, 1988" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72CF056FA91FA85B93DFA29" italics="true" pageId="1" pageNumber="12561">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
are sufficiently dissimilar from the embedded tooth to exclude it from candidacy for the tooth-producing taxon. Only one animal studied—
|
||
<taxonomicName id="4C114D3BA72CF056FC08FA70BE6AFA16" authorityName="Osborn" authorityYear="1905" box="[1022,1084,1447,1468]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="12561" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72CF056FC08FA70BE6AFA16" box="[1022,1084,1447,1468]" italics="true" pageId="1" pageNumber="12561">T. rex</emphasis>
|
||
</taxonomicName>
|
||
—bears close resemblance to the tooth in question.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<caption id="DF6E6630A72CF056FF90F8EBBBFAF83F" ID-DOI="http://doi.org/10.5281/zenodo.3748453" ID-Zenodo-Dep="3748453" httpUri="https://zenodo.org/record/3748453/files/figure.png" pageId="1" pageNumber="12561" startId="1.[102,130,1852,1869]" targetBox="[345,1205,1567,1827]" targetPageId="1">
|
||
<paragraph id="8BAE36B8A72CF056FF90F8EBBBFAF83F" blockId="1.[102,1445,1852,1941]" pageId="1" pageNumber="12561">
|
||
<emphasis id="B965EAAAA72CF056FF90F8EBBACBF8E7" bold="true" box="[102,157,1852,1869]" pageId="1" pageNumber="12561">Fig. 2.</emphasis>
|
||
CT scans of the fused hadrosaur vertebrae, showing embedded theropod tooth and reactive bone (
|
||
<emphasis id="B965EAAAA72CF056FBDDF8EBBE1BF8E7" box="[1067,1101,1852,1870]" italics="true" pageId="1" pageNumber="12561">A–c</emphasis>
|
||
). Longitudinal slice through the fused vertebrae shows the substantial overgrowth of reactive bone on the outside of the centra, while the articular surfaces remain largely unaffected (
|
||
<emphasis id="B965EAAAA72CF056FA94F883BF39F8CF" box="[1378,1391,1876,1893]" italics="true" pageId="1" pageNumber="12561">A</emphasis>
|
||
). Two oblique slices through the vertebrae show the embedded theropod tooth in cross-section and the reactive bone that partially surrounds it (
|
||
<emphasis id="B965EAAAA72CF056FAE9F8BBBF7CF8D7" box="[1311,1322,1900,1917]" italics="true" pageId="1" pageNumber="12561">B</emphasis>
|
||
and
|
||
<emphasis id="B965EAAAA72CF056FAA1F8BBBF34F8D7" box="[1367,1378,1900,1917]" italics="true" pageId="1" pageNumber="12561">c</emphasis>
|
||
). (Scale bars:
|
||
<emphasis id="B965EAAAA72CF056FF63F853BAF4F83F" box="[149,162,1924,1941]" italics="true" pageId="1" pageNumber="12561">A</emphasis>
|
||
, 20 mm and
|
||
<emphasis id="B965EAAAA72CF056FEE1F853BB74F83F" box="[279,290,1924,1941]" italics="true" pageId="1" pageNumber="12561">B</emphasis>
|
||
and
|
||
<emphasis id="B965EAAAA72CF056FEB9F853BB0CF83F" box="[335,346,1924,1941]" italics="true" pageId="1" pageNumber="12561">c</emphasis>
|
||
, 10 mm.)
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF6E6630A72FF055FF84FE53B803FE6F" ID-DOI="http://doi.org/10.5281/zenodo.3748455" ID-Zenodo-Dep="3748455" httpUri="https://zenodo.org/record/3748455/files/figure.png" pageId="2" pageNumber="12562" startId="2.[114,142,388,405]" targetBox="[116,767,191,364]" targetPageId="2">
|
||
<paragraph id="8BAE36B8A72FF055FF84FE53B803FE6F" blockId="2.[114,768,388,453]" pageId="2" pageNumber="12562">
|
||
<emphasis id="B965EAAAA72FF055FF84FE53BAFCFE3F" bold="true" box="[114,170,388,405]" pageId="2" pageNumber="12562">Fig. 3.</emphasis>
|
||
Lateral view of embedded tooth crown showing well-developed blood grooves oriented obliquely toward the base and large, chisel-shaped mesial denticles typical of
|
||
<taxonomicName id="4C114D3BA72FF055FEA1FE63BBDEFE6F" authorityName="Osborn" authorityYear="1905" box="[343,392,436,453]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72FF055FEA1FE63BBDEFE6F" box="[343,392,436,453]" italics="true" pageId="2" pageNumber="12562">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="EF804B49A72FF055FE65FE63BBFDFE6F" author="Currie PJ & Rigby JK, Jr. & Sloan RE" box="[403,427,436,453]" editor="Carpenter K & Currie PJ" journalOrPublisher="Cambridge Univ Press, Cambridge, MA" pageId="2" pageNumber="12562" pagination="107 - 125" refId="ref5987" refString="60. Currie PJ, Rigby JK, Jr., Sloan RE (1990) Theropod teeth from the Judith River Formation of southern Alberta, Canada. Dinosaur Systematics: Perspectives and Approaches, eds Carpenter K, Currie PJ (Cambridge Univ Press, Cambridge, MA), pp 107 - 125." title="Theropod teeth from the Judith River Formation of southern Alberta, Canada" type="book chapter" volumeTitle="Dinosaur Systematics: Perspectives and Approaches" year="1990">60</bibRefCitation>
|
||
); (Scale bar: 5 mm.)
