treatments-xml/data/03/E4/77/03E47701FF834210FCACBD8994F3FB9C.xml

<document ID-DOI="10.1371/journal.pone.0013419" ID-GBIF-Dataset="dcb72bb7-896e-44b1-bd40-21f311643008" ID-PMC="PMC2955550" ID-PubMed="20976177" ID-Zenodo-Dep="3898996" approvalRequired="1" approvalRequired_for_treatments="1" checkinTime="1592395800608" checkinUser="jeremy" docAuthor="Nicholas R. Longrich, John R. Horner, Gregory M. Erickson &amp; Philip J. Currie" docDate="2010" docId="03E47701FF834210FCACBD8994F3FB9C" docLanguage="en" docName="Longrichetal2010.PDF.imf" docOrigin="PLoS ONE 5" docStyle="DocumentStyle{}" docTitle="Tyrannosaurus rex Osborn 1905" docType="treatment" docVersion="14" lastPageNumber="5" masterDocId="FFDD0F79FF834214FFCCB9659003FFB8" masterDocTitle="Cannibalism in Tyrannosaurus rex" masterLastPageNumber="e 13419" masterPageNumber="e 13419" pageNumber="1" updateTime="1668135496986" updateUser="ExternalLinkService">
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<mods:title>Cannibalism in Tyrannosaurus rex</mods:title>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Nicholas R. Longrich</mods:namePart>
<mods:affiliation>Department of Geology and Geophysics, Yale University, New Haven, Connecticut, United States of America</mods:affiliation>
<mods:nameIdentifier type="email">nicholas.longrich@yale.edu</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>John R. Horner</mods:namePart>
<mods:affiliation>Museum of the Rockies, Montana State University, Bozeman, Montana, United States of America</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Gregory M. Erickson</mods:namePart>
<mods:affiliation>Department of Biological Science, Florida State University, Tallahassee, Florida, United States of America</mods:affiliation>
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<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Philip J. Currie</mods:namePart>
<mods:affiliation>Department of Biological Sciences, University of Alberta, Edmonton, Alberta, Canada</mods:affiliation>
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<mods:title>PLoS ONE</mods:title>
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<mods:date>2010</mods:date>
<mods:detail type="pubDate">
<mods:number>2010-10-15</mods:number>
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<mods:detail type="volume">
<mods:number>5</mods:number>
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<mods:start>e 13419</mods:start>
<mods:end>e 13419</mods:end>
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<mods:identifier type="DOI">10.1371/journal.pone.0013419</mods:identifier>
<mods:identifier type="GBIF-Dataset">dcb72bb7-896e-44b1-bd40-21f311643008</mods:identifier>
<mods:identifier type="PMC">PMC2955550</mods:identifier>
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Fossils were examined at the American Museum of Natural History (AMNH), the Carnegie Museum of Natural History (CM); the Museum of the Rockies (MOR), the Canadian Museum of Nature (NMC), the Royal Saskatchewan Museum (RSM), the Royal Tyrrell Museum of Palaeontology, (TMP), the University of California Museum of Paleontology (UCMP), and the Yale Peabody Museum (YPM). These include the 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1310,1368,1431,1453]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
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. 
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specimens described here (
</paragraph>
</subSubSection>
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<paragraph blockId="0.[840,1478,1260,1566]" pageId="0" pageNumber="1">
<materialsCitation ID-GBIF-Occurrence="3396409320" box="[994,1146,1459,1481]" collectionCode="UCMP" httpUri="https://ucmpdb.berkeley.edu/cgi/ucmp_query2?admin=&amp;query_src=ucmp_index&amp;table=ucmp2&amp;spec_id=V137538&amp;one=T " pageId="0" pageNumber="1" specimenCode="UCMP 137538">UCMP 137538</materialsCitation>
, 
<materialsCitation ID-GBIF-Occurrence="3396409318" box="[1156,1260,1459,1481]" collectionCode="MOR" pageId="0" pageNumber="1" specimenCode="MOR 920">MOR 902</materialsCitation>
; 
<materialsCitation ID-GBIF-Occurrence="3396409313" box="[1271,1388,1459,1481]" collectionCode="MOR" pageId="0" pageNumber="1" specimenCode="MOR 1126">MOR 1126</materialsCitation>
; 
<materialsCitation ID-GBIF-Occurrence="3396409303" box="[1398,1445,1459,1481]" collectionCode="MOR" pageId="0" pageNumber="1" specimenCode="MOR 1602">1602</materialsCitation>
) as well as all 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[943,996,1487,1509]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
<emphasis box="[943,959,1487,1508]" italics="true" pageId="0" pageNumber="1">T</emphasis>
. 
<emphasis box="[972,996,1487,1508]" italics="true" pageId="0" pageNumber="1">rex</emphasis>
</taxonomicName>
specimens in the CM, RSM, TMP, UCMP, and YPM collections, although in some cases, not all elements of a skeleton were accessible for study.
</paragraph>
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<subSubSection pageId="0" pageNumber="1" type="description">
<paragraph blockId="0.[840,931,1603,1627]" box="[840,931,1603,1627]" pageId="0" pageNumber="1">
<heading bold="true" box="[840,931,1603,1627]" fontSize="10" level="1" pageId="0" pageNumber="1" reason="0">
<emphasis bold="true" box="[840,931,1603,1627]" pageId="0" pageNumber="1">Results</emphasis>
</heading>
</paragraph>
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Including previously described specimens, a total of 17 specimens are identified as bearing tooth marks made by 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[840,963,1707,1728]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">
<emphasis box="[840,963,1707,1728]" italics="true" pageId="0" pageNumber="1">Tyrannosaurus</emphasis>
</taxonomicName>
(
<figureCitation box="[977,1059,1707,1729]" captionStart="Figure 1" captionStartId="2.[155,220,1634,1653]" captionTargetBox="[160,1000,167,1595]" captionTargetId="figure@2.[155,1005,162,1600]" captionTargetPageId="2" captionText="Figure 1. Tooth marks made by Tyrannosaurus rex. Aı hadrosaurid metatarsal (UCMP uncatalogued) and closeup of tooth marks on distal articular surface. Bı fragment of hadrosaurid pubis (CM 105) showing tooth marks on prepubic process. Cı ceratopsid? frill element (TMP 1998.102.2) showing tooth mark. Dı Triceratops right squamosal (YPM 53263) showing tooth marks on edge. doi:10.1371/journal.pone.0013419.g001" figureDoi="http://doi.org/10.5281/zenodo.3898998" httpUri="https://zenodo.org/record/3898998/files/figure.png" pageId="0" pageNumber="1">Figure 1</figureCitation>
; 
<tableCitation box="[1073,1148,1707,1729]" captionStart="Table 1" captionStartId="1.[171,235,174,195]" captionTargetBox="[171,1478,286,942]" captionTargetId="graphics@1.[171,1478,604,660]" captionTargetPageId="1" captionText="Table 1. Specimens showing tooth marks that are attributable to Tyrannosaurus rexı including previously described specimens (15) and specimens previously unidentified or unpublished (asterisk)." httpUri="http://table.plazi.org/id/DF32969FFF824215FF67B9CB9368FF67" pageId="0" pageNumber="1" tableUuid="DF32969FFF824215FF67B9CB9368FF67">Table 1</tableCitation>
). These traces consist of deep U- and V-shaped gouges and shallower scores. None of the traces described here resemble the puncture marks found on a pelvis of 
<taxonomicName authorityName="Marsh" authorityYear="1889" box="[840,932,1792,1813]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">
<emphasis box="[840,932,1792,1813]" italics="true" pageId="0" pageNumber="1">Triceratops</emphasis>
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, but they closely resemble the furrowed ‘puncture and pull’ traces that have previously been attributed to 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1341,1394,1820,1842]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
<emphasis box="[1341,1357,1820,1841]" italics="true" pageId="0" pageNumber="1">T</emphasis>
. 
<emphasis box="[1370,1394,1820,1841]" italics="true" pageId="0" pageNumber="1">rex</emphasis>
</taxonomicName>
[10,15].