|
||
</paragraph>
|
||
</caption>
|
||
<subSubSection id="C30B6533A72FF054FF84FE2ABFD1FC86" lastPageId="3" lastPageNumber="12563" pageId="2" pageNumber="12562" type="discussion">
|
||
<paragraph id="8BAE36B8A72FF055FF84FE2ABA89FDB9" blockId="2.[114,769,509,954]" box="[114,223,509,531]" pageId="2" pageNumber="12562">
|
||
<heading id="D0E681D4A72FF055FF84FE2ABA89FDB9" bold="true" box="[114,223,509,531]" fontSize="9" level="2" pageId="2" pageNumber="12562" reason="0">
|
||
<emphasis id="B965EAAAA72FF055FF84FE2ABA89FDB9" bold="true" box="[114,223,509,531]" pageId="2" pageNumber="12562">Discussion</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8BAE36B8A72FF055FF84FDCABB5DFC29" blockId="2.[114,769,509,954]" pageId="2" pageNumber="12562">
|
||
The rugose structures observed on the hadrosaur vertebrae are consistent in morphology with bone growth associated with healing injuries, as observed in modern and extinct animals (
|
||
<figureCitation id="132A2A3DA72FF055FD60FD82B885FDC1" box="[662,723,597,619]" captionStart="Fig. 1" captionStartId="1.[102,130,803,820]" captionTargetBox="[104,753,143,777]" captionTargetId="figure@1.[103,755,141,779]" captionTargetPageId="1" captionText="Fig. 1. Depiction of a hadrosaur skeleton showing the position in the tail of the fused vertebrae (A) and a lateral view of the affected vertebrae with the circle indicating the location of the theropod tooth (B)." figureDoi="http://doi.org/10.5281/zenodo.3748451" httpUri="https://zenodo.org/record/3748451/files/figure.png" pageId="2" pageNumber="12562">Fig. 1</figureCitation>
|
||
). In modern endothermic animals, trauma to bone is followed by signs of insipient bone healing within the first several weeks. The first macroscopic evidence of bone healing in mammals can be recognized 7–10 d after the injury (
|
||
<bibRefCitation id="EF804B49A72FF055FE25FD12BBA6FD71" author="Pritchard JJ & Ruzicka AJ" box="[467,496,709,731]" journalOrPublisher="J Anat" pageId="2" pageNumber="12562" pagination="236 - 261" part="84" refId="ref5679" refString="50. Pritchard JJ, Ruzicka AJ (1950) Comparison of fracture repair in the frog, lizard and rat. J Anat 84 (3): 236 - 261." title="Comparison of fracture repair in the frog, lizard and rat" type="journal article" year="1950">50</bibRefCitation>
|
||
). Healing of reptilian bone is much slower (
|
||
<bibRefCitation id="EF804B49A72FF055FEECFD36BB63FD5D" author="Pritchard JJ & Ruzicka AJ" box="[282,309,737,759]" journalOrPublisher="J Anat" pageId="2" pageNumber="12562" pagination="236 - 261" part="84" refId="ref5679" refString="50. Pritchard JJ, Ruzicka AJ (1950) Comparison of fracture repair in the frog, lizard and rat. J Anat 84 (3): 236 - 261." title="Comparison of fracture repair in the frog, lizard and rat" type="journal article" year="1950">50</bibRefCitation>
|
||
) and takes longer in reptiles (
|
||
<bibRefCitation id="EF804B49A72FF055FD9BFD36B8D1FD5D" author="Pritchard JJ & Ruzicka AJ" box="[621,647,737,759]" journalOrPublisher="J Anat" pageId="2" pageNumber="12562" pagination="236 - 261" part="84" refId="ref5679" refString="50. Pritchard JJ, Ruzicka AJ (1950) Comparison of fracture repair in the frog, lizard and rat. J Anat 84 (3): 236 - 261." title="Comparison of fracture repair in the frog, lizard and rat" type="journal article" year="1950">50</bibRefCitation>
|
||
–
|
||
<bibRefCitation id="EF804B49A72FF055FD65FD36B8F8FD5D" author="Robertson DR" box="[659,686,737,759]" journalOrPublisher="J Exp Zool" pageId="2" pageNumber="12562" pagination="425 - 441" part="172" refId="ref5772" refString="53. Robertson DR (1969) The ultimobranchial body of Rana pipiens. X. Effect of glandular extirpation on fracture healing. J Exp Zool 172 (4): 425 - 441." title="The ultimobranchial body of Rana pipiens. X. Effect of glandular extirpation on fracture healing" type="journal article" year="1969">53</bibRefCitation>
|
||
), significantly delaying opportunity for its recognition on macroscopic examination. The massive bone reaction in this report suggests survival of the injury for a significant period, perhaps even years. The injury does not appear to have contributed to the demise of this hadrosaur.