</paragraph>
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<paragraph blockId="0.[840,1479,1650,1956]" lastBlockId="1.[155,793,1030,1764]" lastPageId="1" lastPageNumber="2" pageId="0" pageNumber="1">
Of these sixteen specimens, four represent 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1272,1395,1849,1870]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">
<emphasis box="[1272,1395,1849,1870]" italics="true" pageId="0" pageNumber="1">Tyrannosaurus</emphasis>
</taxonomicName>
[
<figureCitation box="[1407,1467,1849,1871]" captionStart="Figure 2" captionStartId="3.[155,220,1343,1362]" captionTargetBox="[160,1001,167,1304]" captionTargetId="figure@3.[155,1006,162,1309]" captionTargetPageId="3" captionText="Figure 2. Tyrannosaurus rex bones bearing tooth marks made by Tyrannosaurus rex. A1ı A2: UCMP 137538ı pedal phalanx in dorsal view. B1ı B2: Pedal phalanxı MOR 1126ı dorsal view. C1ı C2ı Humerus of MOR 902 in caudal view. D1ı D2 metatarsal III of T. rex MOR 1602ı medial view. doi:10.1371/journal.pone.0013419.g002" figureDoi="http://doi.org/10.5281/zenodo.3899000" httpUri="https://zenodo.org/record/3899000/files/figure.png" pageId="0" pageNumber="1">Fig. 2</figureCitation>
]. The first is 
<materialsCitation ID-GBIF-Occurrence="3396409316" box="[959,1114,1877,1899]" collectionCode="UCMP" httpUri="https://ucmpdb.berkeley.edu/cgi/ucmp_query2?admin=&amp;query_src=ucmp_index&amp;table=ucmp2&amp;spec_id=V137538&amp;one=T " pageId="0" pageNumber="1" specimenCode="UCMP 137538">UCMP 137538</materialsCitation>
, a large (13 cm long) pedal phalanx found in isolation (
<figureCitation box="[1029,1104,1906,1928]" captionStart="Figure 2" captionStartId="3.[155,220,1343,1362]" captionTargetBox="[160,1001,167,1304]" captionTargetId="figure@3.[155,1006,162,1309]" captionTargetPageId="3" captionText="Figure 2. Tyrannosaurus rex bones bearing tooth marks made by Tyrannosaurus rex. A1ı A2: UCMP 137538ı pedal phalanx in dorsal view. B1ı B2: Pedal phalanxı MOR 1126ı dorsal view. C1ı C2ı Humerus of MOR 902 in caudal view. D1ı D2 metatarsal III of T. rex MOR 1602ı medial view. doi:10.1371/journal.pone.0013419.g002" figureDoi="http://doi.org/10.5281/zenodo.3899000" httpUri="https://zenodo.org/record/3899000/files/figure.png" pageId="0" pageNumber="1">Fig. 2A</figureCitation>
). It is identified as a theropod by the gynglymous articular surfaces and deep collateral ligament pits, and is referred to 
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<emphasis box="[328,451,1030,1052]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus</emphasis>
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on the basis of its large size, robust construction, and provenance. Comparisons with 
<materialsCitation ID-GBIF-Occurrence="3396409305" collectionCode="FMNH" pageId="1" pageNumber="2" specimenCode="FMNH PR 2081">FMNH PR 2081</materialsCitation>
[
<bibRefCitation author="Brochu CA" box="[218,232,1087,1109]" journalOrPublisher="Journal of Vertebrate Paleontology, Memoirs" pageId="1" pageNumber="2" pagination="1 - 138" part="7" refId="ref3850" refString="3. Brochu CA (2002) Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. Journal of Vertebrate Paleontology, Memoirs 7: 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="2002">3</bibRefCitation>
] show that the bone is a left pedal phalanx IV-2 from a large, adult animal. The proximal end bears four gouges dorsally, and one ventrally, oriented at an oblique angle relative to the axis of the bone. The largest tooth mark is 25 mm long and 7 mm wide.
</paragraph>
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<emphasis bold="true" box="[171,263,174,195]" pageId="1" pageNumber="2">Table 1.</emphasis>
Specimens showing tooth marks that are attributable to 
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<emphasis box="[855,1035,174,195]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus rex</emphasis>
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ı including previously described specimens (15) and specimens previously unidentified or unpublished (asterisk).
</paragraph>
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<emphasis bold="true" box="[171,226,286,303]" pageId="1" pageNumber="2">Taxon</emphasis>
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<emphasis bold="true" box="[471,715,286,303]" pageId="1" pageNumber="2">Accession/Locality Number</emphasis>
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<emphasis bold="true" box="[864,1065,286,303]" pageId="1" pageNumber="2">Toothmarked element</emphasis>
</th>
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<emphasis bold="true" box="[1138,1243,286,303]" pageId="1" pageNumber="2">Provenance</emphasis>
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<th box="[171,372,326,343]" gridcol="0" gridrow="1" pageId="1" pageNumber="2">
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[171,310,326,343]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">
<emphasis box="[171,310,326,343]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus rex</emphasis>
</taxonomicName>
</th>
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<materialsCitation ID-GBIF-Occurrence="3396409319" box="[471,584,326,343]" collectionCode="UCMP" httpUri="https://ucmpdb.berkeley.edu/cgi/ucmp_query2?admin=&amp;query_src=ucmp_index&amp;table=ucmp2&amp;spec_id=V137538&amp;one=T " pageId="1" pageNumber="2" specimenCode="UCMP 137538">UCMP 137538</materialsCitation>
*
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<td box="[864,1065,326,343]" gridcol="2" gridrow="1" pageId="1" pageNumber="2">pedal phalanx</td>
<td box="[1138,1478,326,343]" gridcol="3" gridrow="1" pageId="1" pageNumber="2">Hell Creek Fm.ı Montanaı late Maastrichtian</td>
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<th box="[171,372,359,376]" gridcol="0" gridrow="2" pageId="1" pageNumber="2">
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[171,310,359,376]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">
<emphasis box="[171,310,359,376]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus rex</emphasis>
</taxonomicName>
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<materialsCitation ID-GBIF-Occurrence="3396409308" box="[471,556,359,376]" collectionCode="MOR" pageId="1" pageNumber="2" specimenCode="MOR 1126">MOR 1126</materialsCitation>
*
</td>
<td box="[864,1065,359,376]" gridcol="2" gridrow="2" pageId="1" pageNumber="2">skeleton</td>
<td box="[1138,1478,359,376]" gridcol="3" gridrow="2" pageId="1" pageNumber="2">Hell Creek Fm.ı Montana</td>
</tr>
<tr box="[171,1478,393,410]" gridrow="3" pageId="1" pageNumber="2">
<th box="[171,372,393,410]" gridcol="0" gridrow="3" pageId="1" pageNumber="2">
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[171,310,393,410]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">
<emphasis box="[171,310,393,410]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus rex</emphasis>
</taxonomicName>
</th>
<td box="[471,715,393,410]" gridcol="1" gridrow="3" pageId="1" pageNumber="2">
<materialsCitation ID-GBIF-Occurrence="3396409309" box="[471,546,393,410]" collectionCode="MOR" pageId="1" pageNumber="2" specimenCode="MOR 920">MOR 920</materialsCitation>
*
</td>
<td box="[864,1065,393,410]" gridcol="2" gridrow="3" pageId="1" pageNumber="2">skeleton</td>
<td box="[1138,1478,393,410]" gridcol="3" gridrow="3" pageId="1" pageNumber="2">Hell Creek Fm.ı Montanaı late Maastrichtian</td>
</tr>
<tr box="[171,1478,426,443]" gridrow="4" pageId="1" pageNumber="2">
<th box="[171,372,426,443]" gridcol="0" gridrow="4" pageId="1" pageNumber="2">
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[171,310,426,443]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">
<emphasis box="[171,310,426,443]" italics="true" pageId="1" pageNumber="2">
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[171,282,426,443]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
rex
</emphasis>
</taxonomicName>
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<materialsCitation ID-GBIF-Occurrence="3396409301" box="[471,556,426,443]" collectionCode="MOR" pageId="1" pageNumber="2" specimenCode="MOR 1602">MOR 1602</materialsCitation>
*
</td>
<td box="[864,1065,426,443]" gridcol="2" gridrow="4" pageId="1" pageNumber="2">metatarsal III</td>
<td box="[1138,1478,426,443]" gridcol="3" gridrow="4" pageId="1" pageNumber="2">Hell Creek Fm.ı Montanaı late Maastrichtian</td>
</tr>
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<th box="[171,372,459,476]" gridcol="0" gridrow="5" pageId="1" pageNumber="2">
<taxonomicName authorityName="Marsh" authorityYear="1889" box="[171,254,459,476]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis box="[171,254,459,476]" italics="true" pageId="1" pageNumber="2">Triceratops</emphasis>
</taxonomicName>
sp.
</th>
<td box="[471,715,459,476]" gridcol="1" gridrow="5" pageId="1" pageNumber="2">YPM 53263*</td>
<td box="[864,1065,459,476]" gridcol="2" gridrow="5" pageId="1" pageNumber="2">squamosal</td>
<td box="[1138,1478,459,476]" gridcol="3" gridrow="5" pageId="1" pageNumber="2">Lance Fm. Wyomingı late Maastrichtian</td>
</tr>
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<th box="[171,372,492,509]" gridcol="0" gridrow="6" pageId="1" pageNumber="2">
<taxonomicName authorityName="Marsh" authorityYear="1888" box="[171,273,492,509]" class="Reptilia" family="Ceratopsidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Ceratopsidae</taxonomicName>
indet.
</th>
<td box="[471,715,492,509]" gridcol="1" gridrow="6" pageId="1" pageNumber="2">TMP 1998.102.0005*</td>
<td box="[864,1065,492,509]" gridcol="2" gridrow="6" pageId="1" pageNumber="2">frill fragment</td>
<td box="[1138,1478,492,509]" gridcol="3" gridrow="6" pageId="1" pageNumber="2">Scollard Fm.ı Albertaı late Maastrichtian</td>
</tr>
<tr box="[171,1478,525,542]" gridrow="7" pageId="1" pageNumber="2">
<th box="[171,372,525,542]" gridcol="0" gridrow="7" pageId="1" pageNumber="2">
<taxonomicName authorityName="Marsh" authorityYear="1888" box="[171,273,525,542]" class="Reptilia" family="Ceratopsidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Ceratopsidae</taxonomicName>
indet.