|
||
</paragraph>
|
||
<paragraph id="8BAE36B8A72FF055FF7CFC5FBE5AFE81" blockId="2.[114,769,509,954]" lastBlockId="2.[804,1458,141,954]" pageId="2" pageNumber="12562">
|
||
The usefulness of individual theropod teeth in determining their taxonomic affinities has been recognized by other researchers (
|
||
<bibRefCitation id="EF804B49A72FF055FCDDFF5AB910FF09" author="Smith JB & Vann DR & Dodson P" box="[811,838,141,163]" journalOrPublisher="Anat Rec A Discov Mol cell Evol Biol" pageId="2" pageNumber="12562" pagination="699 - 736" part="285" refId="ref5509" refString="46. Smith JB, Vann DR, Dodson P (2005) Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth. Anat Rec A Discov Mol cell Evol Biol 285 (2): 699 - 736." title="Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth" type="journal article" year="2005">46</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EF804B49A72FF055FCB9FF5AB93EFF09" author="Currie PJ & Fiorillo AR" box="[847,872,141,163]" journalOrPublisher="J Vert Paleont" pageId="2" pageNumber="12562" pagination="74 - 80" part="14" refId="ref5808" refString="54. Currie PJ, Fiorillo AR (1994) Theropod teeth from the Judith River Formation (Upper Cretaceous) of south-central Montana. J Vert Paleont 14 (1): 74 - 80." title="Theropod teeth from the Judith River Formation (Upper Cretaceous) of south-central Montana" type="journal article" year="1994">54</bibRefCitation>
|
||
) and detailed, quantitative methods of examining theropod teeth have recently been developed and refined (
|
||
<bibRefCitation id="EF804B49A72FF055FAF4FF7FBF48FF14" author="Smith JB & Vann DR & Dodson P" box="[1282,1310,168,190]" journalOrPublisher="Anat Rec A Discov Mol cell Evol Biol" pageId="2" pageNumber="12562" pagination="699 - 736" part="285" refId="ref5509" refString="46. Smith JB, Vann DR, Dodson P (2005) Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth. Anat Rec A Discov Mol cell Evol Biol 285 (2): 699 - 736." title="Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth" type="journal article" year="2005">46</bibRefCitation>
|
||
). The methods developed by Smith et al. (
|
||
<bibRefCitation id="EF804B49A72FF055FBB1FF14BE34FF73" author="Smith JB & Vann DR & Dodson P" box="[1095,1122,195,217]" journalOrPublisher="Anat Rec A Discov Mol cell Evol Biol" pageId="2" pageNumber="12562" pagination="699 - 736" part="285" refId="ref5509" refString="46. Smith JB, Vann DR, Dodson P (2005) Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth. Anat Rec A Discov Mol cell Evol Biol 285 (2): 699 - 736." title="Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth" type="journal article" year="2005">46</bibRefCitation>
|
||
) demonstrate the extent of the discriminating potential of theropod dental characters and enabled identification of isolated
|
||
<taxonomicName id="4C114D3BA72FF055FBACFF2CBEC6FEBA" authorityName="Osborn" authorityYear="1905" box="[1114,1168,251,272]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72FF055FBACFF2CBEC6FEBA" box="[1114,1168,251,272]" italics="true" pageId="2" pageNumber="12562">T. rex</emphasis>
|
||
</taxonomicName>
|
||
tooth crowns with over 96% accuracy in their study.
|
||
</paragraph>
|
||
<paragraph id="8BAE36B8A72FF055FCCAFEE7BF1AFDFD" blockId="2.[804,1458,141,954]" pageId="2" pageNumber="12562">
|
||
The localization of the tyrannosaur tooth in the midcaudal area of the hadrosaur is consistent with that noted in modern pursuit predator attacks (
|
||
<bibRefCitation id="EF804B49A72FF055FC2FFEB1B9A2FED6" author="Stuart CT & Stuart MD" box="[985,1012,358,380]" journalOrPublisher="Struik Publishers, Cape Town, South Africa" pageId="2" pageNumber="12562" refId="ref4793" refString="27. Stuart CT, Stuart MD (1997) The Larger Mammals of Africa (Struik Publishers, Cape Town, South Africa)." title="The Larger Mammals of Africa" type="book" year="1997">27</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EF804B49A72FF055FBF7FEB1BE4AFED6" author="Stander PE" box="[1025,1052,358,380]" journalOrPublisher="can J Zool" pageId="2" pageNumber="12562" pagination="8 - 21" part="70" refId="ref5845" refString="55. Stander PE (1992) Foraging dynamics of lions in a semi-arid environment. can J Zool 70: 8 - 21." title="Foraging dynamics of lions in a semi-arid environment" type="journal article" year="1992">55</bibRefCitation>
|
||
). Kalahari lions have been observed to initially target the hindquarters of the prey animal, in an attempt to immobilize it. This hadrosaur escaped and began to heal around the tooth lodged in its caudal vertebrae, providing evidence of the attack and the identity of the attacker. Such evidence is rare in the fossil record for good reason—prey rarely escapes. As such, it is plausible that many of the purported
|
||
<taxonomicName id="4C114D3BA72FF055FB69FDDCBE81FD8A" authorityName="Osborn" authorityYear="1905" box="[1183,1239,523,544]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72FF055FB69FDDCBE81FD8A" box="[1183,1239,523,544]" italics="true" pageId="2" pageNumber="12562">T. rex</emphasis>
|
||
</taxonomicName>
|
||
bite marks that show no healed bone are actually indicative of successful kills. What is clear is that the
|
||
<taxonomicName id="4C114D3BA72FF055FC3EFD95B9A9FDFD" authorityName="Osborn" authorityYear="1905" box="[968,1023,578,599]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72FF055FC3EFD95B9A9FDFD" box="[968,1023,578,599]" italics="true" pageId="2" pageNumber="12562">T. rex</emphasis>
|
||
</taxonomicName>
|
||
did not finish off this hadrosaur.
|
||
</paragraph>
|
||
<paragraph id="8BAE36B8A72FF054FCCAFD8BBBEFFE8D" blockId="2.[804,1458,141,954]" lastBlockId="3.[102,757,141,534]" lastPageId="3" lastPageNumber="12563" pageId="2" pageNumber="12562">
|
||
The fused vertebrae and embedded tooth are the result of an attack on a live hadrosaur, not the scavenging of a carcass, and represent unequivocal evidence of a predator–prey relationship (
|
||
<bibRefCitation id="EF804B49A72FF055FCDAFD79B96FFD6E" author="Farlow JO & Holtz TR" box="[812,825,686,708]" journalOrPublisher="Paleontological Society Papers" pageId="2" pageNumber="12562" pagination="251 - 266" part="8" refId="ref3943" refString="3. Farlow JO, Holtz TR (2002) The fossil record of predation in dinosaurs. Paleontological Society Papers 8: 251 - 266." title="The fossil record of predation in dinosaurs" type="journal article" year="2002">3</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EF804B49A72FF055FCBFFD79B933FD6E" author="Lebedev OA & Mark-Kurik E & Karataj ‾ ute-Talimaa VN & Luk˘ sevi ˘cs E & Ivanov A" box="[841,869,686,708]" journalOrPublisher="Acta Zool" pageId="2" pageNumber="12562" pagination="344 - 356" part="90 (Supplement 1)" refId="ref4998" refString="33. Lebedev OA, Mark-Kurik E, Karataj ‾ ute-Talimaa VN, Luk ˘ sevi ˘ cs E, Ivanov A (2009) Bite marks as evidence of predation in early vertebrates. Acta Zool 90 (Supplement 1): 344 - 356." title="Bite marks as evidence of predation in early vertebrates" type="journal article" year="2009">33</bibRefCitation>
|
||
). Rugose bone growth encrusting and partially covering the embedded tyrannosaur tooth in the hadrosaur vertebrae indicates that the hadrosaur was unquestionably injured while it was alive and survived the attack long enough to partially heal. The duration of time between the attack and time of death is unknown, however healing rates from modern ectotherms and endotherms suggest that the hadrosaur could have lived from several months to perhaps a number of years after the attack (
|
||
<bibRefCitation id="EF804B49A72FF055FCDDFC5FB910FC34" author="Pritchard JJ & Ruzicka AJ" box="[811,838,904,926]" journalOrPublisher="J Anat" pageId="2" pageNumber="12562" pagination="236 - 261" part="84" refId="ref5679" refString="50. Pritchard JJ, Ruzicka AJ (1950) Comparison of fracture repair in the frog, lizard and rat. J Anat 84 (3): 236 - 261." title="Comparison of fracture repair in the frog, lizard and rat" type="journal article" year="1950">50</bibRefCitation>
|
||
–
|
||
<bibRefCitation id="EF804B49A72FF055FCA5FC5FB926FC34" author="Robertson DR" box="[851,880,904,926]" journalOrPublisher="J Exp Zool" pageId="2" pageNumber="12562" pagination="425 - 441" part="172" refId="ref5772" refString="53. Robertson DR (1969) The ultimobranchial body of Rana pipiens. X. Effect of glandular extirpation on fracture healing. J Exp Zool 172 (4): 425 - 441." title="The ultimobranchial body of Rana pipiens. X. Effect of glandular extirpation on fracture healing" type="journal article" year="1969">53</bibRefCitation>
|
||
). Prey animals that survive attacks by some modern predators, such as lion, hyena, and coyote attacks, frequently can survive long enough to heal (
|
||
<bibRefCitation id="EF804B49A72EF054FE57FF5ABBEAFF09" author="Stuart CT & Stuart MD" box="[417,444,141,163]" journalOrPublisher="Struik Publishers, Cape Town, South Africa" pageId="3" pageNumber="12563" refId="ref4793" refString="27. Stuart CT, Stuart MD (1997) The Larger Mammals of Africa (Struik Publishers, Cape Town, South Africa)." title="The Larger Mammals of Africa" type="book" year="1997">27</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EF804B49A72EF054FE3DFF5ABBB5FF09" author="Stander PE" box="[459,483,141,163]" journalOrPublisher="can J Zool" pageId="3" pageNumber="12563" pagination="8 - 21" part="70" refId="ref5845" refString="55. Stander PE (1992) Foraging dynamics of lions in a semi-arid environment. can J Zool 70: 8 - 21." title="Foraging dynamics of lions in a semi-arid environment" type="journal article" year="1992">55</bibRefCitation>
|
||
–
|
||
<bibRefCitation id="EF804B49A72EF054FE19FF5AB85DFF09" author="Kingdon J" box="[495,523,141,163]" journalOrPublisher="Princeton Univ Press, Princeton" pageId="3" pageNumber="12563" refId="ref5939" refString="58. Kingdon J (1997) The Kingdon Field Guide to African Mammals (Princeton Univ Press, Princeton)." title="The Kingdon Field Guide to African Mammals" type="book" year="1997">58</bibRefCitation>
|
||
). Whereas the healed vertebrae demonstrate in this particular case that the attack on the hadrosaur was not consummated by a kill, they are not a sign that
|
||