</th>
<td box="[471,715,525,542]" gridcol="1" gridrow="7" pageId="1" pageNumber="2">MOR 799</td>
<td box="[864,1065,525,542]" gridcol="2" gridrow="7" pageId="1" pageNumber="2">pelvis</td>
<td box="[1138,1478,525,542]" gridcol="3" gridrow="7" pageId="1" pageNumber="2">Hell Creek Fm.ı Montanaı late Maastrichtian</td>
</tr>
<tr box="[171,1478,559,598]" gridrow="8" pageId="1" pageNumber="2">
<th box="[171,372,559,598]" gridcol="0" gridrow="8" pageId="1" pageNumber="2">
<taxonomicName authorityName="Marsh" authorityYear="1888" box="[171,273,559,576]" class="Reptilia" family="Ceratopsidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Ceratopsidae</taxonomicName>
indet.
</th>
<td box="[471,715,559,598]" gridcol="1" gridrow="8" pageId="1" pageNumber="2">NMC 53370*</td>
<td box="[864,1065,559,598]" gridcol="2" gridrow="8" pageId="1" pageNumber="2">ischium</td>
<td box="[1138,1478,559,598]" gridcol="3" gridrow="8" pageId="1" pageNumber="2">Frenchman Fm.ı Saskatchewanı late Maastrichtian</td>
</tr>
<tr box="[171,1478,614,654]" gridrow="9" pageId="1" pageNumber="2">
<th box="[171,372,614,654]" gridcol="0" gridrow="9" pageId="1" pageNumber="2">
<taxonomicName authorityName="Marsh" authorityYear="1888" box="[171,273,614,631]" class="Reptilia" family="Ceratopsidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Ceratopsidae</taxonomicName>
indet.
</th>
<td box="[471,715,614,654]" gridcol="1" gridrow="9" pageId="1" pageNumber="2">UCMP 130385*</td>
<td box="[864,1065,614,654]" gridcol="2" gridrow="9" pageId="1" pageNumber="2">left dentary</td>
<td box="[1138,1478,614,654]" gridcol="3" gridrow="9" pageId="1" pageNumber="2">Hell Creek Formationı Montanaı late Maastrichtian</td>
</tr>
<tr box="[171,1478,670,710]" gridrow="10" pageId="1" pageNumber="2">
<th box="[171,372,670,710]" gridcol="0" gridrow="10" pageId="1" pageNumber="2">
<taxonomicName authorityName="Marsh" authorityYear="1888" box="[171,273,670,687]" class="Reptilia" family="Ceratopsidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Ceratopsidae</taxonomicName>
indet.
</th>
<td box="[471,715,670,710]" gridcol="1" gridrow="10" pageId="1" pageNumber="2">UCMP V86061*</td>
<td box="[864,1065,670,710]" gridcol="2" gridrow="10" pageId="1" pageNumber="2">limb bone fragment</td>
<td box="[1138,1478,670,710]" gridcol="3" gridrow="10" pageId="1" pageNumber="2">Hell Creek Formationı Montanaı late Maastrichtian</td>
</tr>
<tr box="[171,1478,726,743]" gridrow="11" pageId="1" pageNumber="2">
<th box="[171,372,726,743]" gridcol="0" gridrow="11" pageId="1" pageNumber="2">
<taxonomicName baseAuthorityName="Marsh" baseAuthorityYear="1892" box="[171,372,726,743]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="annectens">
<emphasis box="[171,372,726,743]" italics="true" pageId="1" pageNumber="2">Edmontosaurus annectens</emphasis>
</taxonomicName>
</th>
<td box="[471,715,726,743]" gridcol="1" gridrow="11" pageId="1" pageNumber="2">AMNH 5041*</td>
<td box="[864,1065,726,743]" gridcol="2" gridrow="11" pageId="1" pageNumber="2">dentary</td>
<td box="[1138,1478,726,743]" gridcol="3" gridrow="11" pageId="1" pageNumber="2">Hell Creek Fm.ı Montanaı late Maastrichtian</td>
</tr>
<tr box="[171,1478,759,776]" gridrow="12" pageId="1" pageNumber="2">
<th box="[171,372,759,776]" gridcol="0" gridrow="12" pageId="1" pageNumber="2">
<taxonomicName baseAuthorityName="Cope" baseAuthorityYear="1869" box="[171,284,759,776]" class="Reptilia" family="Hadrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Hadrosauridae</taxonomicName>
</th>
<td box="[471,715,759,776]" gridcol="1" gridrow="12" pageId="1" pageNumber="2">UCMP 140601</td>
<td box="[864,1065,759,776]" gridcol="2" gridrow="12" pageId="1" pageNumber="2">pedal phalanx</td>
<td box="[1138,1478,759,776]" gridcol="3" gridrow="12" pageId="1" pageNumber="2">Hell Creek Fm.ı Montanaı late Maastrichtian</td>
</tr>
<tr box="[171,1478,792,809]" gridrow="13" pageId="1" pageNumber="2">
<th box="[171,372,792,809]" gridcol="0" gridrow="13" pageId="1" pageNumber="2">
<taxonomicName baseAuthorityName="Cope" baseAuthorityYear="1869" box="[171,284,792,809]" class="Reptilia" family="Hadrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Hadrosauridae</taxonomicName>
</th>
<td box="[471,715,792,809]" gridcol="1" gridrow="13" pageId="1" pageNumber="2">UCMP uncatalogued*</td>
<td box="[864,1065,792,809]" gridcol="2" gridrow="13" pageId="1" pageNumber="2">metatarsal</td>
<td box="[1138,1478,792,809]" gridcol="3" gridrow="13" pageId="1" pageNumber="2">Hell Creek Fm.ı Montanaı late Maastrichtian</td>
</tr>
<tr box="[171,1478,826,843]" gridrow="14" pageId="1" pageNumber="2">
<th box="[171,372,826,843]" gridcol="0" gridrow="14" pageId="1" pageNumber="2">
<taxonomicName baseAuthorityName="Cope" baseAuthorityYear="1869" box="[171,284,826,843]" class="Reptilia" family="Hadrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Hadrosauridae</taxonomicName>
</th>
<td box="[471,715,826,843]" gridcol="1" gridrow="14" pageId="1" pageNumber="2">CM 105*</td>
<td box="[864,1065,826,843]" gridcol="2" gridrow="14" pageId="1" pageNumber="2">pubis</td>
<td box="[1138,1478,826,843]" gridcol="3" gridrow="14" pageId="1" pageNumber="2">Lance Fm.ı Wyomingı late Maastrichtian</td>
</tr>
<tr box="[171,1478,859,876]" gridrow="15" pageId="1" pageNumber="2">
<th box="[171,372,859,876]" gridcol="0" gridrow="15" pageId="1" pageNumber="2">
<taxonomicName baseAuthorityName="Cope" baseAuthorityYear="1869" box="[171,284,859,876]" class="Reptilia" family="Hadrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Hadrosauridae</taxonomicName>
</th>
<td box="[471,715,859,876]" gridcol="1" gridrow="15" pageId="1" pageNumber="2">UCMP V86026*</td>
<td box="[864,1065,859,876]" gridcol="2" gridrow="15" pageId="1" pageNumber="2">caudal vertebra</td>
<td box="[1138,1478,859,876]" gridcol="3" gridrow="15" pageId="1" pageNumber="2">Hell Creek Fm.ı Montanaı late Maastrichtian</td>
</tr>
<tr box="[171,1478,892,909]" gridrow="16" pageId="1" pageNumber="2">
<th box="[171,372,892,909]" gridcol="0" gridrow="16" pageId="1" pageNumber="2">
<taxonomicName authorityName="Gilmore" authorityYear="1913" box="[171,362,892,909]" class="Reptilia" family="Thescelosauridae" genus="Thescelosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="neglectus">
<emphasis box="[171,362,892,909]" italics="true" pageId="1" pageNumber="2">Thescelosaurus neglectus</emphasis>
</taxonomicName>
</th>
<td box="[471,715,892,909]" gridcol="1" gridrow="16" pageId="1" pageNumber="2">MOR 1161*</td>
<td box="[864,1065,892,909]" gridcol="2" gridrow="16" pageId="1" pageNumber="2">Femur</td>
<td box="[1138,1478,892,909]" gridcol="3" gridrow="16" pageId="1" pageNumber="2">Hell Creek Fm.ı Montanaı late Maastrichtian</td>
</tr>
<tr box="[171,1478,925,942]" gridrow="17" pageId="1" pageNumber="2">
<th box="[171,372,925,942]" gridcol="0" gridrow="17" pageId="1" pageNumber="2">
<taxonomicName box="[171,264,925,942]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="1" pageNumber="2" phylum="Chordata" rank="order">Ornithischia</taxonomicName>
indet.