<taxonomicName id="4C114D3BA72EF054FF6DFF0ABA8EFF58" authorityName="Osborn" authorityYear="1905" box="[155,216,221,242]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="12563" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72EF054FF6DFF0ABA8EFF58" box="[155,216,221,242]" italics="true" pageId="3" pageNumber="12563">T. rex</emphasis>
|
||
</taxonomicName>
|
||
was a poorly skilled predator. Modern terrestrial predators (e.g., coyotes and lions) fail or abort between 45% and 62% of their attacks (
|
||
<bibRefCitation id="EF804B49A72EF054FEB1FEC6BB34FE8D" author="Bakker RT & Currie PJ & Williams M" box="[327,354,273,295]" journalOrPublisher="Hunteria" pageId="3" pageNumber="12563" pagination="1 - 29" part="1" refId="ref5556" refString="47. Bakker RT, Currie PJ, Williams M (1985) Nanotyrannus, a new genus of pigmy tyrannosaur from the Latest Cretaceous of Montana. Hunteria 1 (5): 1 - 29." title="Nanotyrannus, a new genus of pigmy tyrannosaur from the Latest Cretaceous of Montana" type="journal article" year="1985">47</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EF804B49A72EF054FE98FEC6BBD0FE8D" author="Stander PE" box="[366,390,273,295]" journalOrPublisher="can J Zool" pageId="3" pageNumber="12563" pagination="8 - 21" part="70" refId="ref5845" refString="55. Stander PE (1992) Foraging dynamics of lions in a semi-arid environment. can J Zool 70: 8 - 21." title="Foraging dynamics of lions in a semi-arid environment" type="journal article" year="1992">55</bibRefCitation>
|
||
–
|
||
<bibRefCitation id="EF804B49A72EF054FE64FEC6BBF8FE8D" author="Kingdon J" box="[402,430,273,295]" journalOrPublisher="Princeton Univ Press, Princeton" pageId="3" pageNumber="12563" refId="ref5939" refString="58. Kingdon J (1997) The Kingdon Field Guide to African Mammals (Princeton Univ Press, Princeton)." title="The Kingdon Field Guide to African Mammals" type="book" year="1997">58</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="DF6E6630A72FF055FF84F8F4B8AFF83E" ID-DOI="http://doi.org/10.5281/zenodo.3748457" ID-Zenodo-Dep="3748457" httpUri="https://zenodo.org/record/3748457/files/figure.png" pageId="2" pageNumber="12562" startId="2.[114,142,1827,1844]" targetBox="[356,1216,1024,1801]" targetPageId="2">
|
||
<paragraph id="8BAE36B8A72FF055FF84F8F4B8AFF83E" blockId="2.[114,1458,1827,1940]" pageId="2" pageNumber="12562">
|
||
<emphasis id="B965EAAAA72FF055FF84F8F4BAF0F89E" bold="true" box="[114,166,1827,1844]" pageId="2" pageNumber="12562">Fig. 4.</emphasis>
|
||
Graph of tooth dimensions for
|
||
<taxonomicName id="4C114D3BA72FF055FE32F8F4BBA2F89E" authorityName="Osborn" authorityYear="1905" box="[452,500,1827,1844]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72FF055FE32F8F4BBA2F89E" box="[452,500,1827,1844]" italics="true" pageId="2" pageNumber="12562">T. rex</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C114D3BA72FF055FE08F8F4B820F89E" authorityName="Osborn" authorityYear="1905" box="[510,630,1827,1844]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72FF055FE08F8F4B820F89E" box="[510,630,1827,1844]" italics="true" pageId="2" pageNumber="12562">Albertosaurus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C114D3BA72FF055FD50F8F4B949F89E" authority="Bakker, Currie & Williams, 1988" box="[678,799,1827,1844]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72FF055FD50F8F4B949F89E" box="[678,799,1827,1844]" italics="true" pageId="2" pageNumber="12562">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
, compared with that of the embedded theropod tooth (
|
||
<emphasis id="B965EAAAA72FF055FAF1F8F4BF42F89E" box="[1287,1300,1827,1844]" italics="true" pageId="2" pageNumber="12562">A</emphasis>
|
||
and
|
||
<emphasis id="B965EAAAA72FF055FAB6F8F4BF1DF89E" box="[1344,1355,1827,1844]" italics="true" pageId="2" pageNumber="12562">B</emphasis>
|
||
). When the ICL is plotted against the DCT (
|
||
<emphasis id="B965EAAAA72FF055FE75F8ECBBC6F8E6" box="[387,400,1851,1868]" italics="true" pageId="2" pageNumber="12562">A</emphasis>
|
||
), the three examined taxa are clearly separated, with a large gap existing between
|
||
<taxonomicName id="4C114D3BA72FF055FB83F8ECBEF3F8E6" authorityName="Osborn" authorityYear="1905" box="[1141,1189,1851,1868]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72FF055FB83F8ECBEF3F8E6" box="[1141,1189,1851,1868]" italics="true" pageId="2" pageNumber="12562">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C114D3BA72FF055FB25F8ECBF1DF8E6" authorityName="Osborn" authorityYear="1905" box="[1235,1355,1851,1868]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72FF055FB25F8ECBF1DF8E6" box="[1235,1355,1851,1868]" italics="true" pageId="2" pageNumber="12562">Albertosaurus</emphasis>
|
||
</taxonomicName>
|
||
, and minor overlap between
|
||
<taxonomicName id="4C114D3BA72FF055FEFCF884BBD4F8CE" authorityName="Osborn" authorityYear="1905" box="[266,386,1875,1892]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72FF055FEFCF884BBD4F8CE" box="[266,386,1875,1892]" italics="true" pageId="2" pageNumber="12562">Albertosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C114D3BA72FF055FE58F884B871F8CE" authority="Bakker, Currie & Williams, 1988" box="[430,551,1875,1892]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72FF055FE58F884B871F8CE" box="[430,551,1875,1892]" italics="true" pageId="2" pageNumber="12562">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
. The embedded theropod tooth falls only within the region occupied by
|
||
<taxonomicName id="4C114D3BA72FF055FB54F884BE84F8CE" authorityName="Osborn" authorityYear="1905" box="[1186,1234,1875,1892]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72FF055FB54F884BE84F8CE" box="[1186,1234,1875,1892]" italics="true" pageId="2" pageNumber="12562">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(bold black line in
|
||
<emphasis id="B965EAAAA72FF055FA81F884BFD2F8CE" box="[1399,1412,1875,1892]" italics="true" pageId="2" pageNumber="12562">A</emphasis>
|
||
) and does not overlap with the other two taxa.Graphed ranges of tooth denticles per cm (
|
||
<emphasis id="B965EAAAA72FF055FCBFF8BCB902F8D6" box="[841,852,1899,1916]" italics="true" pageId="2" pageNumber="12562">B</emphasis>
|
||
) indicates a similar trend, in which the embedded theropod tooth only overlaps
|
||
<taxonomicName id="4C114D3BA72FF055FF36F854BAA7F83E" authorityName="Osborn" authorityYear="1905" box="[192,241,1923,1940]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72FF055FF36F854BAA7F83E" box="[192,241,1923,1940]" italics="true" pageId="2" pageNumber="12562">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and does not match either
|
||
<taxonomicName id="4C114D3BA72FF055FE12F854B80AF83E" authorityName="Osborn" authorityYear="1905" box="[484,604,1923,1940]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72FF055FE12F854B80AF83E" box="[484,604,1923,1940]" italics="true" pageId="2" pageNumber="12562">Albertosaurus</emphasis>
|
||
</taxonomicName>
|
||
or
|
||
<taxonomicName id="4C114D3BA72FF055FD8DF854B8A2F83E" authority="Bakker, Currie & Williams, 1988" box="[635,756,1923,1940]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="12562" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72FF055FD8DF854B8A2F83E" box="[635,756,1923,1940]" italics="true" pageId="2" pageNumber="12562">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BAE36B8A72EF054FF88FEFBB894FDBC" blockId="3.[102,757,141,534]" pageId="3" pageNumber="12563">
|
||
Evidence that a
|
||
<taxonomicName id="4C114D3BA72EF054FEC6FEFABB3CFEE8" authorityName="Osborn" authorityYear="1905" box="[304,362,301,322]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="12563" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72EF054FEC6FEFABB3CFEE8" box="[304,362,301,322]" italics="true" pageId="3" pageNumber="12563">T. rex</emphasis>
|
||
</taxonomicName>
|
||
attacked a living hadrosaur indicates predatory behavior, at least in this one instance, and elucidates our view of Western North American Upper Cretaceous paleoecology. Previous assertions that
|
||
<taxonomicName id="4C114D3BA72EF054FEAAFEAABBC3FE38" authorityName="Osborn" authorityYear="1905" box="[348,405,381,402]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="12563" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72EF054FEAAFEAABBC3FE38" box="[348,405,381,402]" italics="true" pageId="3" pageNumber="12563">T. rex</emphasis>
|
||
</taxonomicName>
|
||
was exclusively a scavenger would require us to modify our reconstructions of the paleoecology to accommodate a scavenger of such significant proportions. The fact that
|
||
<taxonomicName id="4C114D3BA72EF054FF65FE1BBA9EFE4B" authorityName="Osborn" authorityYear="1905" box="[147,200,460,481]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="12563" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72EF054FF65FE1BBA9EFE4B" box="[147,200,460,481]" italics="true" pageId="3" pageNumber="12563">T. rex</emphasis>
|
||
</taxonomicName>
|
||
engaged in predatory behavior enables a more plausible paleoecological reconstruction, suggesting an ecology similar to those observed today, such as the African savannah (
|
||