</th>
<td box="[471,715,925,942]" gridcol="1" gridrow="17" pageId="1" pageNumber="2">TMP 1994.125.0102*</td>
<td box="[864,1065,925,942]" gridcol="2" gridrow="17" pageId="1" pageNumber="2">Rib</td>
<td box="[1138,1478,925,942]" gridcol="3" gridrow="17" pageId="1" pageNumber="2">Scollard Fm.ı Albertaı late Maastrichtian</td>
</tr>
</table>
</paragraph>
<paragraph box="[171,475,972,989]" pageId="1" pageNumber="2">
<tableNote box="[171,475,972,989]" pageId="1" pageNumber="2" targetBox="[171,1478,286,942]" targetPageId="1">doi:10.1371/journal.pone.0013419.t001</tableNote>
</paragraph>
<paragraph blockId="1.[155,793,1030,1764]" pageId="1" pageNumber="2">
A second set of 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[343,466,1229,1251]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis box="[343,466,1229,1251]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus</emphasis>
</taxonomicName>
bite marks was found on 
<materialsCitation ID-GBIF-Occurrence="3396409310" collectionCode="MOR" pageId="1" pageNumber="2" specimenCode="MOR 1126">MOR 1126</materialsCitation>
, a partial skeleton of 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[427,550,1258,1280]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis box="[427,550,1258,1280]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus</emphasis>
</taxonomicName>
. On the left foot, pedal phalanx II-2 (
<figureCitation box="[291,360,1286,1308]" captionStart="Figure 2" captionStartId="3.[155,220,1343,1362]" captionTargetBox="[160,1001,167,1304]" captionTargetId="figure@3.[155,1006,162,1309]" captionTargetPageId="3" captionText="Figure 2. Tyrannosaurus rex bones bearing tooth marks made by Tyrannosaurus rex. A1ı A2: UCMP 137538ı pedal phalanx in dorsal view. B1ı B2: Pedal phalanxı MOR 1126ı dorsal view. C1ı C2ı Humerus of MOR 902 in caudal view. D1ı D2 metatarsal III of T. rex MOR 1602ı medial view. doi:10.1371/journal.pone.0013419.g002" figureDoi="http://doi.org/10.5281/zenodo.3899000" httpUri="https://zenodo.org/record/3899000/files/figure.png" pageId="1" pageNumber="2">Fig. 2B</figureCitation>
) bears gouges on the dorsal, lateral, medial, proximal, and distal surfaces. In one tooth mark, long striae left by denticles run through the gouge. The largest tooth mark is on the distal articular surface; it is approximately 70 mm long and 3.5 mm wide. Again, the bone comes from a large adult.
</paragraph>
<paragraph blockId="1.[155,793,1030,1764]" pageId="1" pageNumber="2">
The third example comes from 
<materialsCitation ID-GBIF-Occurrence="3396409311" box="[527,639,1429,1451]" collectionCode="MOR" pageId="1" pageNumber="2" specimenCode="MOR 920">MOR 920</materialsCitation>
, an associated skeleton of an adult 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[347,470,1458,1479]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis box="[347,470,1458,1479]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus</emphasis>
</taxonomicName>
. The left humerus (
<figureCitation box="[659,729,1458,1480]" captionStart="Figure 2" captionStartId="3.[155,220,1343,1362]" captionTargetBox="[160,1001,167,1304]" captionTargetId="figure@3.[155,1006,162,1309]" captionTargetPageId="3" captionText="Figure 2. Tyrannosaurus rex bones bearing tooth marks made by Tyrannosaurus rex. A1ı A2: UCMP 137538ı pedal phalanx in dorsal view. B1ı B2: Pedal phalanxı MOR 1126ı dorsal view. C1ı C2ı Humerus of MOR 902 in caudal view. D1ı D2 metatarsal III of T. rex MOR 1602ı medial view. doi:10.1371/journal.pone.0013419.g002" figureDoi="http://doi.org/10.5281/zenodo.3899000" httpUri="https://zenodo.org/record/3899000/files/figure.png" pageId="1" pageNumber="2">Fig. 2C</figureCitation>
) bears a series of scores on its posterior surface. They are up to 36 mm long and 3 mm wide.
</paragraph>
<paragraph blockId="1.[155,793,1030,1764]" pageId="1" pageNumber="2">
A fourth 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[270,393,1544,1565]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis box="[270,393,1544,1565]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus</emphasis>
</taxonomicName>
specimen bearing conspecific bite marks is 
<materialsCitation ID-GBIF-Occurrence="3396409321" box="[185,312,1572,1594]" collectionCode="MOR" pageId="1" pageNumber="2" specimenCode="MOR 1602">MOR 1602</materialsCitation>
. The specimen consists of an isolated right metatarsal III missing the proximal half of the shaft (
<figureCitation box="[682,758,1600,1622]" captionStart="Figure 2" captionStartId="3.[155,220,1343,1362]" captionTargetBox="[160,1001,167,1304]" captionTargetId="figure@3.[155,1006,162,1309]" captionTargetPageId="3" captionText="Figure 2. Tyrannosaurus rex bones bearing tooth marks made by Tyrannosaurus rex. A1ı A2: UCMP 137538ı pedal phalanx in dorsal view. B1ı B2: Pedal phalanxı MOR 1126ı dorsal view. C1ı C2ı Humerus of MOR 902 in caudal view. D1ı D2 metatarsal III of T. rex MOR 1602ı medial view. doi:10.1371/journal.pone.0013419.g002" figureDoi="http://doi.org/10.5281/zenodo.3899000" httpUri="https://zenodo.org/record/3899000/files/figure.png" pageId="1" pageNumber="2">Fig. 2D</figureCitation>
). It is identified as 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[303,426,1629,1650]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis box="[303,426,1629,1650]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus</emphasis>
</taxonomicName>
by its large size, the triangular shaft, and the splint-like proximal end. It bears two scores on medial surface; the larger is 44 mm by 4 mm. The distal end of the bone is approximately 11 cm across; again the bone is from a large adult.
</paragraph>
</subSubSection>
<subSubSection lastPageId="4" lastPageNumber="5" pageId="1" pageNumber="2" type="discussion">
<paragraph blockId="1.[155,291,1801,1825]" box="[155,291,1801,1825]" pageId="1" pageNumber="2">
<heading bold="true" box="[155,291,1801,1825]" fontSize="10" level="1" pageId="1" pageNumber="2" reason="0">
<emphasis bold="true" box="[155,291,1801,1825]" pageId="1" pageNumber="2">Discussion</emphasis>
</heading>
</paragraph>
<paragraph blockId="1.[155,793,1848,1955]" lastBlockId="1.[840,1479,1030,1956]" pageId="1" pageNumber="2">
The scores and gouges described here closely match the tooth marks of 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[244,367,1877,1898]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
<emphasis box="[244,367,1877,1898]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus</emphasis>
</taxonomicName>
[10,15], and other theropods [
<bibRefCitation author="Rogers RR &amp; Krause DW &amp; Curry Rogers K" box="[663,690,1877,1899]" journalOrPublisher="Nature" pageId="1" pageNumber="2" pagination="515 - 518" part="422" refId="ref4324" refString="16. Rogers RR, Krause DW, Curry Rogers K (2003) Cannibalism in the Madagascan dinosaur Majungatholus atopus. Nature 422: 515 - 518." title="Cannibalism in the Madagascan dinosaur Majungatholus atopus" type="journal article" year="2003">16</bibRefCitation>
] in having a U to V-shaped section that tapers at either end. Given that the tooth marks are relatively deep and narrow, they were probably made by the laterally compressed teeth of the dentary and maxilla, rather than the incisiform premaxillary teeth; the tightly packed premaxillary teeth would also be expected to have left a series of closely spaced scores. An exception is 
<materialsCitation ID-GBIF-Occurrence="3396409314" box="[1212,1365,1115,1137]" collectionCode="UCMP" httpUri="https://ucmpdb.berkeley.edu/cgi/ucmp_query2?admin=&amp;query_src=ucmp_index&amp;table=ucmp2&amp;spec_id=V137538&amp;one=T " pageId="1" pageNumber="2" specimenCode="UCMP 137538">UCMP 137538</materialsCitation>
, where two closely spaced, subparallel gouges are found; these could conceivably have been made by premaxillary teeth.
</paragraph>
<paragraph blockId="1.[840,1479,1030,1956]" pageId="1" pageNumber="2">
It is extremely unlikely that any animal other than a theropod could have produced these traces. Crocodylians would not have produced the tooth marks described here. Whereas the serrated, laterally compressed teeth of theropods carve down into bone (as seen here) the subconical, unserrated teeth of crocodylians produce shallower score marks [
<bibRefCitation author="Njau JK &amp; Blumenschine RJ" box="[1156,1183,1341,1363]" journalOrPublisher="Journal of Human Evolution" pageId="1" pageNumber="2" pagination="142 - 162" part="50" refId="ref4353" refString="17. Njau JK, Blumenschine RJ (2005) A diagnosis of crocodile feeding traces on larger mammal bone, with fossil examples from the Plio-Pleistocene Olduvai Basin, Tanzania. Journal of Human Evolution 50: 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="2005">17</bibRefCitation>
] or deep pits where the tooth punctures the bone [
<bibRefCitation author="Njau JK &amp; Blumenschine RJ" box="[1046,1074,1369,1391]" journalOrPublisher="Journal of Human Evolution" pageId="1" pageNumber="2" pagination="142 - 162" part="50" refId="ref4353" refString="17. Njau JK, Blumenschine RJ (2005) A diagnosis of crocodile feeding traces on larger mammal bone, with fossil examples from the Plio-Pleistocene Olduvai Basin, Tanzania. Journal of Human Evolution 50: 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="2005">17</bibRefCitation>
]; such puncture marks are absent in the bones described here. Some lizards do have ziphodont teeth that can produce tooth marks resembling theropod tooth marks, notably 
<taxonomicName box="[924,1094,1453,1475]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="komodensis">
<emphasis box="[924,1094,1453,1475]" italics="true" pageId="1" pageNumber="2">Varanus komodensis</emphasis>
</taxonomicName>
[
<bibRefCitation author="D'Amore DC &amp; Blumenschine RJ" box="[1112,1140,1454,1476]" journalOrPublisher="Paleobiology" pageId="1" pageNumber="2" pagination="525 - 552" part="35" refId="ref4395" refString="18. D'Amore DC, Blumenschine RJ (2009) Komodo monitor (Varanus komodoensis) feeding behavior and dental function reflected through tooth marks on bone surfaces as the application to ziphodont paleobiology. Paleobiology 35: 525 - 552." title="Komodo monitor (Varanus komodoensis) feeding behavior and dental function reflected through tooth marks on bone surfaces as the application to ziphodont paleobiology" type="journal article" year="2009">18</bibRefCitation>
]. The Hell Creek and Lance do contain a large lizard with ziphodont dentition, 
<emphasis box="[1311,1424,1482,1503]" italics="true" pageId="1" pageNumber="2">Palaeosaniaea</emphasis>
[
<bibRefCitation author="Estes R" box="[1439,1467,1482,1504]" journalOrPublisher="University of California Publications, Department of Geological Sciences" pageId="1" pageNumber="2" pagination="1 - 180" part="49" refId="ref4437" refString="19. Estes R (1964) Fossil vertebrates from the Late Cretaceous Lance Formation, Eastern Wyoming. University of California Publications, Department of Geological Sciences 49: 1 - 180." title="Fossil vertebrates from the Late Cretaceous Lance Formation, Eastern Wyoming" type="journal article" year="1964">19</bibRefCitation>
]. However, even tooth marks made by the large 
<taxonomicName box="[1291,1416,1510,1532]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="komodoensis">
<emphasis box="[1291,1305,1511,1532]" italics="true" pageId="1" pageNumber="2">V</emphasis>
. 
<emphasis box="[1317,1416,1510,1531]" italics="true" pageId="1" pageNumber="2">komodoensis</emphasis>
</taxonomicName>
rarely exceed 1 mm in width [
<bibRefCitation author="D'Amore DC &amp; Blumenschine RJ" box="[1073,1101,1539,1561]" journalOrPublisher="Paleobiology" pageId="1" pageNumber="2" pagination="525 - 552" part="35" refId="ref4395" refString="18. D'Amore DC, Blumenschine RJ (2009) Komodo monitor (Varanus komodoensis) feeding behavior and dental function reflected through tooth marks on bone surfaces as the application to ziphodont paleobiology. Paleobiology 35: 525 - 552." title="Komodo monitor (Varanus komodoensis) feeding behavior and dental function reflected through tooth marks on bone surfaces as the application to ziphodont paleobiology" type="journal article" year="2009">18</bibRefCitation>
]. 
<emphasis box="[1119,1232,1538,1559]" italics="true" pageId="1" pageNumber="2">Palaeosaniaea</emphasis>
was considerably smaller than 
<taxonomicName box="[893,1022,1566,1589]" class="Reptilia" family="Varanidae" genus="Varanus" kingdom="Animalia" order="Squamata" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="komodoensis">
<emphasis box="[893,907,1567,1588]" italics="true" pageId="1" pageNumber="2">V</emphasis>
. 
<emphasis box="[923,1022,1566,1587]" italics="true" pageId="1" pageNumber="2">komodoensis</emphasis>
</taxonomicName>
, probably between 1 and 2 meters long, and therefore too small to have produced the traces described here. Mammals are known to gnaw on dinosaur bone [
<bibRefCitation author="Longrich NR &amp; Ryan MJ" box="[1390,1419,1623,1645]" journalOrPublisher="Palaeontology" pageId="1" pageNumber="2" pagination="703 - 709" part="53" refId="ref4471" refString="20. Longrich NR, Ryan MJ (2010) Mammalian tooth marks on the bones of dinosaurs and other Late Cretaceous vertebrates. Palaeontology 53: 703 - 709." title="Mammalian tooth marks on the bones of dinosaurs and other Late Cretaceous vertebrates" type="journal article" year="2010">20</bibRefCitation>
], but mammalian gnaw traces are far smaller, and consist of closely spaced, paired tooth marks.
</paragraph>
<paragraph blockId="1.[840,1479,1030,1956]" lastBlockId="2.[155,793,1765,1956]" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">
Insects can modify bone, but traces left by dermestid beetles are small and characterized by minute scratches left by the mandibles; termites produce meandering tunnels [
<bibRefCitation author="Britt BB &amp; Scheetz RD &amp; Dangerfield A" box="[1222,1251,1765,1787]" journalOrPublisher="Ichnos" pageId="1" pageNumber="2" pagination="59 - 71" part="15" refId="ref4502" refString="21. Britt BB, Scheetz RD, Dangerfield A (2008) A suite of dermestid beetle traces on dinosaur bone from the Upper Jurassic Morrison Formation, Wyoming, USA. Ichnos 15: 59 - 71." title="A suite of dermestid beetle traces on dinosaur bone from the Upper Jurassic Morrison Formation, Wyoming, USA" type="journal article" year="2008">21</bibRefCitation>
]. The large trace fossil 
<emphasis box="[840,929,1793,1814]" italics="true" pageId="1" pageNumber="2">Cubiculum</emphasis>
[
<bibRefCitation author="Roberts EM &amp; Rogers RR &amp; Foreman BZ" box="[943,972,1793,1815]" journalOrPublisher="Journal of Paleontology" pageId="1" pageNumber="2" pagination="201 - 208" part="81" refId="ref4542" refString="22. Roberts EM, Rogers RR, Foreman BZ (2007) Continental insect borings in dinosaur bone: examples from the Late Cretaceous of Madagascar and Utah. Journal of Paleontology 81: 201 - 208." title="Continental insect borings in dinosaur bone: examples from the Late Cretaceous of Madagascar and Utah" type="journal article" year="2007">22</bibRefCitation>
], which is made by burrowing mayfly nymphs [
<bibRefCitation author="Thenius vE" box="[1438,1467,1793,1815]" journalOrPublisher="Annalen Naturhistorischen Museum Wien" pageId="1" pageNumber="2" pagination="177 - 188" part="82" refId="ref4581" refString="23. Thenius vE (1979) Lebensspuren von ephemeropteren-larven aus dem Jung- Tertiar des wiener beckens. Annalen Naturhistorischen Museum Wien 82: 177 - 188." title="Lebensspuren von ephemeropteren-larven aus dem Jung- Tertiar des wiener beckens" type="journal article" year="1979">23</bibRefCitation>
], (rather than carrion beetles), is common in the Hell Creek and Lance formations (NRL, pers. obs.) but consists of broad channels with a U-shaped section, which do not resemble the traces described here. Neither can these marks be accounted for by nonbiological mechanisms: trample marks are often seen on bones, but these consist of numerous small and closely spaced grooves (NRL, pers. obs.). These traces do not represent tool marks made during excavation, because tools could not penetrate deeply into the fossil without shattering the brittle bone.
</paragraph>
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<paragraph blockId="2.[155,1478,1634,1725]" pageId="2" pageNumber="3">
<emphasis bold="true" box="[155,676,1634,1653]" pageId="2" pageNumber="3">
Figure 1. Tooth marks made by 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[496,670,1634,1653]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
.
</emphasis>
Aı hadrosaurid metatarsal (UCMP uncatalogued) and closeup of tooth marks on distal articular surface. Bı fragment of hadrosaurid pubis (CM 105) showing tooth marks on prepubic process. Cı ceratopsid? frill element (TMP 1998.102.2) showing tooth mark. Dı 
<taxonomicName authorityName="Marsh" authorityYear="1889" box="[376,471,1682,1701]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis box="[376,471,1682,1701]" italics="true" pageId="2" pageNumber="3">Triceratops</emphasis>
</taxonomicName>
right squamosal (YPM 53263) showing tooth marks on edge. doi:10.1371/journal.pone.0013419.g001
</paragraph>
</caption>
<paragraph blockId="2.[155,793,1765,1956]" lastBlockId="3.[155,793,1450,1956]" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3">
It is highly unlikely that a non-tyrannosaurid theropod could have made the bite marks described here. Dromaeosaurids and troodontids are known from the late Maastrichtian of North America, but these are relatively small animals [
<bibRefCitation author="Longrich NR" box="[1323,1353,1765,1787]" editor="Sankey JT &amp; Baszio S" journalOrPublisher="Indiana: University Press" pageId="2" pageNumber="3" pagination="135 - 158" refId="ref4609" refString="24. Longrich NR (2008) Small theropod teeth from the Lance Formation of Wyoming. In: Sankey JT, Baszio S, eds. The Unique Role of Vertebrate Microfossil Assemblages in Paleoecology and Paleobiology:. Indiana: University Press. pp 135 - 158." title="Small theropod teeth from the Lance Formation of Wyoming" type="book chapter" volumeTitle="The Unique Role of Vertebrate Microfossil Assemblages in Paleoecology and Paleobiology" year="2008">24</bibRefCitation>
]. Given that tooth scores made by the much larger dromaeosaurid 
<taxonomicName authorityName="Ostrom" authorityYear="1969" box="[1376,1478,1793,1814]" class="Reptilia" family="Dromaeosauridae" genus="Deinonychus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
<emphasis box="[1376,1478,1793,1814]" italics="true" pageId="2" pageNumber="3">Deinonychus</emphasis>
</taxonomicName>
are just 1 mm wide [
<bibRefCitation author="Gignac PM &amp; Makovicky PJ &amp; Erickson GM &amp; Walsh RP" box="[1049,1077,1821,1843]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="2" pageNumber="3" pagination="1169 - 1177" part="30" refId="ref4659" refString="25. Gignac PM, Makovicky PJ, Erickson GM, Walsh RP (2010) A description of Deinonychus antirrhopus bite marks and estimates of bite force using tooth indentation simulations. Journal of Vertebrate Paleontology 30: 1169 - 1177." title="A description of Deinonychus antirrhopus bite marks and estimates of bite force using tooth indentation simulations" type="journal article" year="2010">25</bibRefCitation>
] the small deinonychosaurs in the fauna could not have made the traces described here. Furthermore, bones bitten by dromaeosaurids are extremely rare, [
<bibRefCitation author="Gignac PM &amp; Makovicky PJ &amp; Erickson GM &amp; Walsh RP" box="[1395,1423,1878,1900]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="2" pageNumber="3" pagination="1169 - 1177" part="30" refId="ref4659" refString="25. Gignac PM, Makovicky PJ, Erickson GM, Walsh RP (2010) A description of Deinonychus antirrhopus bite marks and estimates of bite force using tooth indentation simulations. Journal of Vertebrate Paleontology 30: 1169 - 1177." title="A description of Deinonychus antirrhopus bite marks and estimates of bite force using tooth indentation simulations" type="journal article" year="2010">25</bibRefCitation>
] and dromaeosaurid teeth exhibit little or no wear [
<bibRefCitation author="Longrich NR" box="[1305,1334,1906,1928]" editor="Sankey JT &amp; Baszio S" journalOrPublisher="Indiana: University Press" pageId="2" pageNumber="3" pagination="135 - 158" refId="ref4609" refString="24. Longrich NR (2008) Small theropod teeth from the Lance Formation of Wyoming. In: Sankey JT, Baszio S, eds. The Unique Role of Vertebrate Microfossil Assemblages in Paleoecology and Paleobiology:. Indiana: University Press. pp 135 - 158." title="Small theropod teeth from the Lance Formation of Wyoming" type="book chapter" volumeTitle="The Unique Role of Vertebrate Microfossil Assemblages in Paleoecology and Paleobiology" year="2008">24</bibRefCitation>
], which shows that they avoided biting into bone. In contrast, tyrannosaurids frequently bit in to bone, as demonstrated by spalling of the teeth [
<bibRefCitation author="Schubert BW &amp; Ungar PS" box="[161,190,1478,1500]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="3" pageNumber="4" pagination="93 - 99" part="50" refId="ref4702" refString="26. Schubert BW, Ungar PS (2005) Wear facets and enamel spalling in tyrannosaurid dinosaurs. Acta Palaeontologica Polonica 50: 93 - 99." title="Wear facets and enamel spalling in tyrannosaurid dinosaurs" type="journal article" year="2005">26</bibRefCitation>
], heavy wear on the tooth apices and the carinae [
<bibRefCitation author="Farlow JO &amp; Brinkman D" box="[753,782,1478,1500]" editor="Rosenberg GD &amp; Wolberg DL" journalOrPublisher="Geological Sciences, Univ. of Tennessee" pageId="3" pageNumber="4" pagination="165 - 175" refId="ref4730" refString="27. Farlow JO, Brinkman D (1994) Wear surfaces on the teeth of tyrannosaurs. In: Rosenberg GD, Wolberg DL, eds. The Paleontological Society Special Publication; 1994. Knoxville: Dept. Geological Sciences, Univ. of Tennessee. pp 165 - 175." title="Wear surfaces on the teeth of tyrannosaurs" type="book chapter" volumeTitle="The Paleontological Society Special Publication; 1994. Knoxville: Dept" year="1994">27</bibRefCitation>
], previous identification of bite marks [
<bibRefCitation author="Erickson GM &amp; Olson KH" box="[512,539,1507,1529]" pageId="3" pageNumber="4" pagination="175 - 178" part="16" refId="ref4296" refString="15. Erickson GM, Olson KH (1996) Bite marks attributable to Tyrannosaurus rex: preliminary description and implications. 16: 175 - 178." title="Bite marks attributable to Tyrannosaurus rex: preliminary description and implications" type="journal article" year="1996">15</bibRefCitation>
] and the presence of bone in a tyrannosaur coprolite [
<bibRefCitation author="Chin K &amp; Tokaryk TT &amp; Erickson GM &amp; Calk LC" box="[423,452,1535,1557]" journalOrPublisher="Nature" pageId="3" pageNumber="4" pagination="680 - 682" part="393" refId="ref4783" refString="28. Chin K, Tokaryk TT, Erickson GM, Calk LC (1998) A king-sized theropod coprolite. Nature 393: 680 - 682." title="A king-sized theropod coprolite" type="journal article" year="1998">28</bibRefCitation>
].
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3899000" ID-Zenodo-Dep="3899000" httpUri="https://zenodo.org/record/3899000/files/figure.png" pageId="3" pageNumber="4" startId="3.[155,220,1343,1362]" targetBox="[160,1001,167,1304]" targetPageId="3">
<paragraph blockId="3.[155,1479,1343,1409]" pageId="3" pageNumber="4">
<emphasis bold="true" box="[155,979,1343,1362]" pageId="3" pageNumber="4">
Figure 2. 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[250,421,1343,1362]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
bones bearing tooth marks made by 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[802,973,1343,1362]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
.
</emphasis>
A1ı A2: 
<materialsCitation ID-GBIF-Occurrence="3396409306" box="[1052,1180,1343,1362]" collectionCode="UCMP" httpUri="https://ucmpdb.berkeley.edu/cgi/ucmp_query2?admin=&amp;query_src=ucmp_index&amp;table=ucmp2&amp;spec_id=V137538&amp;one=T " pageId="3" pageNumber="4" specimenCode="UCMP 137538">UCMP 137538</materialsCitation>
ı pedal phalanx in dorsal view. B1ı B2: Pedal phalanxı 
<materialsCitation ID-GBIF-Occurrence="3396409317" box="[326,423,1366,1385]" collectionCode="MOR" pageId="3" pageNumber="4" specimenCode="MOR 1126">MOR 1126</materialsCitation>
ı dorsal view. C1ı C2ı Humerus of 
<materialsCitation ID-GBIF-Occurrence="3396409312" box="[727,810,1366,1385]" collectionCode="MOR" pageId="3" pageNumber="4" specimenCode="MOR 920">MOR 902</materialsCitation>
in caudal view. D1ı D2 metatarsal III of 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1175,1222,1366,1385]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="rex">
<emphasis box="[1175,1185,1366,1385]" italics="true" pageId="3" pageNumber="4">T</emphasis>
. 
<emphasis box="[1196,1222,1366,1385]" italics="true" pageId="3" pageNumber="4">rex</emphasis>
</taxonomicName>
<materialsCitation ID-GBIF-Occurrence="3396409315" box="[1229,1325,1366,1385]" collectionCode="MOR" pageId="3" pageNumber="4" specimenCode="MOR 1602">MOR 1602</materialsCitation>
ı medial view. doi:10.1371/journal.pone.0013419.g002
</paragraph>
</caption>
<paragraph blockId="3.[155,793,1450,1956]" lastBlockId="3.[840,1478,1450,1956]" pageId="3" pageNumber="4">
It is usually impossible to refer tooth marks to a particular species, and here, the traces themselves preserve no distinctive features other than their size. However, 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[557,680,1621,1642]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[557,680,1621,1642]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
</taxonomicName>
is the only large theropod known from the Late Maastrichtian of the Western Interior [
<bibRefCitation author="Carr TD &amp; Williamson TE" box="[250,279,1678,1700]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="3" pageNumber="4" pagination="479 - 523" part="142" refId="ref4259" refString="14. Carr TD, Williamson TE (2004) Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America. Zoological Journal of the Linnean Society 142: 479 - 523." title="Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America" type="journal article" year="2004">14</bibRefCitation>
]. The holotype of ‘‘ 
<taxonomicName baseAuthorityName="Gilmore" baseAuthorityYear="1946" box="[499,698,1677,1699]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="lancensis">
<emphasis box="[499,698,1677,1699]" italics="true" pageId="3" pageNumber="4">Nanotyrannus lancensis</emphasis>
</taxonomicName>
’’ [
<bibRefCitation author="Bakker R &amp; Williams M &amp; Currie PJ" box="[730,758,1678,1700]" journalOrPublisher="Hunteria" pageId="3" pageNumber="4" pagination="1 - 30" part="1" refId="ref4811" refString="29. Bakker R, Williams M, Currie PJ (1988) Nanotyrannus, a new genus of pygmy tyrannosaur, from the latest Cretaceous of Montana. Hunteria 1: 1 - 30." title="Nanotyrannus, a new genus of pygmy tyrannosaur, from the latest Cretaceous of Montana" type="journal article" year="1988">29</bibRefCitation>
] is immature and displays virtually all the features expected for a juvenile 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[242,365,1735,1756]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[242,365,1735,1756]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
</taxonomicName>
[14,30], including a skull with a narrow snout and a broad temporal region, a deep mandible, and an elongate sagittal crest of the frontal [
<bibRefCitation author="Carr TD" box="[587,616,1792,1814]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="3" pageNumber="4" pagination="497 - 520" part="19" refId="ref4847" refString="30. Carr TD (1999) Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology 19: 497 - 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">30</bibRefCitation>
]. No adults of ‘‘ 
<taxonomicName baseAuthorityName="Bakker, Currie &amp; Williams, 1988" baseAuthorityYear="1988" box="[167,281,1820,1841]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[167,281,1820,1841]" italics="true" pageId="3" pageNumber="4">Nanotyrannus</emphasis>
</taxonomicName>
’’ are known, or juveniles of 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[563,617,1820,1842]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="rex">
<emphasis box="[563,579,1820,1841]" italics="true" pageId="3" pageNumber="4">T</emphasis>
. 
<emphasis box="[593,617,1820,1841]" italics="true" pageId="3" pageNumber="4">rex</emphasis>
</taxonomicName>
that clearly differ from ‘‘ 
<taxonomicName baseAuthorityName="Bakker, Currie &amp; Williams, 1988" baseAuthorityYear="1988" box="[221,335,1849,1870]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[221,335,1849,1870]" italics="true" pageId="3" pageNumber="4">Nanotyrannus</emphasis>
</taxonomicName>
’’. Thus, ‘‘ 
<taxonomicName baseAuthorityName="Bakker, Currie &amp; Williams, 1988" baseAuthorityYear="1988" box="[437,551,1849,1870]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[437,551,1849,1870]" italics="true" pageId="3" pageNumber="4">Nanotyrannus</emphasis>
</taxonomicName>
’’ is most parsimoniously considered a juvenile of 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[406,529,1877,1898]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[406,529,1877,1898]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
</taxonomicName>
. ‘‘ 
<taxonomicName baseAuthorityName="Bakker, Currie &amp; Williams, 1988" baseAuthorityYear="1988" box="[557,671,1877,1898]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[557,671,1877,1898]" italics="true" pageId="3" pageNumber="4">Nanotyrannus</emphasis>
</taxonomicName>
’’ does have more maxillary teeth than other specimens of 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[598,650,1905,1927]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="rex">
<emphasis box="[598,614,1905,1926]" italics="true" pageId="3" pageNumber="4">T</emphasis>
. 
<emphasis box="[626,650,1905,1926]" italics="true" pageId="3" pageNumber="4">rex</emphasis>
</taxonomicName>
(fifteen, versus eleven to twelve for other 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[425,481,1934,1956]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="rex">
<emphasis box="[425,441,1934,1955]" italics="true" pageId="3" pageNumber="4">T</emphasis>
. 
<emphasis box="[457,481,1934,1955]" italics="true" pageId="3" pageNumber="4">rex</emphasis>
</taxonomicName>
) [29,30,31] but given that this feature is highly variable within species, and even between the left and right maxillae in a single individual [
<bibRefCitation author="Currie PJ" box="[1252,1280,1478,1500]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="3" pageNumber="4" pagination="191 - 226" part="48" refId="ref4874" refString="31. Currie PJ (2003) Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologica Polonica 48: 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">31</bibRefCitation>
], it is insufficient to warrant the recognition of a separate species. Because there is no compelling evidence for more than one tyrannosaurid in the fauna, then by default, the traces described above can be attributed to 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[970,1093,1592,1613]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[970,1093,1592,1613]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="3.[840,1478,1450,1956]" pageId="3" pageNumber="4">
Most of the traces described here are smaller than previously described 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[941,1064,1650,1671]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[941,1064,1650,1671]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
</taxonomicName>
tooth marks, which are up to 25 mm in width [
<bibRefCitation author="Erickson GM &amp; Olson KH" box="[909,937,1678,1700]" pageId="3" pageNumber="4" pagination="175 - 178" part="16" refId="ref4296" refString="15. Erickson GM, Olson KH (1996) Bite marks attributable to Tyrannosaurus rex: preliminary description and implications. 16: 175 - 178." title="Bite marks attributable to Tyrannosaurus rex: preliminary description and implications" type="journal article" year="1996">15</bibRefCitation>
]. This suggests that they were made by juvenile or subadult 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[897,1020,1706,1727]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis box="[897,1020,1706,1727]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
</taxonomicName>
, although it is also conceivable that they were made by the smaller posterior teeth of a large individual. However, the broad, shallow tooth marks in 
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may have been made by a large individual that was not biting at full force.
</paragraph>
<paragraph blockId="3.[840,1478,1450,1956]" lastBlockId="4.[155,794,163,1385]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
We argue that these traces result from feeding, rather than intraspecific combat. First, these traces would have been difficult to inflict on a live animal. In the case of 
<materialsCitation ID-GBIF-Occurrence="3396409304" box="[1246,1364,1877,1899]" collectionCode="MOR" pageId="3" pageNumber="4" specimenCode="MOR 1126">MOR 1126</materialsCitation>
, bite marks occur on both the proximal and distal ends of the bone and the shaft, suggesting that the bone was bitten two or three times. It seems unlikely that a small 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[446,569,163,185]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[446,569,163,185]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
</taxonomicName>
would be allowed to repeatedly bite a much larger individual several times on a single toe. In the case of the metatarsal, 
<materialsCitation ID-GBIF-Occurrence="3396409307" box="[477,592,219,241]" collectionCode="MOR" pageId="4" pageNumber="5" specimenCode="MOR 1602">MOR 1602</materialsCitation>
, the tooth mark runs across the bone’s articulation with metatarsal II. Because the metatarsus was tightly bound in life, it would have been difficult to inflict such a mark on the articulated foot of a living animal. Furthermore, fighting animals would be expected to inflict wounds to the head [
<bibRefCitation author="Tanke DH &amp; Currie PJ" box="[281,309,359,381]" journalOrPublisher="Gaia" pageId="4" pageNumber="5" pagination="167 - 184" part="15" refId="ref4904" refString="32. Tanke DH, Currie PJ (1998) Head-biting behavior in the theropod dinosaurs: paleopathological evidence. Gaia 15: 167 - 184." title="Head-biting behavior in the theropod dinosaurs: paleopathological evidence" type="journal article" year="1998">32</bibRefCitation>
] or vulnerable areas such as the neck and flanks, and not the feet or arms. Finally, the absence of healing in any of these specimens is also consistent with the hypothesis that the tooth marks were made on carcasses.
</paragraph>
<paragraph blockId="4.[155,794,163,1385]" pageId="4" pageNumber="5">
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[179,302,470,492]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[179,302,470,492]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
</taxonomicName>
therefore seems to have been an indiscriminate and opportunistic feeder, feeding not only on herbivorous dinosaurs, but also on members of its own species. The traces described here likely result from opportunistic scavenging, and were probably made after most of the flesh and organs had been removed from the carcass. Presumably, an animal feeding on a fresh kill would instead be expected to focus on viscera and large muscle masses, which would provide more food with less effort. For feet, toes, and arms to be an appealing source of food, most of the carcass must already have been defleshed. It is somewhat perplexing why so few tooth marks are found on other elements, however. Tooth marks made by Komodo dragons [
<bibRefCitation author="D'Amore DC &amp; Blumenschine RJ" box="[650,677,777,799]" journalOrPublisher="Paleobiology" pageId="4" pageNumber="5" pagination="525 - 552" part="35" refId="ref4395" refString="18. D'Amore DC, Blumenschine RJ (2009) Komodo monitor (Varanus komodoensis) feeding behavior and dental function reflected through tooth marks on bone surfaces as the application to ziphodont paleobiology. Paleobiology 35: 525 - 552." title="Komodo monitor (Varanus komodoensis) feeding behavior and dental function reflected through tooth marks on bone surfaces as the application to ziphodont paleobiology" type="journal article" year="2009">18</bibRefCitation>
] and extant carnivorans [
<bibRefCitation author="Shipman P" box="[285,312,805,827]" journalOrPublisher="American Anthropologist" pageId="4" pageNumber="5" pagination="27 - 43" part="88" refId="ref4931" refString="33. Shipman P (1986) Scavenging or hunting in early hominids: theoretical framework and tests. American Anthropologist 88: 27 - 43." title="Scavenging or hunting in early hominids: theoretical framework and tests" type="journal article" year="1986">33</bibRefCitation>
] tend to be concentrated on elements bearing more meat, and it is therefore surprising not to find more traces made during the initial defleshing of the carcass.
</paragraph>
<paragraph blockId="4.[155,794,163,1385]" pageId="4" pageNumber="5">
While we interpret these traces as the results of scavenging, we cannot entirely rule out the possibility that these traces result from an individual slowly consuming a kill over an extended period of time. It does seem improbable that 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[502,625,972,994]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[502,625,972,994]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
</taxonomicName>
routinely hunted full-grown members of its own species; however, it is possible that intraspecific combat led to casualties, with the dead becoming a convenient source of food for the victors. Still, compelling evidence for predation in 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[320,443,1083,1105]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[320,443,1083,1105]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
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remains elusive. Healed injuries in herbivorous dinosaurs are consistent with failed predation [11,13] but it is debatable whether these traces are actually the results of bites, or some other form of trauma.
</paragraph>
<paragraph blockId="4.[155,794,163,1385]" lastBlockId="4.[840,1479,164,1060]" pageId="4" pageNumber="5">
Four examples of cannibalism are known from a relatively limited sample of tooth-marked bones. Given this, cannibalism must have been common in 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[479,602,1251,1273]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[479,602,1251,1273]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
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. If anything, the frequency of cannibalism is easily underestimated, for several reasons. First, the act of feeding on a carcass tends to destroy the evidence, because bones may be ingested, broken up, or dragged off and left to weather away out in the open. Second, cannibalism can only be observed on a carcass where the animals leave tooth marks; where 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[988,1111,192,214]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[988,1111,192,214]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
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fed around the bones, such events would not be recognized. Third, many 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1225,1348,220,242]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1225,1348,220,242]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
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skeletons are mounted, preventing detailed examination of the bones for tooth marks. Fourth, although we examined as many bones in as many museums as possible, it was not possible to examine all specimens of 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[865,988,332,354]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[865,988,332,354]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
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in all museums. Given this, it is perhaps surprising to find even a single instance of cannibalism, let alone multiple examples.
</paragraph>
<paragraph blockId="4.[840,1479,164,1060]" pageId="4" pageNumber="5">
Recent studies have questioned whether cannibalism was widespread in dinosaurs [
<bibRefCitation author="Nesbitt SJ &amp; Turner AH &amp; Erickson GM &amp; Norell MA" box="[1090,1119,446,468]" journalOrPublisher="Biology Letters" pageId="4" pageNumber="5" pagination="611 - 614" part="2" refId="ref4958" refString="34. Nesbitt SJ, Turner AH, Erickson GM, Norell MA (2006) Prey choice and cannibalistic behavior in the theropod Coelophysis. Biology Letters 2: 611 - 614." title="Prey choice and cannibalistic behavior in the theropod Coelophysis" type="journal article" year="2006">34</bibRefCitation>
], but the traces described here show that 
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<emphasis box="[890,1013,474,496]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
</taxonomicName>
was indisputably a cannibal. The only other dinosaur known to have engaged in cannibalism is the abelisaurid 
<taxonomicName authorityName="Sues &amp; Taquet" authorityYear="1979" box="[840,962,530,552]" class="Reptilia" family="Pachycephalosauridae" genus="Majungatholus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[840,962,530,552]" italics="true" pageId="4" pageNumber="5">Majungatholus</emphasis>
</taxonomicName>
[
<bibRefCitation author="Rogers RR &amp; Krause DW &amp; Curry Rogers K" box="[979,1007,531,553]" journalOrPublisher="Nature" pageId="4" pageNumber="5" pagination="515 - 518" part="422" refId="ref4324" refString="16. Rogers RR, Krause DW, Curry Rogers K (2003) Cannibalism in the Madagascan dinosaur Majungatholus atopus. Nature 422: 515 - 518." title="Cannibalism in the Madagascan dinosaur Majungatholus atopus" type="journal article" year="2003">16</bibRefCitation>
], however theropod tooth marks also occur on tyrannosaurid bones from the Dinosaur Park Formation [32,35]. Because two tyrannosaurids- 
<taxonomicName box="[1130,1230,587,609]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1130,1230,587,609]" italics="true" pageId="4" pageNumber="5">Gorgosaurus</emphasis>
</taxonomicName>
and 
<taxonomicName authorityName="Russell" authorityYear="1970" box="[1285,1408,587,609]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1285,1408,587,609]" italics="true" pageId="4" pageNumber="5">Daspletosaurus</emphasis>
</taxonomicName>
- occur here, it is impossible to definitively state that these traces represent cannibalism [
<bibRefCitation author="Nesbitt SJ &amp; Turner AH &amp; Erickson GM &amp; Norell MA" box="[980,1009,643,665]" journalOrPublisher="Biology Letters" pageId="4" pageNumber="5" pagination="611 - 614" part="2" refId="ref4958" refString="34. Nesbitt SJ, Turner AH, Erickson GM, Norell MA (2006) Prey choice and cannibalistic behavior in the theropod Coelophysis. Biology Letters 2: 611 - 614." title="Prey choice and cannibalistic behavior in the theropod Coelophysis" type="journal article" year="2006">34</bibRefCitation>
]. However, because 
<taxonomicName box="[1244,1344,643,665]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1244,1344,643,665]" italics="true" pageId="4" pageNumber="5">Gorgosaurus</emphasis>
</taxonomicName>
outnumbers 
<taxonomicName authorityName="Russell" authorityYear="1970" box="[840,963,671,693]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[840,963,671,693]" italics="true" pageId="4" pageNumber="5">Daspletosaurus</emphasis>
</taxonomicName>
by three-to-one in this environment [
<bibRefCitation author="Currie PJ &amp; Russell DA" box="[1352,1381,671,693]" editor="Currie PJ &amp; Koppelhus EB" journalOrPublisher="Bloomington: Indiana University Press" pageId="4" pageNumber="5" pagination="202 - 220" refId="ref5021" refString="36. Currie PJ, Russell DA (2005) The geographic and stratigraphic distribution of articulated and associated dinosaur remains. In: Currie PJ, Koppelhus EB, eds. Dinosaur Provincial Park: a spectacular ancient ecosystem revealed. Bloomington: Indiana University Press. pp 202 - 220." title="The geographic and stratigraphic distribution of articulated and associated dinosaur remains" type="book chapter" volumeTitle="Dinosaur Provincial Park: a spectacular ancient ecosystem revealed" year="2005">36</bibRefCitation>
], most of the bones and feeding traces probably represent 
<taxonomicName box="[1331,1431,699,721]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1331,1431,699,721]" italics="true" pageId="4" pageNumber="5">Gorgosaurus</emphasis>
</taxonomicName>
and therefore it is probable that at least some of these traces represent cannibalism.
</paragraph>
<paragraph blockId="4.[840,1479,164,1060]" pageId="4" pageNumber="5">
Cannibalism is common in nature [
<bibRefCitation author="Fox LR" box="[1237,1266,785,807]" journalOrPublisher="Annual Reviews of Ecology and Systematics" pageId="4" pageNumber="5" pagination="87 - 106" part="6" refId="ref5074" refString="37. Fox LR (1975) Cannibalism in natural populations. Annual Reviews of Ecology and Systematics 6: 87 - 106." title="Cannibalism in natural populations" type="journal article" year="1975">37</bibRefCitation>
], particularly among large carnivores, including bears [38,39,40], hyenas [
<bibRefCitation author="Kruuk H" bookContentInfo="335 p." box="[1380,1408,813,835]" journalOrPublisher="University of Chicago Press" pageId="4" pageNumber="5" refId="ref5205" refString="41. Kruuk H (1972) The Spotted Hyena: A Study of Predation and Social Behavior: University of Chicago Press. 335 p." title="The Spotted Hyena: A Study of Predation and Social Behavior" type="book" year="1972">41</bibRefCitation>
], large felids [42,43], Komodo dragons [
<bibRefCitation author="Auffenberg W" box="[1175,1204,841,863]" journalOrPublisher="Gainesville: University Presses of Florida" pageId="4" pageNumber="5" refId="ref5275" refString="44. Auffenberg W (1981) The Behavioral Ecology of the Komodo Monitor. Gainesville: University Presses of Florida." title="The Behavioral Ecology of the Komodo Monitor" type="book" year="1981">44</bibRefCitation>
], and crocodylians [45,46]. Notably, most documented cases of cannibalism in large carnivores involve predation, rather than scavenging. Cannibalism is especially common in the American alligator, and may account for more than half of the juvenile mortality each year [
<bibRefCitation author="Rootes WL &amp; Chabreck RH" box="[1372,1402,954,976]" pageId="4" pageNumber="5" pagination="99 - 107" refId="ref5334" refString="46. Rootes WL, Chabreck RH (1993) Cannibalism in the American alligator. Herpetologica 49: 99 - 107." type="journal article" year="1993">46</bibRefCitation>
]. Given that cannibalism is known in 
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1119,1242,981,1003]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1119,1242,981,1003]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
</taxonomicName>
, 
<taxonomicName authorityName="Sues &amp; Taquet" authorityYear="1979" box="[1254,1376,981,1003]" class="Reptilia" family="Pachycephalosauridae" genus="Majungatholus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
<emphasis box="[1254,1376,981,1003]" italics="true" pageId="4" pageNumber="5">Majungatholus</emphasis>
</taxonomicName>
and many extant, large-bodied carnivores, this behavior is likely to have been widespread in large, carnivorous dinosaurs.
</paragraph>
</subSubSection>
</treatment>
</document>