<bibRefCitation id="EF804B49A72EF054FD81FDD7B8C7FDBC" author="Stuart CT & Stuart MD" box="[631,657,512,534]" journalOrPublisher="Struik Publishers, Cape Town, South Africa" pageId="3" pageNumber="12563" refId="ref4793" refString="27. Stuart CT, Stuart MD (1997) The Larger Mammals of Africa (Struik Publishers, Cape Town, South Africa)." title="The Larger Mammals of Africa" type="book" year="1997">27</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EF804B49A72EF054FD6AFDD7B8E1FDBC" author="Kingdon J" box="[668,695,512,534]" journalOrPublisher="Princeton Univ Press, Princeton" pageId="3" pageNumber="12563" refId="ref5939" refString="58. Kingdon J (1997) The Kingdon Field Guide to African Mammals (Princeton Univ Press, Princeton)." title="The Kingdon Field Guide to African Mammals" type="book" year="1997">58</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BAE36B8A72EF054FF90FDE7BA8EFDEC" blockId="3.[102,756,560,906]" box="[102,216,560,582]" pageId="3" pageNumber="12563">
|
||
<heading id="D0E681D4A72EF054FF90FDE7BA8EFDEC" bold="true" box="[102,216,560,582]" fontSize="9" level="2" pageId="3" pageNumber="12563" reason="0">
|
||
<emphasis id="B965EAAAA72EF054FF90FDE7BA8EFDEC" bold="true" box="[102,216,560,582]" pageId="3" pageNumber="12563">Conclusion</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8BAE36B8A72EF054FF90FD86BF61FF71" blockId="3.[102,756,560,906]" lastBlockId="3.[792,1445,141,219]" pageId="3" pageNumber="12563">
|
||
We now have conclusive evidence that
|
||
<taxonomicName id="4C114D3BA72EF054FE0EFD86B867FDCC" authorityName="Osborn" authorityYear="1905" box="[504,561,593,614]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="12563" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72EF054FE0EFD86B867FDCC" box="[504,561,593,614]" italics="true" pageId="3" pageNumber="12563">T. rex</emphasis>
|
||
</taxonomicName>
|
||
indeed engaged in predatory behavior. Whereas previously cited fossil evidence, such as isolated tooth marks, might have been easily misconstrued as the result of scavenging behavior, or predatory behavior carried out by another large theropod, our specimen includes the identities of the prey animal and the attacker and the fact that the prey was alive when attacked. Moreover, the position of the injury—the tail—suggests that
|
||
<taxonomicName id="4C114D3BA72EF054FED1FCDCBB36FC8A" authorityName="Osborn" authorityYear="1905" box="[295,352,779,800]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="12563" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72EF054FED1FCDCBB36FC8A" box="[295,352,779,800]" italics="true" pageId="3" pageNumber="12563">T. rex</emphasis>
|
||
</taxonomicName>
|
||
could possibly have engaged in pursuit predation. As a significant component of the Late Maastrichtian Western North American ecology, perspectives of
|
||
<taxonomicName id="4C114D3BA72EF054FD95FC97B8CCFCFF" authorityName="Osborn" authorityYear="1905" box="[611,666,832,853]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="12563" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72EF054FD95FC97B8CCFCFF" box="[611,666,832,853]" italics="true" pageId="3" pageNumber="12563">T. rex</emphasis>
|
||
</taxonomicName>
|
||
’s role as either a scavenger or predator has a profound effect on our view of the paleoecology because it is such a massive animal. As such, our view of this large theropod as a predator enables us to speculate with more confidence on more accurate paleoecological reconstructions for Laramidia in the Late Cretaceous.
|
||
</paragraph>
|
||
<paragraph id="8BAE36B8A72EF054FCEEFF22BE46FEA1" blockId="3.[792,1447,245,812]" box="[792,1040,245,267]" pageId="3" pageNumber="12563">
|
||
<heading id="D0E681D4A72EF054FCEEFF22BE46FEA1" bold="true" box="[792,1040,245,267]" fontSize="9" level="2" pageId="3" pageNumber="12563" reason="0">
|
||
<emphasis id="B965EAAAA72EF054FCEEFF22BE46FEA1" bold="true" box="[792,1040,245,267]" pageId="3" pageNumber="12563">Materials and Methods</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8BAE36B8A72EF054FCEEFECFBFF1FE5C" blockId="3.[792,1447,245,812]" pageId="3" pageNumber="12563">
|
||
The vertebrae and tooth crown specimen are reposited at the Palm Beach Museum of Natural History (
|
||
<materialsCitation id="3B793CE5A72EF054FBF5FEE5BECEFEE9" ID-GBIF-Occurrence="3333873302" box="[1027,1176,306,323]" collectionCode="PBMNH" pageId="3" pageNumber="12563" specimenCode="PBMNH.P.09.039" specimenCount="1">
|
||
<specimenCode id="DBB79EC3A72EF054FBF5FEE5BECEFEE9" box="[1027,1176,306,323]" pageId="3" pageNumber="12563">PBMNH.P.09.039</specimenCode>
|
||
</materialsCitation>
|
||
).The fossils were stabilized with cyanoacrylate adhesive PaleoBond PB002 and PB100 and prepared using a Comco MB1000 micro air-abrasive unit with sodium bicarbonate powder. Particularly recalcitrant patches of matrix were removed using a PaleoTools MJ5 micro air-scribe. After preparation, the specimen was sealed with polyvinyl acetate stabilizer and later imaged by CT radiography (Siemens SOMATON Sensation 64-slice). CT scanning revealed the tooth’ s orientation within the vertebrae. The tooth crown is well preserved with enamel and denticles intact.
|
||
</paragraph>
|
||
<paragraph id="8BAE36B8A72EF054FCC6FDD7BE60FD6F" blockId="3.[792,1447,245,812]" pageId="3" pageNumber="12563">
|
||
The morphology and morphometric relationships of the embedded theropod tooth in the present study were examined following the methodology of the Smith et al. (
|
||
<bibRefCitation id="EF804B49A72EF054FC33FDE4B98AFDEE" author="Smith JB & Vann DR & Dodson P" box="[965,988,563,580]" journalOrPublisher="Anat Rec A Discov Mol cell Evol Biol" pageId="3" pageNumber="12563" pagination="699 - 736" part="285" refId="ref5509" refString="46. Smith JB, Vann DR, Dodson P (2005) Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth. Anat Rec A Discov Mol cell Evol Biol 285 (2): 699 - 736." title="Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth" type="journal article" year="2005">46</bibRefCitation>
|
||
) study. The measurements were compared with the Smith et al. (
|
||
<bibRefCitation id="EF804B49A72EF054FC7BFD9AB9F2FDF4" author="Smith JB & Vann DR & Dodson P" box="[909,932,589,606]" journalOrPublisher="Anat Rec A Discov Mol cell Evol Biol" pageId="3" pageNumber="12563" pagination="699 - 736" part="285" refId="ref5509" refString="46. Smith JB, Vann DR, Dodson P (2005) Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth. Anat Rec A Discov Mol cell Evol Biol 285 (2): 699 - 736." title="Dental morphology and variation in theropod dinosaurs: Implications for the taxonomic identification of isolated teeth" type="journal article" year="2005">46</bibRefCitation>
|
||
) data, with large theropod taxa documented in the Hell Creek Formation (
|
||
<taxonomicName id="4C114D3BA72EF054FC4EFDB0B9BCFDD2" authorityName="Osborn" authorityYear="1905" box="[952,1002,615,632]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="12563" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B965EAAAA72EF054FC4EFDB0B9BCFDD2" box="[952,1002,615,632]" italics="true" pageId="3" pageNumber="12563">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C114D3BA72EF054FBECFDB0BEC5FDD2" authority="Bakker, Currie & Williams, 1988" box="[1050,1171,615,632]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="12563" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72EF054FBECFDB0BEC5FDD2" box="[1050,1171,615,632]" italics="true" pageId="3" pageNumber="12563">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
), with
|
||
<taxonomicName id="4C114D3BA72EF054FB23FDB0BF1BFDD2" authorityName="Osborn" authorityYear="1905" box="[1237,1357,615,632]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="12563" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72EF054FB23FDB0BF1BFDD2" box="[1237,1357,615,632]" italics="true" pageId="3" pageNumber="12563">Albertosaurus</emphasis>
|
||
</taxonomicName>
|
||
as an exemplar of smaller tyrannosaurids.Initially proposed by Bakker et al. (
|
||
<bibRefCitation id="EF804B49A72EF054FA95FD57BF2AFD3B" author="Bakker RT & Currie PJ & Williams M" box="[1379,1404,640,657]" journalOrPublisher="Hunteria" pageId="3" pageNumber="12563" pagination="1 - 29" part="1" refId="ref5556" refString="47. Bakker RT, Currie PJ, Williams M (1985) Nanotyrannus, a new genus of pigmy tyrannosaur from the Latest Cretaceous of Montana. Hunteria 1 (5): 1 - 29." title="Nanotyrannus, a new genus of pigmy tyrannosaur from the Latest Cretaceous of Montana" type="journal article" year="1985">47</bibRefCitation>
|
||
), the taxonomic status of
|
||
<taxonomicName id="4C114D3BA72EF054FC3AFD4DBE13FD01" authority="Bakker, Currie & Williams, 1988" box="[972,1093,666,683]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="12563" phylum="Chordata" rank="genus">
|
||
<emphasis id="B965EAAAA72EF054FC3AFD4DBE13FD01" box="[972,1093,666,683]" italics="true" pageId="3" pageNumber="12563">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
has been questioned (
|
||
<bibRefCitation id="EF804B49A72EF054FAF8FD4DBF73FD01" author="Carr TD" box="[1294,1317,666,683]" journalOrPublisher="J Vert Paleont" pageId="3" pageNumber="12563" pagination="497 - 520" part="19" refId="ref5961" refString="59. Carr TD (1999) Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). J Vert Paleont 19: 497 - 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">59</bibRefCitation>
|
||
) but is considered a valid taxon for this study.
|
||
</paragraph>
|
||
<paragraph id="8BAE36B8A72EF054FCC6FD19BFD1FC86" blockId="3.[792,1447,245,812]" pageId="3" pageNumber="12563">
|
||
The ratio of two additional sets of measurements for the embedded tooth, DC and ICL, was also examined and compared with the theropod taxa listed above. The DCT is defined in 1cm increments, and the ICL is defined as the specific crown length observed at each of the DCT increments (
|
||
<figureCitation id="132A2A3DA72EF054FAC8FCCCBF2AFC86" box="[1342,1404,795,812]" captionStart="Fig. 1" captionStartId="1.[102,130,803,820]" captionTargetBox="[104,753,143,777]" captionTargetId="figure@1.[103,755,141,779]" captionTargetPageId="1" captionText="Fig. 1. Depiction of a hadrosaur skeleton showing the position in the tail of the fused vertebrae (A) and a lateral view of the affected vertebrae with the circle indicating the location of the theropod tooth (B)." figureDoi="http://doi.org/10.5281/zenodo.3748451" httpUri="https://zenodo.org/record/3748451/files/figure.png" pageId="3" pageNumber="12563">
|
||
Fig. 1
|
||
<emphasis id="B965EAAAA72EF054FA99FCCCBF2AFC86" box="[1391,1404,795,812]" italics="true" pageId="3" pageNumber="12563">A</emphasis>
|
||
</figureCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |