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<document id="3D1089C98DC2ADA4191EB9FEEF6BF041" ID-DOI="10.3390/fossils2010001" ID-Zenodo-Dep="10535269" IM.bibliography_approvedBy="karina" IM.illustrations_approvedBy="karina" IM.materialsCitations_approvedBy="karina" IM.metadata_approvedBy="karina" IM.tables_approvedBy="karina" IM.taxonomicNames_approvedBy="karina" IM.treatments_approvedBy="karina" checkinTime="1705674475442" checkinUser="karina" docAuthor="Longrich, Nicholas R. &amp; Saitta, Evan T." docDate="2024" docId="201187CAFFD8FF80FE21FA7EFDA8FE74" docLanguage="en" docName="FossStud.2024.2.1.1-65.pdf.imf" docOrigin="Fossil Studies 2 (1)" docSource="http://dx.doi.org/10.3390/fossils2010001" docTitle="Nanotyrannus Bakker et al. 1988" docType="treatment" docVersion="4" lastPageNumber="42" masterDocId="DC28FFB2FFC3FFAAFF9DFFB5FFDAFFC4" masterDocTitle="Taxonomic Status of Nanotyrannus lancensis (Dinosauria: Tyrannosauroidea) - A Distinct Taxon of Small-Bodied Tyrannosaur" masterLastPageNumber="65" masterPageNumber="1" pageNumber="28" updateTime="1705687956675" updateUser="ExternalLinkService">
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<mods:title id="E228F9DD4BC8F836281E4F79D58C6F6B">Taxonomic Status of Nanotyrannus lancensis (Dinosauria: Tyrannosauroidea) - A Distinct Taxon of Small-Bodied Tyrannosaur</mods:title>
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<mods:namePart id="D407C171622F9315929EC3002327C872">Longrich, Nicholas R.</mods:namePart>
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<mods:affiliation id="BEF86FD087E7DEB7962D13649C57BD8A">Department of Life Sciences, University of Bath, Claverton Down, Bath BA 2 7 AY, UK</mods:affiliation>
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<mods:namePart id="66CFF390A3B10B60B62DC25FEA4D1317">Saitta, Evan T.</mods:namePart>
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<mods:affiliation id="E0DCAFCD6F901293A089B39429B8D3F6">Department of Organismal Biology and Anatomy, University of Chicago, Chicago, IL 60637, USA</mods:affiliation>
<mods:nameIdentifier id="9BE491C68A056E3C08B260543F9501EA" type="email">evansaitta@gmail.com</mods:nameIdentifier>
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<mods:title id="A4AADA172F5E740BE2C355A2A1AA564B">Fossil Studies</mods:title>
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<mods:date id="993028AED61CEBE108263112F830E57A">2024</mods:date>
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<treatment id="201187CAFFD8FF80FE21FA7EFDA8FE74" ID-DOI="http://doi.org/10.5281/zenodo.10534261" ID-Zenodo-Dep="10534261" LSID="urn:lsid:plazi:treatment:201187CAFFD8FF80FE21FA7EFDA8FE74" httpUri="http://treatment.plazi.org/id/201187CAFFD8FF80FE21FA7EFDA8FE74" lastPageId="42" lastPageNumber="42" pageId="27" pageNumber="28">
<subSubSection id="E0A26557FFD8FFB1FE21FA7EFD15FA3A" pageId="27" pageNumber="28" type="nomenclature">
<paragraph id="A80736DCFFD8FFB1FE21FA7EFD15FA3A" blockId="27.[443,1497,1482,2045]" pageId="27" pageNumber="28">
<emphasis id="9ACCEACEFFD8FFB1FE21FA7EFB3EFA27" box="[444,1252,1482,1507]" italics="true" pageId="27" pageNumber="28">
3.3.
<taxonomicName id="6FB84D5FFFD8FFB1FE71FA7EFD50FA26" authorityName="Bakker et al." authorityYear="1988" box="[492,650,1483,1506]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">Nanotyrannus</taxonomicName>
Morphology Inconsistent with Predicted Morphology of
</emphasis>
<emphasis id="9ACCEACEFFD8FFB1FE21FA53FD15FA3A" box="[444,719,1510,1534]" italics="true" pageId="27" pageNumber="28">Juvenile Tyrannosaurinae</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="E0A26557FFD8FF80FE69F9A5FDA8FE74" lastPageId="42" lastPageNumber="43" pageId="27" pageNumber="28" type="description">
<paragraph id="A80736DCFFD8FFB1FE69F9A5FD1AF934" blockId="27.[443,1497,1482,2045]" pageId="27" pageNumber="28">
The hypothesized “growth series” linking
<taxonomicName id="6FB84D5FFFD8FFB1FC41F9A4FBADF9EC" box="[988,1143,1553,1576]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FC41F9A4FBADF9EC" box="[988,1143,1553,1576]" italics="true" pageId="27" pageNumber="28">Nanotyrannus</emphasis>
</taxonomicName>
to
<taxonomicName id="6FB84D5FFFD8FFB1FB07F9A4FAE0F9EC" box="[1178,1338,1553,1576]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FB07F9A4FAE0F9EC" box="[1178,1338,1553,1576]" italics="true" pageId="27" pageNumber="28">Tyrannosaurus</emphasis>
</taxonomicName>
can be tested by comparing it with the growth series of other tyrannosaurids, especially tyrannosaurines. If
<taxonomicName id="6FB84D5FFFD8FFB1FE49F9E1FD56F9AF" box="[468,652,1619,1643]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="46" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FE49F9E1FDA8F9AF" box="[468,626,1620,1643]" italics="true" pageId="27" pageNumber="28">Nanotyrannus</emphasis>
is
</taxonomicName>
a juvenile of
<taxonomicName id="6FB84D5FFFD8FFB1FCB4F9E1FC16F9AF" box="[809,972,1620,1643]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FCB4F9E1FC16F9AF" box="[809,972,1620,1643]" italics="true" pageId="27" pageNumber="28">Tyrannosaurus</emphasis>
</taxonomicName>
, and its distinctive morphology is the result of immaturity, then features of
<taxonomicName id="6FB84D5FFFD8FFB1FCADF9C0FC0BF948" box="[816,977,1653,1676]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FCADF9C0FC0BF948" box="[816,977,1653,1676]" italics="true" pageId="27" pageNumber="28">Nanotyrannus</emphasis>
</taxonomicName>
are predicted to occur in juveniles of other tyrannosaurs. If
<taxonomicName id="6FB84D5FFFD8FFB1FD19F923FC9AF96A" box="[644,832,1686,1710]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="46" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FD19F923FCFFF969" box="[644,805,1686,1709]" italics="true" pageId="27" pageNumber="28">Nanotyrannus</emphasis>
is
</taxonomicName>
a distinct taxon, then these features will be absent. We argue that juveniles of other tyrannosaurs do not conform to the “growth series” proposed for
<taxonomicName id="6FB84D5FFFD8FFB1FE79F96CFD5DF934" box="[484,647,1753,1776]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FE79F96CFD5DF934" box="[484,647,1753,1776]" italics="true" pageId="27" pageNumber="28">Tyrannosaurus</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFD8FFB1FD0AF96CFD68F935" author="Carr, T. D." box="[663,690,1753,1777]" journalOrPublisher="PeerJ" pageId="27" pageNumber="28" pagination="e 9192" part="8" refId="ref27454" refString="10. Carr, T. D. A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence. PeerJ 2020, 8, e 9192. [CrossRef]" title="A high-resolution growth series of Tyrannosaurus rex obtained from multiple lines of evidence" type="journal article" year="2020">10</bibRefCitation>
].
</paragraph>
<paragraph id="A80736DCFFD8FFB1FE69F94FFC76F87D" blockId="27.[443,1497,1482,2045]" pageId="27" pageNumber="28">
A young juvenile of
<taxonomicName id="6FB84D5FFFD8FFB1FD7CF94FFC71F8D6" box="[737,939,1786,1810]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="bataar">
<emphasis id="9ACCEACEFFD8FFB1FD7CF94FFC71F8D6" box="[737,939,1786,1810]" italics="true" pageId="27" pageNumber="28">Tarbosaurus bataar</emphasis>
</taxonomicName>
, a close relative of
<taxonomicName id="6FB84D5FFFD8FFB1FB1AF94EFAFDF8D6" box="[1159,1319,1787,1810]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FB1AF94EFAFDF8D6" box="[1159,1319,1787,1810]" italics="true" pageId="27" pageNumber="28">Tyrannosaurus</emphasis>
</taxonomicName>
, is known [
<bibRefCitation id="CC294B2DFFD8FFB1FA33F94FFA13F8D6" author="Tsuihiji, T. &amp; Watabe, M. &amp; Tsogtbaatar, K. &amp; Tsubamoto, T. &amp; Barsbold, R. &amp; Suzuki, S. &amp; Lee, A. H. &amp; Ridgely, R. C. &amp; Kawahara, Y. &amp; Witmer, L. M." box="[1454,1481,1786,1810]" journalOrPublisher="J. Vertebr. Paleontol." pageId="27" pageNumber="28" pagination="497 - 517" part="31" refId="ref29685" refString="54. Tsuihiji, T.; Watabe, M.; Tsogtbaatar, K.; Tsubamoto, T.; Barsbold, R.; Suzuki, S.; Lee, A. H.; Ridgely, R. C.; Kawahara, Y.; Witmer, L. M. Cranial osteology of a juvenile specimen of Tarbosaurus bataar (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia. J. Vertebr. Paleontol. 2011, 31, 497 - 517. [CrossRef]" title="Cranial osteology of a juvenile specimen of Tarbosaurus bataar (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia" type="journal article" year="2011">54</bibRefCitation>
]. In several features—posteriorly wide nasals, a gracile postorbital, a slender dentary, and lack of the suborbital process of the orbit—the animal resembles
<taxonomicName id="6FB84D5FFFD8FFB1FB3BF88BFA9BF891" box="[1190,1345,1854,1877]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FB3BF88BFA9BF891" box="[1190,1345,1854,1877]" italics="true" pageId="27" pageNumber="28">Nanotyrannus</emphasis>
</taxonomicName>
. This means some features seen in
<taxonomicName id="6FB84D5FFFD8FFB1FD5AF8EAFCB2F8B2" box="[711,872,1887,1910]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FD5AF8EAFCB2F8B2" box="[711,872,1887,1910]" italics="true" pageId="27" pageNumber="28">Nanotyrannus</emphasis>
</taxonomicName>
could conceivably be juvenile characters, but these features do not necessarily mean that the animals are juvenile since they occur in adults of tyrannosauroids such as
<taxonomicName id="6FB84D5FFFD8FFB1FD7DF814FC88F87D" authorityName="Kurzanov" authorityYear="1976" box="[736,850,1953,1977]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FD7DF814FC88F87D" box="[736,850,1953,1977]" italics="true" pageId="27" pageNumber="28">Alioramus</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFD8FFB1FCFCF817FCA6F87E" author="Brusatte, S. L. &amp; Carr, T. D. &amp; Norell, M. A." box="[865,892,1954,1978]" journalOrPublisher="Bull. Am. Mus. Nat. Hist." pageId="27" pageNumber="28" pagination="1 - 197" part="2012" refId="ref30715" refString="70. Brusatte, S. L.; Carr, T. D.; Norell, M. A. The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bull. Am. Mus. Nat. Hist. 2012, 2012, 1 - 197. [CrossRef]" title="The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia" type="journal article" year="2012">70</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFD8FFB1FC1FF814FC47F87D" author="Brusatte, S. L. &amp; Carr, T. D. &amp; Erickson, G. M. &amp; Bever, G. S. &amp; Norell, M." box="[898,925,1953,1977]" journalOrPublisher="Proc. Natl. Acad. Sci. USA" pageId="27" pageNumber="28" pagination="17251 - 17266" part="106" refId="ref32124" refString="96. Brusatte, S. L.; Carr, T. D.; Erickson, G. M.; Bever, G. S.; Norell, M. A long-snouted, multihorned tyrannosaurid from the Late Cretaceous of Mongolia. Proc. Natl. Acad. Sci. USA 2009, 106, 17251 - 17266. [CrossRef]" title="A long-snouted, multihorned tyrannosaurid from the Late Cretaceous of Mongolia" type="journal article" year="2009">96</bibRefCitation>
].
</paragraph>
<paragraph id="A80736DCFFD8FFB6FE69F876FA03FEA8" blockId="27.[443,1497,1482,2045]" lastBlockId="28.[444,1497,240,364]" lastPageId="28" lastPageNumber="29" pageId="27" pageNumber="28">
However, the juvenile
<taxonomicName id="6FB84D5FFFD8FFB1FD68F876FCACF81F" box="[757,886,1987,2011]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FD68F876FCACF81F" box="[757,886,1987,2011]" italics="true" pageId="27" pageNumber="28">Tarbosaurus</emphasis>
</taxonomicName>
skull differs from that of
<taxonomicName id="6FB84D5FFFD8FFB1FB09F871FAF5F81F" box="[1172,1327,1988,2011]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FB09F871FAF5F81F" box="[1172,1327,1988,2011]" italics="true" pageId="27" pageNumber="28">Nanotyrannus</emphasis>
</taxonomicName>
in many ways while resembling adult
<taxonomicName id="6FB84D5FFFD8FFB1FD50F851FC95F838" box="[717,847,2020,2044]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FD50F851FC95F838" box="[717,847,2020,2044]" italics="true" pageId="27" pageNumber="28">Tarbosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="6FB84D5FFFD8FFB1FC15F850FC1CF838" box="[904,966,2021,2044]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFD8FFB1FC15F850FC42F838" box="[904,920,2021,2044]" italics="true" pageId="27" pageNumber="28">T</emphasis>
.
<emphasis id="9ACCEACEFFD8FFB1FC38F850FC1CF838" box="[933,966,2021,2044]" italics="true" pageId="27" pageNumber="28">rex</emphasis>
</taxonomicName>
(
<figureCitation id="30832A59FFD8FFB1FC48F851FB9BF839" box="[981,1089,2020,2045]" captionStart="Figure 19" captionStartId="28.[444,515,1798,1820]" captionTargetBox="[447,1488,412,1764]" captionTargetId="figure-191@28.[446,1493,411,1765]" captionTargetPageId="28" captionText="Figure 19. Ontogenetically stable characters in juveniles and adults of Tarbosaurus (Tyrannosaurini). Characters: (a) promaxillary fenestra concealed, (b) narial process of premaxilla faces anteriorly, (c) maxilla relatively short and tall, (d) promaxillary fenestra concealed in lateral view, (e) maxillary fenestra positioned anteriorly and ventrally, (f) lacrimal horn low, (g) orbit tall, (h) jugal narrow below orbit, (i) weakly curved orbital bar of lacrimal, (j) large anterior maxillary teeth, (k) first maxillary tooth large, (l) premaxillary teeth with pointed apices and serrated. Scale = 10 cm." figureDoi="http://doi.org/10.5281/zenodo.10535309" httpUri="https://zenodo.org/record/10535309/files/figure.png" pageId="27" pageNumber="28">Figure 19</figureCitation>
). Features shared with
<taxonomicName id="6FB84D5FFFD8FFB1FACFF851FA0EF838" box="[1362,1492,2020,2044]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFD8FFB1FACFF851FA0EF838" box="[1362,1492,2020,2044]" italics="true" pageId="27" pageNumber="28">Tarbosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="6FB84D5FFFDFFFB6FE6DFF44FDF5FECC" box="[496,559,241,264]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="28" pageNumber="29" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFDFFFB6FE6DFF44FDDAFECC" box="[496,512,241,264]" italics="true" pageId="28" pageNumber="29">T</emphasis>
.
<emphasis id="9ACCEACEFFDFFFB6FD90FF44FDF5FECC" box="[525,559,241,264]" italics="true" pageId="28" pageNumber="29">rex</emphasis>
</taxonomicName>
(but not
<taxonomicName id="6FB84D5FFFDFFFB6FD07FF44FCE3FECC" box="[666,825,241,264]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDFFFB6FD07FF44FCE3FECC" box="[666,825,241,264]" italics="true" pageId="28" pageNumber="29">Nanotyrannus</emphasis>
</taxonomicName>
) include the tall and deep maxilla, the narrow rim of the antorbital fossa, an anteriorly placed maxillary fenestra, a large maxillary fenestra, limited contribution of the lacrimal to the antorbital fossa, weak curvature of the ventral ramus of the lacrimal, an anteriorly expanded jugal, and a broad base of the jugal postorbital process.
</paragraph>
<caption id="FCC76654FFDFFFB6FE21F8B3FAC7F803" ID-DOI="http://doi.org/10.5281/zenodo.10535309" ID-Zenodo-Dep="10535309" httpUri="https://zenodo.org/record/10535309/files/figure.png" pageId="28" pageNumber="29" startId="28.[444,515,1798,1820]" targetBox="[447,1488,412,1764]" targetPageId="28" targetType="figure">
<paragraph id="A80736DCFFDFFFB6FE21F8B3FAC7F803" blockId="28.[443,1496,1798,1992]" pageId="28" pageNumber="29">
<emphasis id="9ACCEACEFFDFFFB6FE21F8B3FDFDF8DF" bold="true" box="[444,551,1798,1820]" pageId="28" pageNumber="29">Figure 19.</emphasis>
Ontogenetically stable characters in juveniles and adults of
<taxonomicName id="6FB84D5FFFDFFFB6FB3EF8B2FA0EF8D9" box="[1187,1492,1799,1821]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="28" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDFFFB6FB3EF8B2FAC0F8D8" box="[1187,1306,1799,1820]" italics="true" pageId="28" pageNumber="29">Tarbosaurus</emphasis>
(Tyrannosaurini)
</taxonomicName>
. Characters: (
<emphasis id="9ACCEACEFFDFFFB6FDD4F89CFD8FF8FA" bold="true" box="[585,597,1833,1854]" pageId="28" pageNumber="29">a</emphasis>
) promaxillary fenestra concealed, (
<emphasis id="9ACCEACEFFDFFFB6FC4AF89CFC3CF8FA" bold="true" box="[983,998,1833,1854]" pageId="28" pageNumber="29">b</emphasis>
) narial process of premaxilla faces anteriorly, (
<emphasis id="9ACCEACEFFDFFFB6FE5EF8FEFE14F8A4" bold="true" box="[451,462,1867,1888]" pageId="28" pageNumber="29">c</emphasis>
) maxilla relatively short and tall, (
<emphasis id="9ACCEACEFFDFFFB6FCA4F8FEFC92F8A4" bold="true" box="[825,840,1867,1888]" pageId="28" pageNumber="29">d</emphasis>
) promaxillary fenestra concealed in lateral view, (
<emphasis id="9ACCEACEFFDFFFB6FACBF8FEFAB8F8A4" bold="true" box="[1366,1378,1867,1888]" pageId="28" pageNumber="29">e</emphasis>
) maxillary fenestra positioned anteriorly and ventrally, (
<emphasis id="9ACCEACEFFDFFFB6FC10F8D8FC4CF847" bold="true" box="[909,918,1901,1923]" pageId="28" pageNumber="29">f</emphasis>
) lacrimal horn low, (
<emphasis id="9ACCEACEFFDFFFB6FBF6F8D8FBA2F846" bold="true" box="[1131,1144,1901,1922]" pageId="28" pageNumber="29">g</emphasis>
) orbit tall, (
<emphasis id="9ACCEACEFFDFFFB6FB6DF8D8FB24F846" bold="true" box="[1264,1278,1901,1922]" pageId="28" pageNumber="29">h</emphasis>
) jugal narrow below orbit, (
<emphasis id="9ACCEACEFFDFFFB6FD9EF83AFDD1F861" bold="true" box="[515,523,1935,1957]" pageId="28" pageNumber="29">i</emphasis>
) weakly curved orbital bar of lacrimal, (
<emphasis id="9ACCEACEFFDFFFB6FC21F83AFC1EF861" bold="true" box="[956,964,1935,1957]" pageId="28" pageNumber="29">j</emphasis>
) large anterior maxillary teeth, (
<emphasis id="9ACCEACEFFDFFFB6FABDF825FAF5F861" bold="true" box="[1312,1327,1936,1957]" pageId="28" pageNumber="29">k</emphasis>
) first maxillary tooth large, (
<emphasis id="9ACCEACEFFDFFFB6FDDFF807FD90F803" bold="true" box="[578,586,1970,1991]" pageId="28" pageNumber="29">l</emphasis>
) premaxillary teeth with pointed apices and serrated. Scale = 10 cm.
</paragraph>
</caption>
<paragraph id="A80736DCFFDEFFB7FE69FF45FB09FEA9" blockId="29.[444,1496,240,365]" pageId="29" pageNumber="30">
These features appear early in the ontogeny of
<taxonomicName id="6FB84D5FFFDEFFB7FB88FF45FB4DFECC" box="[1045,1175,240,264]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDEFFB7FB88FF45FB4DFECC" box="[1045,1175,240,264]" italics="true" pageId="29" pageNumber="30">Tarbosaurus</emphasis>
</taxonomicName>
and would presumably occur early in the ontogeny of
<taxonomicName id="6FB84D5FFFDEFFB7FD67FEA7FC40FEED" box="[762,922,274,297]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDEFFB7FD67FEA7FC40FEED" box="[762,922,274,297]" italics="true" pageId="29" pageNumber="30">Tyrannosaurus</emphasis>
</taxonomicName>
. The absence of these features in absolutely larger
<taxonomicName id="6FB84D5FFFDEFFB7FE21FE81FD86FE8F" box="[444,604,308,331]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDEFFB7FE21FE81FD86FE8F" box="[444,604,308,331]" italics="true" pageId="29" pageNumber="30">Nanotyrannus</emphasis>
</taxonomicName>
specimens is difficult to explain in terms of ontogeny unless
<taxonomicName id="6FB84D5FFFDEFFB7FAB2FE81FA0EFE8F" box="[1327,1492,308,331]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDEFFB7FAB2FE81FA0EFE8F" box="[1327,1492,308,331]" italics="true" pageId="29" pageNumber="30">Tyrannosaurus</emphasis>
</taxonomicName>
had a pattern of development unlike that of
<taxonomicName id="6FB84D5FFFDEFFB7FC5EFEE1FB9DFEA8" box="[963,1095,340,364]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDEFFB7FC5EFEE1FB9DFEA8" box="[963,1095,340,364]" italics="true" pageId="29" pageNumber="30">Tarbosaurus</emphasis>
</taxonomicName>
(
<figureCitation id="30832A59FFDEFFB7FBC8FEE1FB1EFEA9" box="[1109,1220,340,365]" captionStart="Figure 20" captionStartId="29.[444,516,1980,2002]" captionTargetBox="[445,1194,397,1954]" captionTargetId="figure-78@29.[512,1093,889,1340]" captionTargetPageId="29" captionText="Figure 20. Growth series of Tarbosaurus. Top to bottom: PIN 551-1 [108], ZPAL MgD-I/4 [53], PIN 553-1 [108], PIN 552-2 [108]. Scale = 10 cm." figureDoi="http://doi.org/10.5281/zenodo.10535311" httpUri="https://zenodo.org/record/10535311/files/figure.png" pageId="29" pageNumber="30">Figure 20</figureCitation>
).
</paragraph>
<caption id="FCC76654FFDEFFB7FE21F809FCA7F830" ID-DOI="http://doi.org/10.5281/zenodo.10535311" ID-Zenodo-Dep="10535311" httpUri="https://zenodo.org/record/10535311/files/figure.png" pageId="29" pageNumber="30" startId="29.[444,516,1980,2002]" targetBox="[445,1194,397,1954]" targetPageId="29" targetType="figure">
<paragraph id="A80736DCFFDEFFB7FE21F809FCA7F830" blockId="29.[444,1491,1980,2036]" pageId="29" pageNumber="30">
<emphasis id="9ACCEACEFFDEFFB7FE21F809FDF3F815" bold="true" box="[444,553,1980,2002]" pageId="29" pageNumber="30">Figure 20.</emphasis>
Growth series of
<taxonomicName id="6FB84D5FFFDEFFB7FD75F809FCBBF815" box="[744,865,1980,2001]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="29" pageNumber="30" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDEFFB7FD75F809FCBBF815" box="[744,865,1980,2001]" italics="true" pageId="29" pageNumber="30">Tarbosaurus</emphasis>
</taxonomicName>
. Top to bottom: PIN 551-1 [
<bibRefCitation id="CC294B2DFFDEFFB7FB13F809FB68F815" author="Maleev, E. A." box="[1166,1202,1980,2001]" journalOrPublisher="Jt. Sov. Mong. Paleontol. Exped." pageId="29" pageNumber="30" pagination="132 - 191" part="1" refId="ref32771" refString="108. Maleev, E. A. Giant carnosaurs of the family Tyrannosauridae. Jt. Sov. Mong. Paleontol. Exped. 1974, 1, 132 - 191." title="Giant carnosaurs of the family Tyrannosauridae" type="journal article" year="1974">108</bibRefCitation>
], ZPAL MgD-I/4 [
<bibRefCitation id="CC294B2DFFDEFFB7FAE1F809FA4EF816" author="Hurum, J. H. &amp; Sabath, K." box="[1404,1428,1980,2002]" journalOrPublisher="Acta Palaeontol. Pol." pageId="29" pageNumber="30" pagination="161 - 190" part="48" refId="ref29641" refString="53. Hurum, J. H.; Sabath, K. Giant theropod dinosaurs from Asia and North America: Skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontol. Pol. 2003, 48, 161 - 190." title="Giant theropod dinosaurs from Asia and North America: Skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">53</bibRefCitation>
], PIN 553-1 [
<bibRefCitation id="CC294B2DFFDEFFB7FD9CF86AFDFFF830" author="Maleev, E. A." box="[513,549,2015,2036]" journalOrPublisher="Jt. Sov. Mong. Paleontol. Exped." pageId="29" pageNumber="30" pagination="132 - 191" part="1" refId="ref32771" refString="108. Maleev, E. A. Giant carnosaurs of the family Tyrannosauridae. Jt. Sov. Mong. Paleontol. Exped. 1974, 1, 132 - 191." title="Giant carnosaurs of the family Tyrannosauridae" type="journal article" year="1974">108</bibRefCitation>
], PIN 552-2 [
<bibRefCitation id="CC294B2DFFDEFFB7FD32F86AFD09F830" author="Maleev, E. A." box="[687,723,2015,2036]" journalOrPublisher="Jt. Sov. Mong. Paleontol. Exped." pageId="29" pageNumber="30" pagination="132 - 191" part="1" refId="ref32771" refString="108. Maleev, E. A. Giant carnosaurs of the family Tyrannosauridae. Jt. Sov. Mong. Paleontol. Exped. 1974, 1, 132 - 191." title="Giant carnosaurs of the family Tyrannosauridae" type="journal article" year="1974">108</bibRefCitation>
]. Scale = 10 cm.
</paragraph>
</caption>
<paragraph id="A80736DCFFDDFFB4FE69FF45FAEDFEA8" blockId="30.[442,1496,240,1101]" pageId="30" pageNumber="31">
Some features of Tyrannosaurinae, especially those related to the skull ornamentation, orbits, and skull roof, appear to develop late, but others appear in even the youngest specimens (
<figureCitation id="30832A59FFDDFFB4FDDFFE86FD63FE8F" box="[578,697,307,331]" captionStart="Figure 19" captionStartId="28.[444,515,1798,1820]" captionTargetBox="[447,1488,412,1764]" captionTargetId="figure-191@28.[446,1493,411,1765]" captionTargetPageId="28" captionText="Figure 19. Ontogenetically stable characters in juveniles and adults of Tarbosaurus (Tyrannosaurini). Characters: (a) promaxillary fenestra concealed, (b) narial process of premaxilla faces anteriorly, (c) maxilla relatively short and tall, (d) promaxillary fenestra concealed in lateral view, (e) maxillary fenestra positioned anteriorly and ventrally, (f) lacrimal horn low, (g) orbit tall, (h) jugal narrow below orbit, (i) weakly curved orbital bar of lacrimal, (j) large anterior maxillary teeth, (k) first maxillary tooth large, (l) premaxillary teeth with pointed apices and serrated. Scale = 10 cm." figureDoi="http://doi.org/10.5281/zenodo.10535309" httpUri="https://zenodo.org/record/10535309/files/figure.png" pageId="30" pageNumber="31">Figures 19</figureCitation>
and
<figureCitation id="30832A59FFDDFFB4FD6FFE86FCD6FE8F" box="[754,780,307,331]" captionStart="Figure 20" captionStartId="29.[444,516,1980,2002]" captionTargetBox="[445,1194,397,1954]" captionTargetId="figure-78@29.[512,1093,889,1340]" captionTargetPageId="29" captionText="Figure 20. Growth series of Tarbosaurus. Top to bottom: PIN 551-1 [108], ZPAL MgD-I/4 [53], PIN 553-1 [108], PIN 552-2 [108]. Scale = 10 cm." figureDoi="http://doi.org/10.5281/zenodo.10535311" httpUri="https://zenodo.org/record/10535311/files/figure.png" pageId="30" pageNumber="31">20</figureCitation>
). At least some tyrannosaurine features would be expected in
<taxonomicName id="6FB84D5FFFDDFFB4FE21FEE0FD80FEA8" box="[444,602,341,364]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FE21FEE0FD80FEA8" box="[444,602,341,364]" italics="true" pageId="30" pageNumber="31">Nanotyrannus</emphasis>
</taxonomicName>
if it was a juvenile tyrannosaurine, but few, if any, are present.
</paragraph>
<paragraph id="A80736DCFFDDFFB4FE69FEC3FDBDFDBC" blockId="30.[442,1496,240,1101]" pageId="30" pageNumber="31">
Juveniles are also known for
<taxonomicName id="6FB84D5FFFDDFFB4FCD3FEC3FBECFE4A" box="[846,1078,374,398]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="species" species="libratus">
<emphasis id="9ACCEACEFFDDFFB4FCD3FEC3FBECFE4A" box="[846,1078,374,398]" italics="true" pageId="30" pageNumber="31">Gorgosaurus libratus</emphasis>
</taxonomicName>
, including skulls [
<bibRefCitation id="CC294B2DFFDDFFB4FA8FFEC3FAE0FE4A" author="Voris, J. T. &amp; Zelenitsky, D. K. &amp; Therrien, F. &amp; Ridgely, R. C. &amp; Currie, P. J. &amp; Witmer, L. M." box="[1298,1338,374,398]" journalOrPublisher="J. Vertebr. Paleontol." pageId="30" pageNumber="31" pagination="2041651" part="41" refId="ref32804" refString="109. Voris, J. T.; Zelenitsky, D. K.; Therrien, F.; Ridgely, R. C.; Currie, P. J.; Witmer, L. M. Two exceptionally preserved juvenile specimens of Gorgosaurus libratus (Tyrannosauridae, Albertosaurinae) provide new insight into the timing of ontogenetic changes in tyrannosaurids. J. Vertebr. Paleontol. 2021, 41, e 2041651. [CrossRef]" title="Two exceptionally preserved juvenile specimens of Gorgosaurus libratus (Tyrannosauridae, Albertosaurinae) provide new insight into the timing of ontogenetic changes in tyrannosaurids" type="journal article" year="2021">109</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFDDFFB4FADCFEC3FAB3FE4A" author="Voris, J. T. &amp; Zelenitsky, D. K. &amp; Therrien, F. &amp; Currie, P. J." box="[1345,1385,374,398]" journalOrPublisher="Sci. Rep." pageId="30" pageNumber="31" pagination="1 - 10" part="9" refId="ref32886" refString="110. Voris, J. T.; Zelenitsky, D. K.; Therrien, F.; Currie, P. J. Reassessment of a juvenile Daspletosaurus from the Late Cretaceous of Alberta, Canada with implications for the identification of immature tyrannosaurids. Sci. Rep. 2019, 9, 1 - 10. [CrossRef] [PubMed]" title="Reassessment of a juvenile Daspletosaurus from the Late Cretaceous of Alberta, Canada with implications for the identification of immature tyrannosaurids" type="journal article" year="2019">110</bibRefCitation>
] and isolated elements [
<bibRefCitation id="CC294B2DFFDDFFB4FDE3FE2DFD51FE74" author="Carr, T. D." box="[638,651,408,432]" journalOrPublisher="J. Vertebr. Paleontol." pageId="30" pageNumber="31" pagination="497 - 520" part="19" refId="ref27419" refString="9. Carr, T. D. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). J. Vertebr. Paleontol. 1999, 19, 497 - 520. [CrossRef]" title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">9</bibRefCitation>
]. Juveniles are remarkably similar to adult
<taxonomicName id="6FB84D5FFFDDFFB4FB37FE2DFAEDFE6B" authorityName="Lambe" authorityYear="1914" box="[1194,1335,408,431]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FB37FE2DFAEDFE6B" box="[1194,1335,408,431]" italics="true" pageId="30" pageNumber="31">Gorgosaurus</emphasis>
</taxonomicName>
, particularly in the shape of the maxilla, antorbital fenestra, antorbital fossa, and maxillary fenestra, implying that
<taxonomicName id="6FB84D5FFFDDFFB4FDF5FE6EFD2FFE36" authorityName="Lambe" authorityYear="1914" box="[616,757,475,498]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FDF5FE6EFD2FFE36" box="[616,757,475,498]" italics="true" pageId="30" pageNumber="31">Gorgosaurus</emphasis>
</taxonomicName>
did not undergo radical changes in skull anatomy as it grew. Neither do juvenile
<taxonomicName id="6FB84D5FFFDDFFB4FD3EFE49FCF1FDD7" authorityName="Lambe" authorityYear="1914" box="[675,811,508,531]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FD3EFE49FCF1FDD7" box="[675,811,508,531]" italics="true" pageId="30" pageNumber="31">Gorgosaurus</emphasis>
</taxonomicName>
exhibit
<taxonomicName id="6FB84D5FFFDDFFB4FC15FE48FBFEFDD0" box="[904,1060,509,532]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FC15FE48FBFEFDD0" box="[904,1060,509,532]" italics="true" pageId="30" pageNumber="31">Nanotyrannus</emphasis>
</taxonomicName>
-like characters such as the expanded antorbital fossa, procumbent premaxillary teeth, or a pneumatized quadratojugal. Growth patterns in
<emphasis id="9ACCEACEFFDDFFB4FDA1FD8AFD10FD92" box="[572,714,575,598]" italics="true" pageId="30" pageNumber="31">
<taxonomicName id="6FB84D5FFFDDFFB4FDA1FD8AFD1CFD92" authorityName="Lambe" authorityYear="1914" box="[572,710,575,598]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
,
</emphasis>
therefore, argue against
<taxonomicName id="6FB84D5FFFDDFFB4FC78FDF5FB5AFD93" box="[997,1152,576,599]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FC78FDF5FB5AFD93" box="[997,1152,576,599]" italics="true" pageId="30" pageNumber="31">Nanotyrannus</emphasis>
</taxonomicName>
morphology being the result of immaturity.
</paragraph>
<paragraph id="A80736DCFFDDFFB4FE69FD37FCE6FD3A" blockId="30.[442,1496,240,1101]" pageId="30" pageNumber="31">
For
<taxonomicName id="6FB84D5FFFDDFFB4FDB8FD36FD1CFD5E" box="[549,710,643,666]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FDB8FD36FD1CFD5E" box="[549,710,643,666]" italics="true" pageId="30" pageNumber="31">Nanotyrannus</emphasis>
</taxonomicName>
to be a juvenile
<taxonomicName id="6FB84D5FFFDDFFB4FC10FD36FBE9FD5E" box="[909,1075,643,666]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FC10FD36FBE9FD5E" box="[909,1075,643,666]" italics="true" pageId="30" pageNumber="31">Tyrannosaurus</emphasis>
</taxonomicName>
,
<taxonomicName id="6FB84D5FFFDDFFB4FBDCFD36FB3DFD5E" box="[1089,1255,643,666]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FBDCFD36FB3DFD5E" box="[1089,1255,643,666]" italics="true" pageId="30" pageNumber="31">Tyrannosaurus</emphasis>
</taxonomicName>
would have had to have a radically different development pattern than
<taxonomicName id="6FB84D5FFFDDFFB4FB86FD16FB47FD7F" box="[1051,1181,675,699]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FB86FD16FB47FD7F" box="[1051,1181,675,699]" italics="true" pageId="30" pageNumber="31">Tarbosaurus</emphasis>
</taxonomicName>
or
<taxonomicName id="6FB84D5FFFDDFFB4FB5EFD11FA96FD7F" authorityName="Lambe" authorityYear="1914" box="[1219,1356,676,699]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FB5EFD11FA96FD7F" box="[1219,1356,676,699]" italics="true" pageId="30" pageNumber="31">Gorgosaurus</emphasis>
</taxonomicName>
. This is not impossible; ontogeny evolves. However, it is more parsimonious to treat
<taxonomicName id="6FB84D5FFFDDFFB4FA9BFD73FA7BFD19" box="[1286,1441,710,733]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FA9BFD73FA7BFD19" box="[1286,1441,710,733]" italics="true" pageId="30" pageNumber="31">Nanotyrannus</emphasis>
</taxonomicName>
and
<taxonomicName id="6FB84D5FFFDDFFB4FE27FD52FD87FD3A" box="[442,605,743,766]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FE27FD52FD87FD3A" box="[442,605,743,766]" italics="true" pageId="30" pageNumber="31">Tyrannosaurus</emphasis>
</taxonomicName>
as distinct species.
</paragraph>
<paragraph id="A80736DCFFDDFFB4FE69FCBDFA4CFB88" blockId="30.[442,1496,240,1101]" pageId="30" pageNumber="31">
Finally, the proportions of the manus in the two animals are inconsistent with
<taxonomicName id="6FB84D5FFFDDFFB4FA02FCBDFDF6FC85" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FA02FCBDFDF6FC85" italics="true" pageId="30" pageNumber="31">Nanotyrannus</emphasis>
</taxonomicName>
developing into
<taxonomicName id="6FB84D5FFFDDFFB4FD68FC9FFC40FC85" box="[757,922,810,833]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FD68FC9FFC40FC85" box="[757,922,810,833]" italics="true" pageId="30" pageNumber="31">Tyrannosaurus</emphasis>
</taxonomicName>
(
<figureCitation id="30832A59FFDDFFB4FC34FC9CFBC3FC85" box="[937,1049,809,833]" captionStart="Figure 21" captionStartId="30.[444,515,1984,2006]" captionTargetBox="[452,1097,1143,1946]" captionTargetId="figure-667@30.[440,1134,1131,1951]" captionTargetPageId="30" captionText="Figure 21. Manual unguals of Tyrannosaurus and Nanotyrannus, shown to scale; small Nanotyrannus specimens have absolutely larger unguals than much larger Tyrannosaurus. Scale = 5 cm." figureDoi="http://doi.org/10.5281/zenodo.10535313" httpUri="https://zenodo.org/record/10535313/files/figure.png" pageId="30" pageNumber="31">Figure 21</figureCitation>
). Despite coming from much smaller animals, approximately
<quantity id="6F409B39FFDDFFB4FD40FCFFFCFEFCA6" box="[733,804,842,866]" metricMagnitude="0" metricUnit="m" metricValue="5.5" metricValueMax="6.0" metricValueMin="5.0" pageId="30" pageNumber="31" unit="m" value="5.5" valueMax="6.0" valueMin="5.0">56 m</quantity>
in length (versus
<quantity id="6F409B39FFDDFFB4FC64FCFEFBE9FCA6" box="[1017,1075,843,867]" metricMagnitude="1" metricUnit="m" metricValue="1.2" pageId="30" pageNumber="31" unit="m" value="12.0">12 m</quantity>
or more in
<taxonomicName id="6FB84D5FFFDDFFB4FB21FCFEFAB8FCA6" box="[1212,1378,843,866]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FB21FCFEFAB8FCA6" box="[1212,1378,843,866]" italics="true" pageId="30" pageNumber="31">Tyrannosaurus</emphasis>
</taxonomicName>
), manual phalanges of
<collectionCode id="CEA9AE19FFDDFFB4FDC8FCD8FD45FC40" box="[597,671,877,900]" collectionName="BMRP" pageId="30" pageNumber="31">BMRP</collectionCode>
<date id="DC06101CFFDDFFB4FD38FCD9FCDBFC40" box="[677,769,876,900]" pageId="30" pageNumber="31" value="2006-04-04">2006.4.4</date>
and
<collectionCode id="CEA9AE19FFDDFFB4FCA7FCD9FCB5FC40" box="[826,879,876,900]" collectionName="HRS" pageId="30" pageNumber="31">HRS</collectionCode>
15001 are significantly larger than those of even very large
<taxonomicName id="6FB84D5FFFDDFFB4FE60FC3BFCBCFC62" box="[509,870,910,934]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FE60FC3BFD45FC61" box="[509,671,910,933]" italics="true" pageId="30" pageNumber="31">Tyrannosaurus</emphasis>
,
<collectionCode id="CEA9AE19FFDDFFB4FD31FC3BFCDAFC61" box="[684,768,910,933]" collectionName="USA, Illinois, Chicago, Field Museum of Natural History (also used by Finnish Museum of Natural History)" pageId="30" pageNumber="31">FMNH</collectionCode>
PR 2081
</taxonomicName>
. While allometric growth is possible, with the manus becoming proportionately smaller, the proportions seen in
<taxonomicName id="6FB84D5FFFDDFFB4FBFDFC05FB21FC03" box="[1120,1275,944,967]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FBFDFC05FB21FC03" box="[1120,1275,944,967]" italics="true" pageId="30" pageNumber="31">Nanotyrannus</emphasis>
</taxonomicName>
require the manus and claws to become absolutely smallerfor bone to be resorbed and elements reduced in lengthas the animal matures. We are unaware of any amniote that develops in this fashion. Another problem is that the tip of the vomer is deeper in
<taxonomicName id="6FB84D5FFFDDFFB4FB4BFBA1FAADFBEF" box="[1238,1399,1044,1067]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FB4BFBA1FAADFBEF" box="[1238,1399,1044,1067]" italics="true" pageId="30" pageNumber="31">Nanotyrannus</emphasis>
</taxonomicName>
than in
<taxonomicName id="6FB84D5FFFDDFFB4FE27FB83FD87FB89" authorityName=", Sue FMNH PR" authorityYear="2081" box="[442,605,1078,1101]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FE27FB83FD87FB89" box="[442,605,1078,1101]" italics="true" pageId="30" pageNumber="31">Tyrannosaurus</emphasis>
</taxonomicName>
; this would require the end of the vomer to shrink or be resorbed [
<bibRefCitation id="CC294B2DFFDDFFB4FAF1FB80FA5DFB89" author="Larson, P." bookContentInfo="Indiana University Press: Bloomington, IN, USA" box="[1388,1415,1077,1101]" editor="Parrish, J. M. &amp; Molnar, R. E. &amp; Currie, P. J. &amp; Koppelhus, E. B." pageId="30" pageNumber="31" pagination="15 - 53" refId="ref28873" refString="40. Larson, P. The case for Nanotyrannus. In Tyrannosaurid Paleobiology; Parrish, J. M., Molnar, R. E., Currie, P. J., Koppelhus, E. B., Eds.; Indiana University Press: Bloomington, IN, USA, 2013; pp. 15 - 53." title="The case for Nanotyrannus" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">40</bibRefCitation>
].
</paragraph>
<caption id="FCC76654FFDDFFB4FE21F875FA85F83C" ID-DOI="http://doi.org/10.5281/zenodo.10535313" ID-Zenodo-Dep="10535313" httpUri="https://zenodo.org/record/10535313/files/figure.png" pageId="30" pageNumber="31" startId="30.[444,515,1984,2006]" targetBox="[452,1097,1143,1946]" targetPageId="30" targetType="figure">
<paragraph id="A80736DCFFDDFFB4FE21F875FA85F83C" blockId="30.[444,1492,1984,2041]" pageId="30" pageNumber="31">
<emphasis id="9ACCEACEFFDDFFB4FE21F875FDFDF812" bold="true" box="[444,551,1984,2006]" pageId="30" pageNumber="31">Figure 21.</emphasis>
Manual unguals of
<taxonomicName id="6FB84D5FFFDDFFB4FD61F874FC55F812" authorityName=", Sue FMNH PR" authorityYear="2081" box="[764,911,1985,2006]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FD61F874FC55F812" box="[764,911,1985,2006]" italics="true" pageId="30" pageNumber="31">Tyrannosaurus</emphasis>
</taxonomicName>
and
<taxonomicName id="6FB84D5FFFDDFFB4FC59F874FB89F812" box="[964,1107,1985,2006]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FC59F874FB89F812" box="[964,1107,1985,2006]" italics="true" pageId="30" pageNumber="31">Nanotyrannus</emphasis>
</taxonomicName>
, shown to scale; small
<taxonomicName id="6FB84D5FFFDDFFB4FAD8F874FA0EF812" box="[1349,1492,1985,2006]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FAD8F874FA0EF812" box="[1349,1492,1985,2006]" italics="true" pageId="30" pageNumber="31">Nanotyrannus</emphasis>
</taxonomicName>
specimens have absolutely larger unguals than much larger
<taxonomicName id="6FB84D5FFFDDFFB4FBAAF851FB10F83D" authorityName=", Sue FMNH PR" authorityYear="2081" box="[1079,1226,2020,2041]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDDFFB4FBAAF851FB10F83D" box="[1079,1226,2020,2041]" italics="true" pageId="30" pageNumber="31">Tyrannosaurus</emphasis>
</taxonomicName>
. Scale = 5 cm.
</paragraph>
</caption>
<paragraph id="A80736DCFFDCFFB5FE69FF44FC2EFE8F" blockId="31.[443,1494,240,331]" pageId="31" pageNumber="32">
Larson [
<bibRefCitation id="CC294B2DFFDCFFB5FDCAFF45FDA8FECC" author="Larson, P." bookContentInfo="Indiana University Press: Bloomington, IN, USA" box="[599,626,240,264]" editor="Parrish, J. M. &amp; Molnar, R. E. &amp; Currie, P. J. &amp; Koppelhus, E. B." pageId="31" pageNumber="32" pagination="15 - 53" refId="ref28873" refString="40. Larson, P. The case for Nanotyrannus. In Tyrannosaurid Paleobiology; Parrish, J. M., Molnar, R. E., Currie, P. J., Koppelhus, E. B., Eds.; Indiana University Press: Bloomington, IN, USA, 2013; pp. 15 - 53." title="The case for Nanotyrannus" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">40</bibRefCitation>
] also notes that patterns of pneumaticity are stable in birds as they grow, which makes differences in the presence and position of pneumatic foramina, such as the maxillary fenestra (
<figureCitation id="30832A59FFDCFFB5FD03FE86FD25FE8F" box="[670,767,307,331]" captionStart="Figure 4" captionStartId="17.[444,516,1929,1951]" captionTargetBox="[173,1417,245,1886]" captionTargetId="figure-54@17.[140,1486,209,1107]" captionTargetPageId="17" captionText="Figure 4. Left maxilla of Tyrannosaurus rex CM 9380 and right maxilla (reversed) of Nanotyrannus BMRP 2002.4.1 in lateral view, showing anatomical characters differentiating the two. Scale = 10 cm." figureDoi="http://doi.org/10.5281/zenodo.10535277" httpUri="https://zenodo.org/record/10535277/files/figure.png" pageId="31" pageNumber="32">Figure 4</figureCitation>
), difficult to explain.
</paragraph>
<paragraph id="A80736DCFFDCFFB5FE21FEDAFBE5FE43" blockId="31.[443,1496,367,1120]" box="[444,1087,367,391]" pageId="31" pageNumber="32">
<emphasis id="9ACCEACEFFDCFFB5FE21FEDAFBE5FE43" box="[444,1087,367,391]" italics="true" pageId="31" pageNumber="32">
3.4. Histology Supports Existence of Mature
<taxonomicName id="6FB84D5FFFDCFFB5FC3CFEC5FBE5FE43" box="[929,1087,368,391]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="32" phylum="Chordata" rank="genus">Nanotyrannus</taxonomicName>
</emphasis>
</paragraph>
<paragraph id="A80736DCFFDCFFB5FE21FE26FBD4FE6F" blockId="31.[443,1496,367,1120]" box="[444,1038,403,427]" pageId="31" pageNumber="32">3.4.1. Use of Histology to Test the Two Hypotheses</paragraph>
<paragraph id="A80736DCFFDCFFB5FE69FE09FCDDFD24" blockId="31.[443,1496,367,1120]" pageId="31" pageNumber="32">
If
<taxonomicName id="6FB84D5FFFDCFFB5FD90FE08FD1DFE10" box="[525,711,444,468]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="46" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDCFFB5FD90FE08FD77FE10" box="[525,685,445,468]" italics="true" pageId="31" pageNumber="32">Nanotyrannus</emphasis>
is
</taxonomicName>
a juvenile of
<taxonomicName id="6FB84D5FFFDCFFB5FCF8FE08FBD0FE10" authorityName=", Sue FMNH PR" authorityYear="2081" box="[869,1034,445,468]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="31" pageNumber="32" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDCFFB5FCF8FE08FBD0FE10" box="[869,1034,445,468]" italics="true" pageId="31" pageNumber="32">Tyrannosaurus</emphasis>
</taxonomicName>
, then all individuals showing the
<taxonomicName id="6FB84D5FFFDCFFB5FA02FE08FDF7FE31" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="32" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDCFFB5FA02FE08FDF7FE31" italics="true" pageId="31" pageNumber="32">Nanotyrannus</emphasis>
</taxonomicName>
morphology must be immature relative to
<taxonomicName id="6FB84D5FFFDCFFB5FBAAFE6BFB07FE31" authorityName=", Sue FMNH PR" authorityYear="2081" box="[1079,1245,478,501]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="31" pageNumber="32" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDCFFB5FBAAFE6BFB07FE31" box="[1079,1245,478,501]" italics="true" pageId="31" pageNumber="32">Tyrannosaurus</emphasis>
</taxonomicName>
. Bone histology can be used to infer the age and maturity of fossils, either to estimate absolute age (i.e., years of age) or relative maturity (e.g., young, rapidly growing juveniles and subadults, slower growing young adults versus old adults with slowed/ceased growth). The study of histology encompasses all aspects of bony tissue development, not simply thin sections and growth lines. To assess whether animals putatively identified as
<taxonomicName id="6FB84D5FFFDCFFB5FB21FD33FA87FD59" box="[1212,1373,646,669]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="32" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDCFFB5FB21FD33FA87FD59" box="[1212,1373,646,669]" italics="true" pageId="31" pageNumber="32">Nanotyrannus</emphasis>
</taxonomicName>
represent juveniles of
<taxonomicName id="6FB84D5FFFDCFFB5FDD4FD12FD50FD7A" box="[585,650,679,702]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="31" pageNumber="32" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
or a distinct, small-bodied tyrannosaur taxon, maturity can be assessed in at least five distinct ways:
</paragraph>
<paragraph id="A80736DCFFDCFFB5FE26FD44FCF4FCCD" blockId="31.[443,1496,367,1120]" box="[443,814,753,777]" pageId="31" pageNumber="32">(i) Patterns of skeletal fusion;</paragraph>
<paragraph id="A80736DCFFDCFFB5FE26FCA6FD35FCEE" blockId="31.[443,1496,367,1120]" box="[443,751,787,811]" pageId="31" pageNumber="32">(ii) Bone surface texture;</paragraph>
<paragraph id="A80736DCFFDCFFB5FE26FC81FB6FFC88" blockId="31.[443,1496,367,1120]" box="[443,1205,820,845]" pageId="31" pageNumber="32">
(iii) Presence/absence of an external fundamental system (
<collectionCode id="CEA9AE19FFDCFFB5FBE5FC81FB70FC88" box="[1144,1194,820,844]" pageId="31" pageNumber="32">EFS</collectionCode>
);
</paragraph>
<paragraph id="A80736DCFFDCFFB5FE26FCE3FD0FFC4B" blockId="31.[443,1496,367,1120]" pageId="31" pageNumber="32">(iv) Patterns of annual growth rates, either in terms of measures of bone deposition or kilograms of mass;</paragraph>
<paragraph id="A80736DCFFDCFFB5FE26FC2CFB52FC74" blockId="31.[443,1496,367,1120]" box="[443,1160,921,945]" pageId="31" pageNumber="32">(v) Predicted adult mass, extrapolated from growth curves.</paragraph>
<paragraph id="A80736DCFFDCFFB5FE69FC76FA8DFBA4" blockId="31.[443,1496,367,1120]" pageId="31" pageNumber="32">
<taxonomicName id="6FB84D5FFFDCFFB5FE69FC76FD55FC1E" box="[500,655,963,986]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="32" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDCFFB5FE69FC76FD55FC1E" box="[500,655,963,986]" italics="true" pageId="31" pageNumber="32">Nanotyrannus</emphasis>
</taxonomicName>
individuals show skeletal fusion and rugose facial bone, suggesting they were approaching maturity. Histology shows that
<taxonomicName id="6FB84D5FFFDCFFB5FC65FC51FB49FC3F" box="[1016,1171,996,1019]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="32" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDCFFB5FC65FC51FB49FC3F" box="[1016,1171,996,1019]" italics="true" pageId="31" pageNumber="32">Nanotyrannus</emphasis>
</taxonomicName>
individuals lack an external fundamental system, meaning that they are not old adults, but they show annual growth rates suggesting maturity. They also have predicted adult masses strongly suggestive of a distinct, small-bodied taxon rather than of juveniles of the giant
<taxonomicName id="6FB84D5FFFDCFFB5FB30FBFCFA8AFBA4" authorityName=", Sue FMNH PR" authorityYear="2081" box="[1197,1360,1097,1120]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="31" pageNumber="32" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDCFFB5FB30FBFCFA8AFBA4" box="[1197,1360,1097,1120]" italics="true" pageId="31" pageNumber="32">Tyrannosaurus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="A80736DCFFDCFFB5FE21FB31FD68FB58" blockId="31.[443,1496,1156,2058]" box="[444,690,1156,1180]" pageId="31" pageNumber="32">3.4.2. Skeletal Fusion</paragraph>
<paragraph id="A80736DCFFDCFFB5FE69FB1BFB39FA37" blockId="31.[443,1496,1156,2058]" pageId="31" pageNumber="32">
In vertebrates, composite elements such as the skull, vertebrae, shoulder girdle, sacrum, and pelvis may fuse late in development when growth slows. In crocodilians, centra and neural arches of vertebrae typically fuse late in life [
<bibRefCitation id="CC294B2DFFDCFFB5FB38FB45FB17FACC" author="Brochu, C. A." box="[1189,1229,1264,1288]" journalOrPublisher="J. Vertebr. Paleontol." pageId="31" pageNumber="32" pagination="49 - 62" part="16" refId="ref32953" refString="111. Brochu, C. A. Closure of neurocentral sutures during crocodylian ontogeny: Implication for maturity assessment in fossil archosaurs. J. Vertebr. Paleontol. 1996, 16, 49 - 62. [CrossRef]" title="Closure of neurocentral sutures during crocodylian ontogeny: Implication for maturity assessment in fossil archosaurs" type="journal article" year="1996">111</bibRefCitation>
]. In ceratopsids, skull elements and their associated osteoderms fuse late in development [
<bibRefCitation id="CC294B2DFFDCFFB5FB45FAA7FADAFAEE" author="Longrich, N. R. &amp; Field, D." box="[1240,1280,1298,1322]" journalOrPublisher="PLoS ONE" pageId="31" pageNumber="32" pagination="e 32623" part="7" refId="ref32729" refString="107. Longrich, N. R.; Field, D. Torosaurus is not Triceratops: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy. PLoS ONE 2012, 7, e 32623. [CrossRef]" title="Torosaurus is not Triceratops: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy" type="journal article" year="2012">107</bibRefCitation>
]. Which elements fuse and the sequence of fusion can vary from taxon to taxon and even individual to individual [
<bibRefCitation id="CC294B2DFFDCFFB5FDD4FAE0FDABFAA9" author="Longrich, N. R. &amp; Field, D." box="[585,625,1365,1389]" journalOrPublisher="PLoS ONE" pageId="31" pageNumber="32" pagination="e 32623" part="7" refId="ref32729" refString="107. Longrich, N. R.; Field, D. Torosaurus is not Triceratops: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy. PLoS ONE 2012, 7, e 32623. [CrossRef]" title="Torosaurus is not Triceratops: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy" type="journal article" year="2012">107</bibRefCitation>
]. Furthermore, some skull elements fuse early in ontogeny. The parietal bones, for example, are fused even in very young individuals in ceratopsids [
<bibRefCitation id="CC294B2DFFDCFFB5FAF1FAC3FA4EFA4A" author="Goodwin, M. B. &amp; Clemens, W. A. &amp; Horner, J. R. &amp; Padian, K." box="[1388,1428,1398,1422]" journalOrPublisher="J. Vertebr. Paleontol." pageId="31" pageNumber="32" pagination="103 - 112" part="26" refId="ref32994" refString="112. Goodwin, M. B.; Clemens, W. A.; Horner, J. R.; Padian, K. The smallest known Triceratops skull: New observations on ceratopsid cranial anatomy and ontogeny. J. Vertebr. Paleontol. 2006, 26, 103 - 112. [CrossRef]" title="The smallest known Triceratops skull: New observations on ceratopsid cranial anatomy and ontogeny" type="journal article" year="2006">112</bibRefCitation>
] and tyrannosaurids [
<bibRefCitation id="CC294B2DFFDCFFB5FDE3FA22FD43FA6B" author="Tsuihiji, T. &amp; Watabe, M. &amp; Tsogtbaatar, K. &amp; Tsubamoto, T. &amp; Barsbold, R. &amp; Suzuki, S. &amp; Lee, A. H. &amp; Ridgely, R. C. &amp; Kawahara, Y. &amp; Witmer, L. M." box="[638,665,1431,1455]" journalOrPublisher="J. Vertebr. Paleontol." pageId="31" pageNumber="32" pagination="497 - 517" part="31" refId="ref29685" refString="54. Tsuihiji, T.; Watabe, M.; Tsogtbaatar, K.; Tsubamoto, T.; Barsbold, R.; Suzuki, S.; Lee, A. H.; Ridgely, R. C.; Kawahara, Y.; Witmer, L. M. Cranial osteology of a juvenile specimen of Tarbosaurus bataar (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia. J. Vertebr. Paleontol. 2011, 31, 497 - 517. [CrossRef]" title="Cranial osteology of a juvenile specimen of Tarbosaurus bataar (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia" type="journal article" year="2011">54</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFDCFFB5FD3DFA2DFD12FA74" author="Sereno, P. C. &amp; Tan, L. &amp; Brusatte, S. L. &amp; Kriegstein, H. J. &amp; Zhao, X. &amp; Cloward, K." box="[672,712,1432,1456]" journalOrPublisher="Science" pageId="31" pageNumber="32" pagination="418 - 422" part="326" refId="ref33053" refString="113. Sereno, P. C.; Tan, L.; Brusatte, S. L.; Kriegstein, H. J.; Zhao, X.; Cloward, K. Tyrannosaurid skeletal design first evolved at small body size. Science 2009, 326, 418 - 422. [CrossRef] [PubMed]" title="Tyrannosaurid skeletal design first evolved at small body size" type="journal article" year="2009">113</bibRefCitation>
], and nasals are fused even in very young tyrannosaurids [
<bibRefCitation id="CC294B2DFFDCFFB5FAE2FA22FA40FA6B" author="Tsuihiji, T. &amp; Watabe, M. &amp; Tsogtbaatar, K. &amp; Tsubamoto, T. &amp; Barsbold, R. &amp; Suzuki, S. &amp; Lee, A. H. &amp; Ridgely, R. C. &amp; Kawahara, Y. &amp; Witmer, L. M." box="[1407,1434,1431,1455]" journalOrPublisher="J. Vertebr. Paleontol." pageId="31" pageNumber="32" pagination="497 - 517" part="31" refId="ref29685" refString="54. Tsuihiji, T.; Watabe, M.; Tsogtbaatar, K.; Tsubamoto, T.; Barsbold, R.; Suzuki, S.; Lee, A. H.; Ridgely, R. C.; Kawahara, Y.; Witmer, L. M. Cranial osteology of a juvenile specimen of Tarbosaurus bataar (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia. J. Vertebr. Paleontol. 2011, 31, 497 - 517. [CrossRef]" title="Cranial osteology of a juvenile specimen of Tarbosaurus bataar (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia" type="journal article" year="2011">54</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFDCFFB5FA3CFA2DFA13FA74" author="Sereno, P. C. &amp; Tan, L. &amp; Brusatte, S. L. &amp; Kriegstein, H. J. &amp; Zhao, X. &amp; Cloward, K." box="[1441,1481,1432,1456]" journalOrPublisher="Science" pageId="31" pageNumber="32" pagination="418 - 422" part="326" refId="ref33053" refString="113. Sereno, P. C.; Tan, L.; Brusatte, S. L.; Kriegstein, H. J.; Zhao, X.; Cloward, K. Tyrannosaurid skeletal design first evolved at small body size. Science 2009, 326, 418 - 422. [CrossRef] [PubMed]" title="Tyrannosaurid skeletal design first evolved at small body size" type="journal article" year="2009">113</bibRefCitation>
]. Therefore, not all fusions signal maturity. Some elements, however, only fuse in large individuals, suggesting their fusion correlates with skeletal maturity.
</paragraph>
<paragraph id="A80736DCFFDCFFB5FE69FA49FC42F97F" blockId="31.[443,1496,1156,2058]" pageId="31" pageNumber="32">
The scapula and coracoid fuse appear to fuse late in many dinosaurs, including
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<emphasis id="9ACCEACEFFDCFFB5FA38FA48FDF6F9F1" italics="true" pageId="31" pageNumber="32">Herrerasaurus</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFDCFFB5FDA6F9ABFDB9F9F2" author="Sereno, P. C." box="[571,611,1566,1590]" journalOrPublisher="J. Vertebr. Paleontol." pageId="31" pageNumber="32" pagination="425 - 450" part="13" refId="ref33115" refString="114. Sereno, P. C. The pectoral girdle and forelimb of the basal theropod Herrerasaurus ischigualastensis. J. Vertebr. Paleontol. 1994, 13, 425 - 450. [CrossRef]" title="The pectoral girdle and forelimb of the basal theropod Herrerasaurus ischigualastensis" type="journal article" year="1994">114</bibRefCitation>
],
<taxonomicName id="6FB84D5FFFDCFFB5FDE5F9A8FCC3F9F1" box="[632,793,1565,1589]" family="Abelisauridae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="32" phylum="Chordata" rank="family">Abelisauridae</taxonomicName>
[
<bibRefCitation id="CC294B2DFFDCFFB5FCBAF9A8FC95F9F1" author="Burch, S. H. &amp; Carrano, M. T." box="[807,847,1565,1589]" journalOrPublisher="J. Vertebr. Paleontol." pageId="31" pageNumber="32" pagination="1 - 16" part="32" refId="ref33152" refString="115. Burch, S. H.; Carrano, M. T. An articulated pectoral girdle and forelimb of the abelisaurid theropod Majungasaurus crenatissimus from the Late Cretaceous of Madagascar. J. Vertebr. Paleontol. 2012, 32, 1 - 16. [CrossRef]" title="An articulated pectoral girdle and forelimb of the abelisaurid theropod Majungasaurus crenatissimus from the Late Cretaceous of Madagascar" type="journal article" year="2012">115</bibRefCitation>
], and at least some dromaeosaurs, including
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<emphasis id="9ACCEACEFFDCFFB5FACEF9A8FD9FF993" italics="true" pageId="31" pageNumber="32">Velociraptor mongoliensis</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFDCFFB5FDCEF98AFDA1F993" author="Norell, M. A. &amp; Makovicky, P. J." box="[595,635,1599,1623]" journalOrPublisher="Am. Mus. Novit." pageId="31" pageNumber="32" pagination="1 - 45" part="3282" refId="ref33203" refString="116. Norell, M. A.; Makovicky, P. J. Important features of the dromaeosaurid skeleton II: Information from newly collected specimens of Velociraptor mongoliensis. Am. Mus. Novit. 1999, 3282, 1 - 45." title="Important features of the dromaeosaurid skeleton II: Information from newly collected specimens of Velociraptor mongoliensis" type="journal article" year="1999">116</bibRefCitation>
] and
<taxonomicName id="6FB84D5FFFDCFFB5FD27F98AFC75F993" box="[698,943,1599,1623]" family="Dromaeosauridae" genus="Achillobator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="32" phylum="Chordata" rank="species" species="giganticus">
<emphasis id="9ACCEACEFFDCFFB5FD27F98AFC75F993" box="[698,943,1599,1623]" italics="true" pageId="31" pageNumber="32">Achillobator giganticus</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFDCFFB5FC23F98AFC3CF993" author="Perle, A. &amp; Norell, M. A. &amp; Clark, J. M." box="[958,998,1599,1623]" journalOrPublisher="Dep. Geol. Natl. Univ. Mong. Ulaanbaatar" pageId="31" pageNumber="32" pagination="1 - 105" part="101" refId="ref33249" refString="117. Perle, A.; Norell, M. A.; Clark, J. M. A new maniraptoran theropod Achillobator giganticus (Dromaeosauridae) from the Upper Cretaceous of Burkhant, Mongolia. Dep. Geol. Natl. Univ. Mong. Ulaanbaatar 1999, 101, 1 - 105." title="A new maniraptoran theropod Achillobator giganticus (Dromaeosauridae) from the Upper Cretaceous of Burkhant, Mongolia" type="journal article" year="1999">117</bibRefCitation>
]. Fusion of the scapulocoracoid also occurs in tyrannosaurs. Partial scapulocoracoid fusion is seen in
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<emphasis id="9ACCEACEFFDCFFB5FBEEF9D5FA46F9BC" box="[1139,1436,1632,1656]" italics="true" pageId="31" pageNumber="32">Albertosaurus sarcophagus</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFDCFFB5FA31F9D4FA1DF9BD" author="Mallon, J. C. &amp; Bura, J. R. &amp; Schumann, D. &amp; Currie, P. J." box="[1452,1479,1633,1657]" journalOrPublisher="Anat. Rec." pageId="31" pageNumber="32" pagination="673 - 690" part="303" refId="ref30780" refString="71. Mallon, J. C.; Bura, J. R.; Schumann, D.; Currie, P. J. A problematic tyrannosaurid (Dinosauria: Theropoda) skeleton and its implications for tyrannosaurid diversity in the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta. Anat. Rec. 2020, 303, 673 - 690. [CrossRef]" title="A problematic tyrannosaurid (Dinosauria: Theropoda) skeleton and its implications for tyrannosaurid diversity in the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta" type="journal article" year="2020">71</bibRefCitation>
]; complete fusion is seen in a large
<taxonomicName id="6FB84D5FFFDCFFB5FCC3F936FC7BF95E" box="[862,929,1666,1690]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="31" pageNumber="32" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFDCFFB5FCC3F936FCB5F95E" box="[862,879,1667,1690]" italics="true" pageId="31" pageNumber="32">T</emphasis>
.
<emphasis id="9ACCEACEFFDCFFB5FCE2F936FC7BF95E" box="[895,929,1667,1690]" italics="true" pageId="31" pageNumber="32">rex</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFDCFFB5FC2EF937FC1AF95D" author="Brochu, C. A." box="[947,960,1666,1689]" journalOrPublisher="J. Vertebr. Paleontol. Mem." pageId="31" pageNumber="32" pagination="1 - 138" part="7" refId="ref27308" refString="7. Brochu, C. A. Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. J. Vertebr. Paleontol. Mem. 2003, 7, 1 - 138. [CrossRef]" title="Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">7</bibRefCitation>
] and a
<taxonomicName id="6FB84D5FFFDCFFB5FBBDF936FB1CF95E" authorityName=", Sue FMNH PR" authorityYear="2081" box="[1056,1222,1667,1690]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="31" pageNumber="32" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDCFFB5FBBDF936FB1CF95E" box="[1056,1222,1667,1690]" italics="true" pageId="31" pageNumber="32">Tyrannosaurus</emphasis>
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from the Naashoibito member of the Kirtland Formation [
<bibRefCitation id="CC294B2DFFDCFFB5FCFCF911FC53F978" author="Jasinski, S. E. &amp; Sullivan, R. M. &amp; Lucas, S. G." box="[865,905,1700,1724]" journalOrPublisher="New Mex. Mus. Nat. Hist. Sci. Bull." pageId="31" pageNumber="32" pagination="216 - 265" part="53" refId="ref33306" refString="118. Jasinski, S. E.; Sullivan, R. M.; Lucas, S. G. Taxonomic composition of the Alamo Wash local fauna from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. New Mex. Mus. Nat. Hist. Sci. Bull. 2011, 53, 216 - 265." title="Taxonomic composition of the Alamo Wash local fauna from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico" type="journal article" year="2011">118</bibRefCitation>
].
</paragraph>
<paragraph id="A80736DCFFDCFFB5FE69F970FAF0F8A7" blockId="31.[443,1496,1156,2058]" pageId="31" pageNumber="32">
In some theropods, the pelvis shows partial or complete fusion in large individuals. The pubis and ilium fuse in the microraptorine
<taxonomicName id="6FB84D5FFFDCFFB5FBABF953FA86F93A" authorityName="Longrich &amp; Currie" authorityYear="2009" box="[1078,1372,1766,1790]" family="Dromaeosauridae" genus="Hesperonychus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="32" phylum="Chordata" rank="species" species="elizabethae">
<emphasis id="9ACCEACEFFDCFFB5FBABF953FA86F93A" box="[1078,1372,1766,1790]" italics="true" pageId="31" pageNumber="32">Hesperonychus elizabethae</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFDCFFB5FAF0F952FA4FF93B" author="Longrich, N. R. &amp; Currie, P. J." box="[1389,1429,1767,1791]" journalOrPublisher="PNAS" pageId="31" pageNumber="32" pagination="5002 - 5008" part="106" refId="ref33376" refString="119. Longrich, N. R.; Currie, P. J. A microraptorine (Dinosauria-Dromaeosauridae) from the Late Cretaceous of North America. PNAS 2009, 106, 5002 - 5008. [CrossRef] [PubMed]" title="A microraptorine (Dinosauria-Dromaeosauridae) from the Late Cretaceous of North America" type="journal article" year="2009">119</bibRefCitation>
]; the ilium, ischium, and pubis fuse in Coelophysoidea [
<bibRefCitation id="CC294B2DFFDCFFB5FB88F8BDFBE7F8E4" author="Raath, M. A. &amp; Carpenter, K. &amp; Currie, P. J." bookContentInfo="Cambridge University Press: Cambridge, UK" box="[1045,1085,1800,1824]" editor="Currie, P. J. &amp; Carpenter, K." pageId="31" pageNumber="32" pagination="91 - 105" refId="ref33419" refString="120. Raath, M. A.; Carpenter, K.; Currie, P. J. Morphological variation in small theropods and its meaning in systematics: Evidence from Syntarsus. In Dinosaur Systematics: Perspectives and Approaches; Currie, P. J., Carpenter, K., Eds.; Cambridge University Press: Cambridge, UK, 1990; pp. 91 - 105." title="Morphological variation in small theropods and its meaning in systematics: Evidence from Syntarsus" type="book chapter" volumeTitle="Dinosaur Systematics: Perspectives and Approaches" year="1990">120</bibRefCitation>
],
<taxonomicName id="6FB84D5FFFDCFFB5FBCEF8BDFB23F8E4" box="[1107,1273,1800,1824]" family="Abelisauridae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="32" phylum="Chordata" rank="family">Abelisauridae</taxonomicName>
[
<bibRefCitation id="CC294B2DFFDCFFB5FA94F8BDFAEBF8E4" author="Iori, F. V. &amp; de Araujo-Junior, H. I. &amp; Tavares, S. A. S. &amp; da Silva Marinho, T. &amp; Martinelli, A. G." box="[1289,1329,1800,1824]" journalOrPublisher="J. South Am. Earth Sci." pageId="31" pageNumber="32" pagination="103551" part="112" refId="ref33493" refString="121. Iori, F. V.; de Araujo-Junior, H. I.; Tavares, S. A. S.; da Silva Marinho, T.; Martinelli, A. G. New theropod dinosaur from the Late Cretaceous of Brazil improves abelisaurid diversity. J. South Am. Earth Sci. 2021, 112, 103551. [CrossRef]" title="New theropod dinosaur from the Late Cretaceous of Brazil improves abelisaurid diversity" type="journal article" year="2021">121</bibRefCitation>
<bibRefCitation id="CC294B2DFFDCFFB5FAA3F8BDFABCF8E4" author="Cerroni, M. A. &amp; Baiano, M. A. &amp; Canale, J. I. &amp; Agnolin, F. L. &amp; Otero, A. &amp; Novas, F. E." box="[1342,1382,1800,1824]" journalOrPublisher="J. Syst. Palaeontol." pageId="31" pageNumber="32" pagination="1 - 32" part="20" refId="ref33615" refString="123. Cerroni, M. A.; Baiano, M. A.; Canale, J. I.; Agnolin, F. L.; Otero, A.; Novas, F. E. Appendicular osteology of Skorpiovenator bustingorryi (Theropoda, Abelisauridae) with comments on phylogenetic features of abelisaurids. J. Syst. Palaeontol. 2022, 20, 1 - 32. [CrossRef]" title="Appendicular osteology of Skorpiovenator bustingorryi (Theropoda, Abelisauridae) with comments on phylogenetic features of abelisaurids" type="journal article" year="2022">123</bibRefCitation>
], and
<taxonomicName id="6FB84D5FFFDCFFB5FA2DF8BDFDBBF885" family="Ornithomimidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="32" phylum="Chordata" rank="family">Ornithomimidae</taxonomicName>
[
<bibRefCitation id="CC294B2DFFDCFFB5FDEDF89CFD42F885" author="Macdonald, I. &amp; Currie, P. J." box="[624,664,1833,1857]" journalOrPublisher="Can. J. Earth Sci." pageId="31" pageNumber="32" pagination="129 - 157" part="56" refId="ref33691" refString="124. Macdonald, I.; Currie, P. J. Description of a partial Dromiceiomimus (Dinosauria: Theropoda) skeleton with comments on the validity of the genus. Can. J. Earth Sci. 2019, 56, 129 - 157. [CrossRef]" title="Description of a partial Dromiceiomimus (Dinosauria: Theropoda) skeleton with comments on the validity of the genus" type="journal article" year="2019">124</bibRefCitation>
<bibRefCitation id="CC294B2DFFDCFFB5FD38F89CFD17F885" author="Xu, L. &amp; Kobayashi, Y. &amp; Lu, J. &amp; Lee, Y. - N. &amp; Liu, Y. &amp; Tanaka, K. &amp; Zhang, X. &amp; Jia, S. &amp; Zhang, J." box="[677,717,1833,1857]" journalOrPublisher="Cretac. Res." pageId="31" pageNumber="32" pagination="213 - 222" part="32" refId="ref33832" refString="127. Xu, L.; Kobayashi, Y.; Lu, J.; Lee, Y. - N.; Liu, Y.; Tanaka, K.; Zhang, X.; Jia, S.; Zhang, J. A new ornithomimid dinosaur with North American affinities from the Late Cretaceous Qiupa Formation in Henan Province of China. Cretac. Res. 2011, 32, 213 - 222. [CrossRef]" title="A new ornithomimid dinosaur with North American affinities from the Late Cretaceous Qiupa Formation in Henan Province of China" type="journal article" year="2011">127</bibRefCitation>
]. Fusion of the pubis and ischium also occurs in a large individual of
<taxonomicName id="6FB84D5FFFDCFFB5FE47F8F9FDC2F8A7" box="[474,536,1867,1891]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="31" pageNumber="32" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFDCFFB5FE47F8F9FE30F8A7" box="[474,490,1868,1891]" italics="true" pageId="31" pageNumber="32">T</emphasis>
.
<emphasis id="9ACCEACEFFDCFFB5FE6AF8F9FDC2F8A7" box="[503,536,1868,1891]" italics="true" pageId="31" pageNumber="32">rex</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFDCFFB5FDB5F8FEFDEFF8A6" author="Brochu, C. A." box="[552,565,1867,1890]" journalOrPublisher="J. Vertebr. Paleontol. Mem." pageId="31" pageNumber="32" pagination="1 - 138" part="7" refId="ref27308" refString="7. Brochu, C. A. Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull. J. Vertebr. Paleontol. Mem. 2003, 7, 1 - 138. [CrossRef]" title="Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" type="journal article" year="2003">7</bibRefCitation>
]; the pubes and ischia are fused in
<taxonomicName id="6FB84D5FFFDCFFB5FC4FF8FEFB00F8A7" box="[978,1242,1867,1891]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="31" pageNumber="32" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFDCFFB5FC4FF8FEFBA3F8A7" box="[978,1145,1867,1891]" italics="true" pageId="31" pageNumber="32">Daspletosaurus</emphasis>
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</taxonomicName>
52981.
</paragraph>
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While not all skeletal fusions are correlated with maturity, fusion of the vertebrae, pectoral girdle, and pelvic girdle do seem to correlate with maturity. Strikingly, a number of fusions occur in
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<emphasis id="9ACCEACEFFDCFFB5FD0BF805FCEBF803" box="[662,817,1968,1991]" italics="true" pageId="31" pageNumber="32">Nanotyrannus</emphasis>
<collectionCode id="CEA9AE19FFDCFFB5FCA5F805FC58F803" box="[824,898,1968,1991]" collectionName="BMRP" pageId="31" pageNumber="32">BMRP</collectionCode>
</taxonomicName>
<date id="DC06101CFFDCFFB5FC15F81AFC39F803" box="[904,995,1967,1991]" pageId="31" pageNumber="32" value="2002-04-01">2002.4.1</date>
[
<bibRefCitation id="CC294B2DFFDCFFB5FC6EF81AFBD4F803" author="Larson, P." bookContentInfo="Indiana University Press: Bloomington, IN, USA" box="[1011,1038,1967,1991]" editor="Parrish, J. M. &amp; Molnar, R. E. &amp; Currie, P. J. &amp; Koppelhus, E. B." pageId="31" pageNumber="32" pagination="15 - 53" refId="ref28873" refString="40. Larson, P. The case for Nanotyrannus. In Tyrannosaurid Paleobiology; Parrish, J. M., Molnar, R. E., Currie, P. J., Koppelhus, E. B., Eds.; Indiana University Press: Bloomington, IN, USA, 2013; pp. 15 - 53." title="The case for Nanotyrannus" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">40</bibRefCitation>
]. These include fusion or partial fusion of neural arches to centra, fusion of the scapulocoracoid, and fusion of the ilium, pubis, and ischium [
<bibRefCitation id="CC294B2DFFDCFFB5FDF8F847FD5AF7CE" author="Larson, P." bookContentInfo="Indiana University Press: Bloomington, IN, USA" box="[613,640,2034,2058]" editor="Parrish, J. M. &amp; Molnar, R. E. &amp; Currie, P. J. &amp; Koppelhus, E. B." pageId="31" pageNumber="32" pagination="15 - 53" refId="ref28873" refString="40. Larson, P. The case for Nanotyrannus. In Tyrannosaurid Paleobiology; Parrish, J. M., Molnar, R. E., Currie, P. J., Koppelhus, E. B., Eds.; Indiana University Press: Bloomington, IN, USA, 2013; pp. 15 - 53." title="The case for Nanotyrannus" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">40</bibRefCitation>
]. This degree of skeletal fusion is consistent with the animal being a nearly full-sized subadult or early adult [
<bibRefCitation id="CC294B2DFFE3FF8AFC2EFF45FC14FECC" author="Larson, P." bookContentInfo="Indiana University Press: Bloomington, IN, USA" box="[947,974,240,264]" editor="Parrish, J. M. &amp; Molnar, R. E. &amp; Currie, P. J. &amp; Koppelhus, E. B." pageId="32" pageNumber="33" pagination="15 - 53" refId="ref28873" refString="40. Larson, P. The case for Nanotyrannus. In Tyrannosaurid Paleobiology; Parrish, J. M., Molnar, R. E., Currie, P. J., Koppelhus, E. B., Eds.; Indiana University Press: Bloomington, IN, USA, 2013; pp. 15 - 53." title="The case for Nanotyrannus" type="book chapter" volumeTitle="Tyrannosaurid Paleobiology" year="2013">40</bibRefCitation>
]. Further study of skeletal fusion patterns is needed for tyrannosaurs (and dinosaurs more generally), but evidence from skeletal fusions suggests that
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<emphasis id="9ACCEACEFFE3FF8AFD25FE81FC8CFE8F" box="[696,854,308,331]" italics="true" pageId="32" pageNumber="33">Nanotyrannus</emphasis>
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are not juveniles of
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<emphasis id="9ACCEACEFFE3FF8AFBDFFE81FB3FFE8F" box="[1090,1253,308,331]" italics="true" pageId="32" pageNumber="33">Tyrannosaurus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="A80736DCFFE3FF8AFE21FEDAFD6FFE42" blockId="32.[443,1497,367,1370]" box="[444,693,367,391]" pageId="32" pageNumber="33">3.4.3. Surface Texture</paragraph>
<paragraph id="A80736DCFFE3FF8AFE69FE2CFCD1FD5F" blockId="32.[443,1497,367,1370]" pageId="32" pageNumber="33">
In many dinosaurs, the adult skull bones take on a rugose to gnarled surface texture and may develop sculpturing. In chasmosaurine ceratopsians, for example, juveniles and subadults have smooth, striated skull bones. In adults, the bone takes on a gnarled texture, resembling tree bark, often with extensive, high-relief rugosity [
<bibRefCitation id="CC294B2DFFE3FF8AFB30FE48FB0FFDD1" author="Longrich, N. R. &amp; Field, D." box="[1197,1237,509,533]" journalOrPublisher="PLoS ONE" pageId="32" pageNumber="33" pagination="e 32623" part="7" refId="ref32729" refString="107. Longrich, N. R.; Field, D. Torosaurus is not Triceratops: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy. PLoS ONE 2012, 7, e 32623. [CrossRef]" title="Torosaurus is not Triceratops: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy" type="journal article" year="2012">107</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE3FF8AFB41FE48FADEFDD1" author="Brown, C. M. &amp; Russell, A. P. &amp; Ryan, M. J." box="[1244,1284,509,533]" journalOrPublisher="J. Vertebr. Paleontol." pageId="32" pageNumber="33" pagination="132 - 141" part="29" refId="ref33916" refString="128. Brown, C. M.; Russell, A. P.; Ryan, M. J. Pattern and transition of surficial bone texture of the centrosaurine frill and their ontogenetic and taxonomic implications. J. Vertebr. Paleontol. 2009, 29, 132 - 141. [CrossRef]" title="Pattern and transition of surficial bone texture of the centrosaurine frill and their ontogenetic and taxonomic implications" type="journal article" year="2009">128</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE3FF8AFA97FE48FAE8FDD1" author="Sampson, S. D. &amp; Ryan, M. J. &amp; Tanke, D. H." box="[1290,1330,509,533]" journalOrPublisher="Zooogical J. Linn. Soc." pageId="32" pageNumber="33" pagination="293 - 337" part="121" refId="ref33973" refString="129. Sampson, S. D.; Ryan, M. J.; Tanke, D. H. Craniofacial ontogeny in centrosaurine dinosaurs (Ornithischia: Ceratopsidae): Taxonomic and behavioral implications. Zooogical J. Linn. Soc. 1997, 121, 293 - 337. [CrossRef]" title="Craniofacial ontogeny in centrosaurine dinosaurs (Ornithischia: Ceratopsidae): Taxonomic and behavioral implications" type="journal article" year="1997">129</bibRefCitation>
] and grooves for blood vessels. The appearance of rugose bone texture can be used as a rough proxy for maturity in
<taxonomicName id="6FB84D5FFFE3FF8AFDD6FDF5FD37FD9C" box="[587,749,576,600]" family="Ceratopsidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="family">Ceratopsidae</taxonomicName>
. Striated bone is not seen in the very oldest individuals but is seen in very large individuals of
<taxonomicName id="6FB84D5FFFE3FF8AFCA6FDD7FC68FDBD" authorityName="Marsh" authorityYear="1891" box="[827,946,610,633]" family="Ceratopsidae" genus="Torosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE3FF8AFCA6FDD7FC68FDBD" box="[827,946,610,633]" italics="true" pageId="32" pageNumber="33">Torosaurus</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFE3FF8AFC5FFDD4FC30FDBD" author="Longrich, N. R. &amp; Field, D." box="[962,1002,609,633]" journalOrPublisher="PLoS ONE" pageId="32" pageNumber="33" pagination="e 32623" part="7" refId="ref32729" refString="107. Longrich, N. R.; Field, D. Torosaurus is not Triceratops: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy. PLoS ONE 2012, 7, e 32623. [CrossRef]" title="Torosaurus is not Triceratops: Ontogeny in Chasmosaurine Ceratopsids as a Case Study in Dinosaur Taxonomy" type="journal article" year="2012">107</bibRefCitation>
], showing that it persists relatively late in subadults and young adults.
</paragraph>
<paragraph id="A80736DCFFE3FF8AFE69FD11FD38FC40" blockId="32.[443,1497,367,1370]" pageId="32" pageNumber="33">
A similar pattern is seen in tyrannosaurids. In
<taxonomicName id="6FB84D5FFFE3FF8AFBB4FD11FB6CFD7F" authorityName="Lambe" authorityYear="1914" box="[1065,1206,676,699]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE3FF8AFBB4FD11FB6CFD7F" box="[1065,1206,676,699]" italics="true" pageId="32" pageNumber="33">Gorgosaurus</emphasis>
</taxonomicName>
, nasals [
<bibRefCitation id="CC294B2DFFE3FF8AFA82FD11FAF6FD78" author="Carr, T. D." box="[1311,1324,676,700]" journalOrPublisher="J. Vertebr. Paleontol." pageId="32" pageNumber="33" pagination="497 - 520" part="19" refId="ref27419" refString="9. Carr, T. D. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). J. Vertebr. Paleontol. 1999, 19, 497 - 520. [CrossRef]" title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">9</bibRefCitation>
], maxillae [
<bibRefCitation id="CC294B2DFFE3FF8AFA27FD11FA1DFD78" author="Carr, T. D." box="[1466,1479,676,700]" journalOrPublisher="J. Vertebr. Paleontol." pageId="32" pageNumber="33" pagination="497 - 520" part="19" refId="ref27419" refString="9. Carr, T. D. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). J. Vertebr. Paleontol. 1999, 19, 497 - 520. [CrossRef]" title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">9</bibRefCitation>
], and postorbitals [
<bibRefCitation id="CC294B2DFFE3FF8AFD09FD73FD66FD1A" author="Voris, J. T. &amp; Zelenitsky, D. K. &amp; Therrien, F. &amp; Currie, P. J." box="[660,700,710,734]" journalOrPublisher="Sci. Rep." pageId="32" pageNumber="33" pagination="1 - 10" part="9" refId="ref32886" refString="110. Voris, J. T.; Zelenitsky, D. K.; Therrien, F.; Currie, P. J. Reassessment of a juvenile Daspletosaurus from the Late Cretaceous of Alberta, Canada with implications for the identification of immature tyrannosaurids. Sci. Rep. 2019, 9, 1 - 10. [CrossRef] [PubMed]" title="Reassessment of a juvenile Daspletosaurus from the Late Cretaceous of Alberta, Canada with implications for the identification of immature tyrannosaurids" type="journal article" year="2019">110</bibRefCitation>
] are relatively smooth in juveniles, and become more rugose in subadults and adults. A similar pattern occurs in postorbitals of
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<emphasis id="9ACCEACEFFE3FF8AFB79FD52FA54FD3B" box="[1252,1422,743,767]" italics="true" pageId="32" pageNumber="33">Daspletosaurus</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFE3FF8AFA3DFD52FA12FD3B" author="Voris, J. T. &amp; Zelenitsky, D. K. &amp; Therrien, F. &amp; Currie, P. J." box="[1440,1480,743,767]" journalOrPublisher="Sci. Rep." pageId="32" pageNumber="33" pagination="1 - 10" part="9" refId="ref32886" refString="110. Voris, J. T.; Zelenitsky, D. K.; Therrien, F.; Currie, P. J. Reassessment of a juvenile Daspletosaurus from the Late Cretaceous of Alberta, Canada with implications for the identification of immature tyrannosaurids. Sci. Rep. 2019, 9, 1 - 10. [CrossRef] [PubMed]" title="Reassessment of a juvenile Daspletosaurus from the Late Cretaceous of Alberta, Canada with implications for the identification of immature tyrannosaurids" type="journal article" year="2019">110</bibRefCitation>
]. Young
<taxonomicName id="6FB84D5FFFE3FF8AFD92FCBDFD4CFCE4" box="[527,662,776,800]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE3FF8AFD92FCBDFD4CFCE4" box="[527,662,776,800]" italics="true" pageId="32" pageNumber="33">Tarbosaurus</emphasis>
</taxonomicName>
show weak sculpturing of the maxilla, while nasals and lacrimals are almost smooth [
<bibRefCitation id="CC294B2DFFE3FF8AFDE6FC9FFD4CFC86" author="Tsuihiji, T. &amp; Watabe, M. &amp; Tsogtbaatar, K. &amp; Tsubamoto, T. &amp; Barsbold, R. &amp; Suzuki, S. &amp; Lee, A. H. &amp; Ridgely, R. C. &amp; Kawahara, Y. &amp; Witmer, L. M." box="[635,662,810,834]" journalOrPublisher="J. Vertebr. Paleontol." pageId="32" pageNumber="33" pagination="497 - 517" part="31" refId="ref29685" refString="54. Tsuihiji, T.; Watabe, M.; Tsogtbaatar, K.; Tsubamoto, T.; Barsbold, R.; Suzuki, S.; Lee, A. H.; Ridgely, R. C.; Kawahara, Y.; Witmer, L. M. Cranial osteology of a juvenile specimen of Tarbosaurus bataar (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia. J. Vertebr. Paleontol. 2011, 31, 497 - 517. [CrossRef]" title="Cranial osteology of a juvenile specimen of Tarbosaurus bataar (Theropoda, Tyrannosauridae) from the Nemegt Formation (Upper Cretaceous) of Bugin Tsav, Mongolia" type="journal article" year="2011">54</bibRefCitation>
]; smooth facial bones are seen in another juvenile tyrannosaurine, the
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of “
<taxonomicName id="6FB84D5FFFE3FF8AFDCCFCF9FCFEFCA7" box="[593,804,843,867]" class="Reptilia" family="Tyrannosauridae" genus="Raptorex" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="kriegsteini">
<emphasis id="9ACCEACEFFE3FF8AFDCCFCF9FCFEFCA7" box="[593,804,843,867]" italics="true" pageId="32" pageNumber="33">Raptorex kriegsteini</emphasis>
</taxonomicName>
” [
<bibRefCitation id="CC294B2DFFE3FF8AFCDDFCF9FCB2FCA0" author="Sereno, P. C. &amp; Tan, L. &amp; Brusatte, S. L. &amp; Kriegstein, H. J. &amp; Zhao, X. &amp; Cloward, K." box="[832,872,844,868]" journalOrPublisher="Science" pageId="32" pageNumber="33" pagination="418 - 422" part="326" refId="ref33053" refString="113. Sereno, P. C.; Tan, L.; Brusatte, S. L.; Kriegstein, H. J.; Zhao, X.; Cloward, K. Tyrannosaurid skeletal design first evolved at small body size. Science 2009, 326, 418 - 422. [CrossRef] [PubMed]" title="Tyrannosaurid skeletal design first evolved at small body size" type="journal article" year="2009">113</bibRefCitation>
], likely a juvenile
<taxonomicName id="6FB84D5FFFE3FF8AFBAAFCFEFB62FCA7" box="[1079,1208,843,867]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE3FF8AFBAAFCFEFB62FCA7" box="[1079,1208,843,867]" italics="true" pageId="32" pageNumber="33">Tarbosaurus</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFE3FF8AFB5BFCFEFB3BFCA7" author="Fowler, D. W. &amp; Woodward, H. N. &amp; Freedman, E. A. &amp; Larson, P. L. &amp; Horner, J. R." box="[1222,1249,843,867]" journalOrPublisher="PLoS ONE" pageId="32" pageNumber="33" pagination="e 21376" part="6" refId="ref29785" refString="55. Fowler, D. W.; Woodward, H. N.; Freedman, E. A.; Larson, P. L.; Horner, J. R. Reanalysis of &quot; Raptorex kriegsteini &quot;: A juvenile tyrannosaurid dinosaur from Mongolia. PLoS ONE 2011, 6, e 21376. [CrossRef]" title="Reanalysis of &quot; Raptorex kriegsteini &quot;: A juvenile tyrannosaurid dinosaur from Mongolia" type="journal article" year="2011">55</bibRefCitation>
]. Adults have highly rugose facial bones [
<bibRefCitation id="CC294B2DFFE3FF8AFD36FCD8FD09FC41" author="Maleev, E. A." box="[683,723,877,901]" journalOrPublisher="Dokl. Akad. Nauk. SSSR" pageId="32" pageNumber="33" pagination="779 - 782" part="104" refId="ref32390" refString="101. Maleev, E. A. New carnivorous dinosaurs from the Upper Cretaceous of Mongolia. Dokl. Akad. Nauk. SSSR 1955, 104, 779 - 782." title="New carnivorous dinosaurs from the Upper Cretaceous of Mongolia" type="journal article" year="1955">101</bibRefCitation>
].
</paragraph>
<paragraph id="A80736DCFFE3FF8AFE69FC3BFA03FBAB" blockId="32.[443,1497,367,1370]" pageId="32" pageNumber="33">
These patterns are hard to quantify or characterize objectively, but overall, it appears that rugosity of facial elements increases as animals mature, providing a rough proxy for maturity. As in
<taxonomicName id="6FB84D5FFFE3FF8AFD02FC64FC9AFC2D" box="[671,832,977,1001]" family="Ceratopsidae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="family">Ceratopsidae</taxonomicName>
, striated bone persists relatively late in ontogeny, being seen in subadult
<taxonomicName id="6FB84D5FFFE3FF8AFD11FC46FCDAFBCF" authorityName="Kurzanov" authorityYear="1976" box="[652,768,1011,1035]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="32" pageNumber="33" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE3FF8AFD11FC46FCDAFBCF" box="[652,768,1011,1035]" italics="true" pageId="32" pageNumber="33">Alioramus</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFE3FF8AFC8CFC46FCF6FBCF" author="Brusatte, S. L. &amp; Carr, T. D. &amp; Norell, M. A." box="[785,812,1011,1035]" journalOrPublisher="Bull. Am. Mus. Nat. Hist." pageId="32" pageNumber="33" pagination="1 - 197" part="2012" refId="ref30715" refString="70. Brusatte, S. L.; Carr, T. D.; Norell, M. A. The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bull. Am. Mus. Nat. Hist. 2012, 2012, 1 - 197. [CrossRef]" title="The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia" type="journal article" year="2012">70</bibRefCitation>
], young adult
<taxonomicName id="6FB84D5FFFE3FF8AFC7CFC46FBB4FBCE" authorityName="Lambe" authorityYear="1914" box="[993,1134,1011,1034]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE3FF8AFC7CFC46FBB4FBCE" box="[993,1134,1011,1034]" italics="true" pageId="32" pageNumber="33">Gorgosaurus</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFE3FF8AFB1DFC46FB72FBCF" author="Voris, J. T. &amp; Zelenitsky, D. K. &amp; Therrien, F. &amp; Ridgely, R. C. &amp; Currie, P. J. &amp; Witmer, L. M." box="[1152,1192,1011,1035]" journalOrPublisher="J. Vertebr. Paleontol." pageId="32" pageNumber="33" pagination="2041651" part="41" refId="ref32804" refString="109. Voris, J. T.; Zelenitsky, D. K.; Therrien, F.; Ridgely, R. C.; Currie, P. J.; Witmer, L. M. Two exceptionally preserved juvenile specimens of Gorgosaurus libratus (Tyrannosauridae, Albertosaurinae) provide new insight into the timing of ontogenetic changes in tyrannosaurids. J. Vertebr. Paleontol. 2021, 41, e 2041651. [CrossRef]" title="Two exceptionally preserved juvenile specimens of Gorgosaurus libratus (Tyrannosauridae, Albertosaurinae) provide new insight into the timing of ontogenetic changes in tyrannosaurids" type="journal article" year="2021">109</bibRefCitation>
], and in the
<typeStatus id="7703887EFFE3FF8AFAD8FC46FA5CFBCF" box="[1349,1414,1011,1035]" pageId="32" pageNumber="33">types</typeStatus>
of the tyrannosaurines
<taxonomicName id="6FB84D5FFFE3FF8AFD1AFBA1FCF8FBE8" authorityName="Carr &amp; Williamson" authorityYear="2010" box="[647,802,1044,1068]" class="Reptilia" genus="Bistahieversor" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE3FF8AFD1AFBA1FCF8FBE8" box="[647,802,1044,1068]" italics="true" pageId="32" pageNumber="33">Bistahieversor</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFE3FF8AFCAEFBA1FC94FBE8" author="Carr, T. D. &amp; Williamson, T. E." box="[819,846,1044,1068]" journalOrPublisher="J. Vertebr. Paleontol." pageId="32" pageNumber="33" pagination="1 - 16" part="30" refId="ref30052" refString="59. Carr, T. D.; Williamson, T. E. Bistahieversor sealeyi, gen. et sp. nov., a new tyrannosauroid from New Mexico and the origin of deep snouts in Tyrannosauroidea. J. Vertebr. Paleontol. 2010, 30, 1 - 16. [CrossRef]" title="Bistahieversor sealeyi, gen. et sp. nov., a new tyrannosauroid from New Mexico and the origin of deep snouts in Tyrannosauroidea." type="journal article" year="2010">59</bibRefCitation>
] and
<taxonomicName id="6FB84D5FFFE3FF8AFC08FBA1FB91FBE8" authorityName="Voris, Therrien, Zelenitsky &amp; Brown" authorityYear="2020" box="[917,1099,1044,1068]" class="Reptilia" family="Tyrannosauridae" genus="Thanatotheristes" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE3FF8AFC08FBA1FB91FBE8" box="[917,1099,1044,1068]" italics="true" pageId="32" pageNumber="33">Thanatotheristes</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFE3FF8AFBC1FBA1FB5EFBE8" author="Voris, J. T. &amp; Zelenitsky, D. K. &amp; Therrien, F. &amp; Currie, P. J." box="[1116,1156,1044,1068]" journalOrPublisher="Sci. Rep." pageId="32" pageNumber="33" pagination="1 - 10" part="9" refId="ref32886" refString="110. Voris, J. T.; Zelenitsky, D. K.; Therrien, F.; Currie, P. J. Reassessment of a juvenile Daspletosaurus from the Late Cretaceous of Alberta, Canada with implications for the identification of immature tyrannosaurids. Sci. Rep. 2019, 9, 1 - 10. [CrossRef] [PubMed]" title="Reassessment of a juvenile Daspletosaurus from the Late Cretaceous of Alberta, Canada with implications for the identification of immature tyrannosaurids" type="journal article" year="2019">110</bibRefCitation>
]. Although the presence of striated bone may show that an animal has not ceased growing entirely, its presence in relatively large young adults suggests that it cannot be used to identify animals as juveniles.
</paragraph>
<paragraph id="A80736DCFFE3FF8AFE69FBCCFAAEFA9E" blockId="32.[443,1497,367,1370]" pageId="32" pageNumber="33">
In the smallest
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<emphasis id="9ACCEACEFFE3FF8AFD2CFBCCFC88FB54" box="[689,850,1145,1168]" italics="true" pageId="32" pageNumber="33">Nanotyrannus</emphasis>
</taxonomicName>
specimen,
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28471, the surface of the maxillae and nasals is smooth, with little sculpture. However, in the larger
<taxonomicName id="6FB84D5FFFE3FF8AFB4AFB2EFA84FB76" box="[1239,1374,1178,1202]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="lancensis">
<emphasis id="9ACCEACEFFE3FF8AFB4AFB2EFB36FB76" box="[1239,1260,1179,1202]" italics="true" pageId="32" pageNumber="33">N</emphasis>
.
<emphasis id="9ACCEACEFFE3FF8AFB67FB2FFA84FB76" box="[1274,1374,1178,1202]" italics="true" pageId="32" pageNumber="33">lancensis</emphasis>
</taxonomicName>
<typeStatus id="7703887EFFE3FF8AFAFBFB2FFA09FB76" box="[1382,1491,1178,1202]" pageId="32" pageNumber="33">holotype</typeStatus>
,
<collectionCode id="CEA9AE19FFE3FF8AFE21FB09FDCCFB10" box="[444,534,1212,1236]" collectionName="USA, Pennsylvania, Pittsburgh, Carnegie Museum of Natural History" pageId="32" pageNumber="33">CMNH</collectionCode>
7541 (
<figureCitation id="30832A59FFE3FF8AFDFEFB09FD0FFB10" box="[611,725,1212,1236]" captionStart="Figure 22" captionStartId="32.[444,514,1992,2014]" captionTargetBox="[448,1318,1416,1958]" captionTargetId="figure-558@32.[447,1319,1415,1958]" captionTargetPageId="32" captionText="Figure 22. Facial bone rugosity in the Nanotyrannus lancensis holotype CMNH 7541. (A), nasal in left lateral view, (B), right maxilla, (C), right dentary. Scale = 5 cm." figureDoi="http://doi.org/10.5281/zenodo.10535315" httpUri="https://zenodo.org/record/10535315/files/figure.png" pageId="32" pageNumber="33">Figure 22</figureCitation>
), much of the nasals, maxillae, and the anteroventral surface of the dentary are rugose, as are the lateral surface of the lacrimal, the descending process of the postorbital, and the jugal ventral surface. Striated bone occurs inside the antorbital fossa, on the dentarys dorsolateral surface, and the dorsal part of the jugal. The specimen, therefore, shows a mixture of textures, as expected for a subadult or young adult.
</paragraph>
<caption id="FCC76654FFE3FF8AFE21F87DFB97F7C4" ID-DOI="http://doi.org/10.5281/zenodo.10535315" ID-Zenodo-Dep="10535315" httpUri="https://zenodo.org/record/10535315/files/figure.png" pageId="32" pageNumber="33" startId="32.[444,514,1992,2014]" targetBox="[448,1318,1416,1958]" targetPageId="32" targetType="figure">
<paragraph id="A80736DCFFE3FF8AFE21F87DFB97F7C4" blockId="32.[444,1492,1992,2048]" pageId="32" pageNumber="33">
<emphasis id="9ACCEACEFFE3FF8AFE21F87DFDFFF819" bold="true" box="[444,549,1992,2014]" pageId="32" pageNumber="33">Figure 22.</emphasis>
Facial bone rugosity in the
<taxonomicName id="6FB84D5FFFE3FF8AFCD5F87CFBE8F819" box="[840,1074,1992,2014]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="lancensis">
<emphasis id="9ACCEACEFFE3FF8AFCD5F87CFBE8F819" box="[840,1074,1992,2014]" italics="true" pageId="32" pageNumber="33">Nanotyrannus lancensis</emphasis>
</taxonomicName>
holotype CMNH 7541. (
<emphasis id="9ACCEACEFFE3FF8AFAA9F87DFA9CF819" bold="true" box="[1332,1350,1992,2013]" pageId="32" pageNumber="33">A</emphasis>
), nasal in left lateral view, (
<emphasis id="9ACCEACEFFE3FF8AFDD7F85EFD80F7C4" bold="true" box="[586,602,2027,2048]" pageId="32" pageNumber="33">B</emphasis>
), right maxilla, (
<emphasis id="9ACCEACEFFE3FF8AFC94F85FFCC0F7C4" bold="true" box="[777,794,2026,2048]" pageId="32" pageNumber="33">C</emphasis>
), right dentary. Scale = 5 cm.
</paragraph>
</caption>
<paragraph id="A80736DCFFE2FF8BFE69FF44FD55FE36" blockId="33.[443,1497,240,498]" pageId="33" pageNumber="34">
In Jane (
<collectionCode id="CEA9AE19FFE2FF8BFDCAFF45FD7FFECC" box="[599,677,240,264]" collectionName="BMRP" pageId="33" pageNumber="34">BMRP</collectionCode>
<date id="DC06101CFFE2FF8BFD31FF45FCD5FECC" box="[684,783,240,264]" pageId="33" pageNumber="34" value="2002-04-01">2002.4.1</date>
), the maxillae, lacrimals, postorbitals, nasals, and the tip of the dentary are highly rugose and covered with grooves, sculpturing, and gnarled bone (
<figureCitation id="30832A59FFE2FF8BFE5EFE86FDE9FE8F" box="[451,563,307,331]" captionStart="Figure 23" captionStartId="33.[444,514,1634,1656]" captionTargetBox="[456,1292,539,1595]" captionTargetId="figure-225@33.[455,1296,533,1602]" captionTargetPageId="33" captionText="Figure 23. Facial bone rugosity in Jane BMRP 2002.4.1. (A), nasals, (B) right maxilla, (C), left dentary, (D), left lacrimal. The facial bones are extensively sculptured, similar to the condition seen in large Tyrannosaurus rex. Scale = 5 cm." figureDoi="http://doi.org/10.5281/zenodo.10535317" httpUri="https://zenodo.org/record/10535317/files/figure.png" pageId="33" pageNumber="34">Figure 23</figureCitation>
); striated bone is found on the antorbital fossa of the maxilla and lacrimal, the posterior end of the nasal, and the posterior end of the dentary. These bone textures suggest a subadult or young adult. The Zuri specimens show highly rugose sculpturing on the maxilla, nasals, lacrimals, and dentary tip. The maxilla of KU 155809 is also highly rugose. Meanwhile, the nasals of
<collectionCode id="CEA9AE19FFE2FF8BFD77FE0CFCE6FE15" box="[746,828,441,465]" collectionName="USA, California, Los Angeles, Los Angeles County Museum of Natural History" pageId="33" pageNumber="34">LACM</collectionCode>
23845, the smallest definitive
<taxonomicName id="6FB84D5FFFE2FF8BFB3DFE0FFA9FFE15" authorityName=", Sue FMNH PR" authorityYear="2081" box="[1184,1349,442,465]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE2FF8BFB3DFE0FFA9FFE15" box="[1184,1349,442,465]" italics="true" pageId="33" pageNumber="34">Tyrannosaurus</emphasis>
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skull, show weak sculpturing.
</paragraph>
<caption id="FCC76654FFE2FF8BFE21F9D7FCDDF978" ID-DOI="http://doi.org/10.5281/zenodo.10535317" ID-Zenodo-Dep="10535317" httpUri="https://zenodo.org/record/10535317/files/figure.png" pageId="33" pageNumber="34" startId="33.[444,514,1634,1656]" targetBox="[456,1292,539,1595]" targetPageId="33" targetType="figure">
<paragraph id="A80736DCFFE2FF8BFE21F9D7FCDDF978" blockId="33.[442,1495,1634,1724]" pageId="33" pageNumber="34">
<emphasis id="9ACCEACEFFE2FF8BFE21F9D7FDFEF9B3" bold="true" box="[444,548,1634,1656]" pageId="33" pageNumber="34">Figure 23.</emphasis>
Facial bone rugosity in Jane BMRP 2002.4.1. (
<emphasis id="9ACCEACEFFE2FF8BFB9EF9D7FBCFF9B3" bold="true" box="[1027,1045,1634,1655]" pageId="33" pageNumber="34">A</emphasis>
), nasals, (
<emphasis id="9ACCEACEFFE2FF8BFBE3F9D7FB54F9B3" bold="true" box="[1150,1166,1634,1655]" pageId="33" pageNumber="34">B</emphasis>
) right maxilla, (
<emphasis id="9ACCEACEFFE2FF8BFAA8F9D7FA9CF9BC" bold="true" box="[1333,1350,1634,1656]" pageId="33" pageNumber="34">C</emphasis>
), left dentary, (
<emphasis id="9ACCEACEFFE2FF8BFE5EF931FE0DF95D" bold="true" box="[451,471,1668,1689]" pageId="33" pageNumber="34">D</emphasis>
), left lacrimal. The facial bones are extensively sculptured, similar to the condition seen in large
<taxonomicName id="6FB84D5FFFE2FF8BFE27F912FDABF978" box="[442,625,1703,1724]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFE2FF8BFE27F912FDABF978" box="[442,625,1703,1724]" italics="true" pageId="33" pageNumber="34">Tyrannosaurus rex</emphasis>
</taxonomicName>
. Scale = 5 cm.
</paragraph>
</caption>
<paragraph id="A80736DCFFE2FF8BFE69F955FCB5F8B9" blockId="33.[443,1492,1760,1918]" pageId="33" pageNumber="34">
Striated surface textures associated with growth occur in the
<typeStatus id="7703887EFFE2FF8BFB32F955FACFF93C" box="[1199,1301,1760,1784]" pageId="33" pageNumber="34">holotype</typeStatus>
of
<taxonomicName id="6FB84D5FFFE2FF8BFAA4F954FDA0F8DE" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="33" pageNumber="34" phylum="Chordata" rank="species" species="lancensis">
<emphasis id="9ACCEACEFFE2FF8BFAA4F954FDC6F8DD" italics="true" pageId="33" pageNumber="34">Nanotyrannus lancensis</emphasis>
<collectionCode id="CEA9AE19FFE2FF8BFDBEF8B7FDA0F8DE" box="[547,634,1794,1818]" collectionName="USA, Pennsylvania, Pittsburgh, Carnegie Museum of Natural History" pageId="33" pageNumber="34">CMNH</collectionCode>
</taxonomicName>
7541 and in
<collectionCode id="CEA9AE19FFE2FF8BFC96F8B7FC8FF8DD" box="[779,853,1794,1817]" collectionName="BMRP" pageId="33" pageNumber="34">BMRP</collectionCode>
2002.41. However, striated bone is seen in subadults or yong adults of other tyrannosaurs [
<bibRefCitation id="CC294B2DFFE2FF8BFCC3F896FCA3F8FF" author="Voris, J. T. &amp; Therrien, F. &amp; Zelenitsky, D. K. &amp; Brown, C. M." box="[862,889,1827,1851]" journalOrPublisher="Cretac. Res." pageId="33" pageNumber="34" pagination="104388" part="110" refId="ref29913" refString="57. Voris, J. T.; Therrien, F.; Zelenitsky, D. K.; Brown, C. M. A new tyrannosaurine (Theropoda: Tyrannosauridae) from the Campanian Foremost Formation of Alberta, Canada, provides insight into the evolution and biogeography of tyrannosaurids. Cretac. Res. 2020, 110, 104388. [CrossRef]" title="A new tyrannosaurine (Theropoda: Tyrannosauridae) from the Campanian Foremost Formation of Alberta, Canada, provides insight into the evolution and biogeography of tyrannosaurids" type="journal article" year="2020">57</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE2FF8BFCE2F896FC40F8FF" author="Carr, T. D. &amp; Williamson, T. E." box="[895,922,1827,1851]" journalOrPublisher="J. Vertebr. Paleontol." pageId="33" pageNumber="34" pagination="1 - 16" part="30" refId="ref30052" refString="59. Carr, T. D.; Williamson, T. E. Bistahieversor sealeyi, gen. et sp. nov., a new tyrannosauroid from New Mexico and the origin of deep snouts in Tyrannosauroidea. J. Vertebr. Paleontol. 2010, 30, 1 - 16. [CrossRef]" title="Bistahieversor sealeyi, gen. et sp. nov., a new tyrannosauroid from New Mexico and the origin of deep snouts in Tyrannosauroidea." type="journal article" year="2010">59</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE2FF8BFC3CF896FC66F8FF" author="Brusatte, S. L. &amp; Carr, T. D. &amp; Norell, M. A." box="[929,956,1827,1851]" journalOrPublisher="Bull. Am. Mus. Nat. Hist." pageId="33" pageNumber="34" pagination="1 - 197" part="2012" refId="ref30715" refString="70. Brusatte, S. L.; Carr, T. D.; Norell, M. A. The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia. Bull. Am. Mus. Nat. Hist. 2012, 2012, 1 - 197. [CrossRef]" title="The osteology of Alioramus, a gracile and long-snouted tyrannosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Mongolia" type="journal article" year="2012">70</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE2FF8BFC5FF896FC30F8FF" author="Voris, J. T. &amp; Zelenitsky, D. K. &amp; Therrien, F. &amp; Ridgely, R. C. &amp; Currie, P. J. &amp; Witmer, L. M." box="[962,1002,1827,1851]" journalOrPublisher="J. Vertebr. Paleontol." pageId="33" pageNumber="34" pagination="2041651" part="41" refId="ref32804" refString="109. Voris, J. T.; Zelenitsky, D. K.; Therrien, F.; Ridgely, R. C.; Currie, P. J.; Witmer, L. M. Two exceptionally preserved juvenile specimens of Gorgosaurus libratus (Tyrannosauridae, Albertosaurinae) provide new insight into the timing of ontogenetic changes in tyrannosaurids. J. Vertebr. Paleontol. 2021, 41, e 2041651. [CrossRef]" title="Two exceptionally preserved juvenile specimens of Gorgosaurus libratus (Tyrannosauridae, Albertosaurinae) provide new insight into the timing of ontogenetic changes in tyrannosaurids" type="journal article" year="2021">109</bibRefCitation>
]. Overall, bone textures suggest a degree of maturity in these animals, suggesting they are subadults or young adults of a distinct taxon, not juveniles of
<taxonomicName id="6FB84D5FFFE2FF8BFD58F8D2FCB2F8BA" authorityName=", Sue FMNH PR" authorityYear="2081" box="[709,872,1895,1918]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="33" pageNumber="34" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE2FF8BFD58F8D2FCB2F8BA" box="[709,872,1895,1918]" italics="true" pageId="33" pageNumber="34">Tyrannosaurus</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="A80736DCFFE2FF8BFE21F817FCBBF87E" blockId="33.[444,1492,1954,2053]" box="[444,865,1954,1978]" pageId="33" pageNumber="34">3.4.4. External Fundamental System</paragraph>
<paragraph id="A80736DCFFE2FF88FE69F87EFDB5FE8F" blockId="33.[444,1492,1954,2053]" lastBlockId="34.[444,1495,240,767]" lastPageId="34" lastPageNumber="35" pageId="33" pageNumber="34">The external fundamental system, or EFS, is an outermost band of very slow-growing bone with multiple, closely spaced lines of arrested growth (LAGs). It is deposited as growth rates slow and plateau late in life. An EFS can be used as an indicator of the cessation of significant growth and the attainment of maximum body size in a highly mature animal.</paragraph>
<paragraph id="A80736DCFFE1FF88FE69FEE1FD5AFDD0" blockId="34.[444,1495,240,767]" pageId="34" pageNumber="35">
The existence of an EFS would be strong evidence that an animal was old and had effectively stopped growing. The absence of an external fundamental system would suggest that the animal had yet to achieve full adult size. It would mean the animal was not an old adult; however, given that the EFS appears late in life, as the animal attains maximum size [
<bibRefCitation id="CC294B2DFFE1FF88FE65FE6EFDDFFE37" author="Erickson, G. M. &amp; Makovicky, P. J. &amp; Currie, P. J. &amp; Norell, M. A. &amp; Yerby, S. A. &amp; Brochu, C. A." box="[504,517,475,499]" journalOrPublisher="Nature" pageId="34" pageNumber="35" pagination="772 - 775" part="430" refId="ref27355" refString="8. Erickson, G. M.; Makovicky, P. J.; Currie, P. J.; Norell, M. A.; Yerby, S. A.; Brochu, C. A. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 2004, 430, 772 - 775. [CrossRef]" title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">8</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE1FF88FD91FE6EFDFDFE37" author="Cullen, T. M. &amp; Canale, J. I &amp; Apesteguia, S. &amp; Smith, N. D. &amp; Hu, D. &amp; Makovicky, P. J." box="[524,551,475,499]" journalOrPublisher="Proc. R. Soc. B" pageId="34" pageNumber="35" pagination="20202258" part="287" refId="ref29372" refString="49 Cullen, T. M.; Canale, J. I.; Apesteguia, S.; Smith, N. D.; Hu, D.; Makovicky, P. J. Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution. Proc. R. Soc. B 2020, 287, 20202258. [CrossRef]" title="Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution." type="journal article" year="2020">49</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE1FF88FDB0FE6EFD8FFE37" author="Horner, J. R. &amp; de Ricqles, A. &amp; Padian, K." box="[557,597,475,499]" journalOrPublisher="Paleobiology" pageId="34" pageNumber="35" pagination="295 - 304" part="25" refId="ref34029" refString="130. Horner, J. R.; de Ricqles, A.; Padian, K. Variation in dinosaur skeletochronology indicators: Implications for age assessment and physiology. Paleobiology 1999, 25, 295 - 304. [CrossRef]" title="Variation in dinosaur skeletochronology indicators: Implications for age assessment and physiology" type="journal article" year="1999">130</bibRefCitation>
], it would not preclude the possibility that an animal was a young adult just short of full size.
</paragraph>
<paragraph id="A80736DCFFE1FF88FE69FDA8FB23FD3A" blockId="34.[444,1495,240,767]" pageId="34" pageNumber="35">
Three putative
<taxonomicName id="6FB84D5FFFE1FF88FD35FDABFC44FDF1" box="[680,926,542,565]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="34" pageNumber="35" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE1FF88FD35FDABFC9DFDF1" box="[680,839,542,565]" italics="true" pageId="34" pageNumber="35">Nanotyrannus</emphasis>
,
<collectionCode id="CEA9AE19FFE1FF88FCCEFDABFC44FDF1" box="[851,926,542,565]" collectionName="BMRP" pageId="34" pageNumber="35">BMRP</collectionCode>
</taxonomicName>
<date id="DC06101CFFE1FF88FC3BFDABFBDFFDF2" box="[934,1029,542,566]" pageId="34" pageNumber="35" value="2002-04-01">2002.4.1</date>
,
<collectionCode id="CEA9AE19FFE1FF88FB8DFDABFB81FDF1" box="[1040,1115,542,565]" collectionName="BMRP" pageId="34" pageNumber="35">BMRP</collectionCode>
<date id="DC06101CFFE1FF88FBFFFDA8FB19FDF1" box="[1122,1219,541,565]" pageId="34" pageNumber="35" value="2006-04-04">2006.4.4</date>
, and
<collectionCode id="CEA9AE19FFE1FF88FA9CFDABFAEDFDF2" box="[1281,1335,542,566]" collectionName="HRS" pageId="34" pageNumber="35">HRS</collectionCode>
081514, have been sectioned and lack an EFS [
<bibRefCitation id="CC294B2DFFE1FF88FCB1FD8AFC9DFD93" author="Woodward, H. N. &amp; Tremaine, K. &amp; Williams, S. A &amp; Zanno, L. E. &amp; Horner, J. R. &amp; Myhrvold, N." box="[812,839,575,599]" journalOrPublisher="Sci. Adv." pageId="34" pageNumber="35" pagination="eaax 6250" part="6" refId="ref29210" refString="46. Woodward, H. N.; Tremaine, K.; Williams, S. A.; Zanno, L. E.; Horner, J. R.; Myhrvold, N. Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Sci. Adv. 2020, 6, eaax 6250. [CrossRef]" title="Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">46</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE1FF88FCD0FD8AFCB2FD93" author="Griffin, C." bookContentInfo="Cedarville, OH, USA," box="[845,872,575,599]" pageId="34" pageNumber="35" refId="ref29498" refString="51. Griffin, C. Using osteohistology to determine the taxonomic validity of the Late Cretaceous dinosaur Nanotyrannus lancensis Bakker et al, 1988 (Theropoda: Tyrannosauridae). In Proceedings of the The Research and Scholarship Symposium, Cedarville, OH, USA, 16 April 2014." title="Using osteohistology to determine the taxonomic validity of the Late Cretaceous dinosaur Nanotyrannus lancensis Bakker et al, 1988 (Theropoda: Tyrannosauridae)" type="proceedings paper" volumeTitle="Proceedings of the The Research and Scholarship Symposium" year="2014">51</bibRefCitation>
]. This shows these animals are not old adults but does not preclude the possibility that these animals are young adults. In
<taxonomicName id="6FB84D5FFFE1FF88FB48FDD4FACFFDBC" box="[1237,1301,609,632]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="34" pageNumber="35" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFE1FF88FB48FDD4FB3FFDBC" box="[1237,1253,609,632]" italics="true" pageId="34" pageNumber="35">T</emphasis>
.
<emphasis id="9ACCEACEFFE1FF88FB6EFDD4FACFFDBC" box="[1267,1301,609,632]" italics="true" pageId="34" pageNumber="35">rex</emphasis>
</taxonomicName>
, individuals are nearly full size before establishing an external fundamental system. Sue,
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PR 2081, grew to an estimated
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before establishing an EFS [
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,
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], then died at
<quantity id="6F409B39FFE1FF88FAD9FD16FA44FD7F" box="[1348,1438,675,699]" metricMagnitude="3" metricUnit="kg" metricValue="8.222999999999999" pageId="34" pageNumber="35" unit="kg" value="8223.0">8223 kg</quantity>
[
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], adding only around
<quantity id="6F409B39FFE1FF88FD25FD70FCDDFD19" box="[696,775,709,733]" metricMagnitude="2" metricUnit="kg" metricValue="3.0" pageId="34" pageNumber="35" unit="kg" value="300.0">300 kg</quantity>
(i.e., &lt;4% increase) after the appearance of the EFS.
<collectionCode id="CEA9AE19FFE1FF88FA1AFD70FA0EFD18" box="[1415,1492,709,732]" collectionName="BMRP" pageId="34" pageNumber="35">BMRP</collectionCode>
<date id="DC06101CFFE1FF88FE21FD53FDC1FD3A" box="[444,539,742,766]" pageId="34" pageNumber="35" value="2002-04-01">2002.4.1</date>
,
<collectionCode id="CEA9AE19FFE1FF88FDBBFD52FDABFD3A" box="[550,625,743,766]" collectionName="BMRP" pageId="34" pageNumber="35">BMRP</collectionCode>
<date id="DC06101CFFE1FF88FDE5FD53FD0DFD3A" box="[632,727,742,766]" pageId="34" pageNumber="35" value="2006-04-04">2006.4.4</date>
, and
<collectionCode id="CEA9AE19FFE1FF88FC8BFD53FC96FD3A" box="[790,844,742,766]" collectionName="HRS" pageId="34" pageNumber="35">HRS</collectionCode>
081514 may represent young adults.
</paragraph>
<paragraph id="A80736DCFFE1FF88FE21FC97FD7BFCFE" blockId="34.[443,1492,802,1135]" box="[444,673,802,826]" pageId="34" pageNumber="35">3.4.5. Growth Rates</paragraph>
<paragraph id="A80736DCFFE1FF88FE69FCFEFCF4FBAB" blockId="34.[443,1492,802,1135]" pageId="34" pageNumber="35">
Lines of arrested growth (LAGs) record changes in bone circumference and diameter over time (
<figureCitation id="30832A59FFE1FF88FDA0FCD8FD77FC41" box="[573,685,877,901]" captionStart="Figure 24" captionStartId="35.[444,513,940,962]" captionTargetId="figure-210@35.[157,1431,-431,904]" captionTargetPageId="35" captionText="Figure 24. Thin sections of the femur of (A), BMRP 2002.4.1 (“Jane”) and (B), BMRP 2006.4.4 (“Petey”). An external fundamental system (EFS) is absent, but lines of arrested growth or LAGs (blue) become closer toward the outer edge of the bone, showing decelerating growth. Modified from [46]. Note that some LAGs are likely split (multi-LAGs) and do not represent a full year of growth." figureDoi="http://doi.org/10.5281/zenodo.10535319" httpUri="https://zenodo.org/record/10535319/files/figure.png" pageId="34" pageNumber="35">Figure 24</figureCitation>
). Assuming that such lines develop annually, as is commonly done in paleohistology, it becomes possible to reconstruct growth rates by using measured and estimated circumferences either reported in or calculated from published data [
<bibRefCitation id="CC294B2DFFE1FF88FAF0FC05FA52FC0C" author="Woodward, H. N. &amp; Tremaine, K. &amp; Williams, S. A &amp; Zanno, L. E. &amp; Horner, J. R. &amp; Myhrvold, N." box="[1389,1416,944,968]" journalOrPublisher="Sci. Adv." pageId="34" pageNumber="35" pagination="eaax 6250" part="6" refId="ref29210" refString="46. Woodward, H. N.; Tremaine, K.; Williams, S. A.; Zanno, L. E.; Horner, J. R.; Myhrvold, N. Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Sci. Adv. 2020, 6, eaax 6250. [CrossRef]" title="Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">46</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE1FF88FA13FC05FA73FC0C" author="Cullen, T. M. &amp; Canale, J. I &amp; Apesteguia, S. &amp; Smith, N. D. &amp; Hu, D. &amp; Makovicky, P. J." box="[1422,1449,944,968]" journalOrPublisher="Proc. R. Soc. B" pageId="34" pageNumber="35" pagination="20202258" part="287" refId="ref29372" refString="49 Cullen, T. M.; Canale, J. I.; Apesteguia, S.; Smith, N. D.; Hu, D.; Makovicky, P. J. Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution. Proc. R. Soc. B 2020, 287, 20202258. [CrossRef]" title="Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution." type="journal article" year="2020">49</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE1FF88FA2DFC05FA11FC0C" author="Campione, N. E &amp; Evans, D. C. &amp; Brown, C. M. &amp; Carrano, M. T." box="[1456,1483,944,968]" pageId="34" pageNumber="35" pagination="913 - 923" part="5" refId="ref29438" refString="50. Campione, N. E.; Evans, D. C.; Brown, C. M.; Carrano, M. T. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol. Evol. 2014, 5, 913 - 923. [CrossRef]" title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" volumeTitle="Methods Ecol. Evol." year="2014">50</bibRefCitation>
] to estimate mass [
<bibRefCitation id="CC294B2DFFE1FF88FD13FC64FD73FC2D" author="Campione, N. E &amp; Evans, D. C. &amp; Brown, C. M. &amp; Carrano, M. T." box="[654,681,977,1001]" pageId="34" pageNumber="35" pagination="913 - 923" part="5" refId="ref29438" refString="50. Campione, N. E.; Evans, D. C.; Brown, C. M.; Carrano, M. T. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol. Evol. 2014, 5, 913 - 923. [CrossRef]" title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" volumeTitle="Methods Ecol. Evol." year="2014">50</bibRefCitation>
] at various points in the individuals lifespan. By converting femoral circumferences into body mass estimates [
<bibRefCitation id="CC294B2DFFE1FF88FC3DFC46FC61FBCF" author="Campione, N. E &amp; Evans, D. C. &amp; Brown, C. M. &amp; Carrano, M. T." box="[928,955,1011,1035]" pageId="34" pageNumber="35" pagination="913 - 923" part="5" refId="ref29438" refString="50. Campione, N. E.; Evans, D. C.; Brown, C. M.; Carrano, M. T. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol. Evol. 2014, 5, 913 - 923. [CrossRef]" title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" volumeTitle="Methods Ecol. Evol." year="2014">50</bibRefCitation>
], one can estimate changes in mass in terms of kilograms per year (
<tableCitation id="E53A0367FFE1FF88FD30FBA1FCD9FBE8" box="[685,771,1044,1068]" captionStart="Table 5" captionStartId="35.[443,502,1103,1124]" captionTargetPageId="35" captionText="Table 5. LAG data, body mass, and growth rate estimates rounded to the nearest kg for Sue (FMNH PR 2081), Petey (BMRP 2006.4.4), and Jane (BMRP 2002.4.4) based on femoral circumferences, including corrected values for Jane based on a split multi-LAG interpretation [49]. Campione et al. [50] estimate Sues mass at death (i.e., after LAG #23) as 7377 kg after applying a correction for non-circular femoral cross-section. Hutchinson et al. [104] estimate Janes mass at death (i.e., after LAG #9) as 954 kg based on 3D modeling. See Supplemental File for equations used in calculations. Note that LAG #1 and the periosteum do not record a full year of growth. Periosteum is assigned a LAG pseudo-number of +0.5 after the final LAG, indicating that death could have occurred at any point after the last yearly marker was deposited while assigning an age at the midpoint of that final year. Jane required estimation of LAG/periosteum circumferences from LAG spacing and femoral width due to the incomplete cross-section of the femur midshaft. (1) LAG (or periosteum) distance from endosteum is measured at the time of death, (2) femoral width at midshaft (including medullary cavity) is measured at time of death and back estimated using LAG spacing, (3) femoral circumference derived from femoral width at a given year is approximated as a circular cross-section. Data sources: Campione et al. [50]; Cullen et al. [49]; Woodward et al. [46]." httpUri="http://table.plazi.org/id/FCC76654FFE0FF89FE26FBFAFC83F987" pageId="34" pageNumber="35" tableUuid="FCC76654FFE0FF89FE26FBFAFC83F987">Table 5</tableCitation>
). Note that Janes femur is incomplete, so the circumference was approximated as a circle using LAG spacing from the endosteum and using femur width (Supplemental Material).
</paragraph>
<caption id="FCC76654FFE1FF88FE21F8F3FDB9F898" box="[444,611,1862,1884]" isContinuationCaption="true" pageId="34" pageNumber="35" startId="34.[444,514,1862,1884]" targetBox="[157,1431,1178,1825]" targetPageId="34" targetType="figure">
<paragraph id="A80736DCFFE1FF88FE21F8F3FDB9F898" blockId="34.[444,611,1862,1884]" box="[444,611,1862,1884]" pageId="34" pageNumber="35">
<emphasis id="9ACCEACEFFE1FF88FE21F8F3FDFCF89F" bold="true" box="[444,550,1862,1884]" pageId="34" pageNumber="35">Figure 24.</emphasis>
<emphasis id="9ACCEACEFFE1FF88FDB0F8F2FD87F898" box="[557,605,1863,1884]" italics="true" pageId="34" pageNumber="35">Cont</emphasis>
.
</paragraph>
</caption>
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<paragraph id="A80736DCFFE0FF89FE21FC19FA87FBED" blockId="35.[443,1496,940,1065]" pageId="35" pageNumber="36">
<emphasis id="9ACCEACEFFE0FF89FE21FC19FDF9FC05" bold="true" box="[444,547,940,962]" pageId="35" pageNumber="36">Figure 24.</emphasis>
Thin sections of the femur of (
<emphasis id="9ACCEACEFFE0FF89FCC3FC19FCAAFC05" bold="true" box="[862,880,940,961]" pageId="35" pageNumber="36">A</emphasis>
), BMRP 2002.4.1 (“Jane”) and (
<emphasis id="9ACCEACEFFE0FF89FB2DFC19FB1AFC05" bold="true" box="[1200,1216,940,961]" pageId="35" pageNumber="36">B</emphasis>
), BMRP 2006.4.4 (“Petey”). An external fundamental system (EFS) is absent, but lines of arrested growth or LAGs (blue) become closer toward the outer edge of the bone, showing decelerating growth. Modified from [
<bibRefCitation id="CC294B2DFFE0FF89FAEDFC45FA52FBC2" author="Woodward, H. N. &amp; Tremaine, K. &amp; Williams, S. A &amp; Zanno, L. E. &amp; Horner, J. R. &amp; Myhrvold, N." box="[1392,1416,1008,1030]" journalOrPublisher="Sci. Adv." pageId="35" pageNumber="36" pagination="eaax 6250" part="6" refId="ref29210" refString="46. Woodward, H. N.; Tremaine, K.; Williams, S. A.; Zanno, L. E.; Horner, J. R.; Myhrvold, N. Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Sci. Adv. 2020, 6, eaax 6250. [CrossRef]" title="Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">46</bibRefCitation>
]. Note that some LAGs are likely split (multi-LAGs) and do not represent a full year of growth.
</paragraph>
</caption>
<caption id="FCC76654FFE0FF89FE26FBFAFC83F987" ID-Table-UUID="FCC76654FFE0FF89FE26FBFAFC83F987" httpUri="http://table.plazi.org/id/FCC76654FFE0FF89FE26FBFAFC83F987" pageId="35" pageNumber="36" startId="35.[443,502,1103,1124]" targetBox="[112,1469,1638,2013]" targetIsTable="true" targetPageId="35" targetType="table">
<paragraph id="A80736DCFFE0FF89FE26FBFAFC83F987" blockId="35.[443,1495,1103,1604]" pageId="35" pageNumber="36">
<emphasis id="9ACCEACEFFE0FF89FE26FBFAFDD6FBA0" bold="true" box="[443,524,1103,1124]" pageId="35" pageNumber="36">Table 5.</emphasis>
LAG data, body mass, and growth rate estimates rounded to the nearest kg for Sue (FMNH PR 2081), Petey (BMRP 2006.4.4), and Jane (BMRP 2002.4.4) based on femoral circumferences, including corrected values for Jane based on a split multi-LAG interpretation [
<bibRefCitation id="CC294B2DFFE0FF89FBE4FB21FB4BFB6D" author="Cullen, T. M. &amp; Canale, J. I &amp; Apesteguia, S. &amp; Smith, N. D. &amp; Hu, D. &amp; Makovicky, P. J." box="[1145,1169,1172,1193]" journalOrPublisher="Proc. R. Soc. B" pageId="35" pageNumber="36" pagination="20202258" part="287" refId="ref29372" refString="49 Cullen, T. M.; Canale, J. I.; Apesteguia, S.; Smith, N. D.; Hu, D.; Makovicky, P. J. Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution. Proc. R. Soc. B 2020, 287, 20202258. [CrossRef]" title="Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution." type="journal article" year="2020">49</bibRefCitation>
]. Campione et al. [
<bibRefCitation id="CC294B2DFFE0FF89FACAFB21FAB5FB6E" author="Campione, N. E &amp; Evans, D. C. &amp; Brown, C. M. &amp; Carrano, M. T." box="[1367,1391,1172,1194]" pageId="35" pageNumber="36" pagination="913 - 923" part="5" refId="ref29438" refString="50. Campione, N. E.; Evans, D. C.; Brown, C. M.; Carrano, M. T. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol. Evol. 2014, 5, 913 - 923. [CrossRef]" title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" volumeTitle="Methods Ecol. Evol." year="2014">50</bibRefCitation>
] estimate Sues mass at death (i.e., after LAG #23) as 7377 kg after applying a correction for non-circular femoral cross-section. Hutchinson et al. [
<bibRefCitation id="CC294B2DFFE0FF89FC90FB6DFCEBFB29" author="Hutchinson, J. R. &amp; Bates, K. T. &amp; Molnar, J. &amp; Allen, V. &amp; Makovicky, P. J." box="[781,817,1240,1261]" journalOrPublisher="PLoS ONE" pageId="35" pageNumber="36" pagination="e 26037" part="6" refId="ref32548" refString="104. Hutchinson, J. R.; Bates, K. T.; Molnar, J.; Allen, V.; Makovicky, P. J. A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth. PLoS ONE 2011, 6, e 26037. [CrossRef]" title="A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth" type="journal article" year="2011">104</bibRefCitation>
] estimate Janes mass at death (i.e., after LAG #9) as 954 kg based on 3D modeling. See Supplemental File for equations used in calculations. Note that LAG #1 and the periosteum do not record a full year of growth. Periosteum is assigned a LAG pseudo-number of +0.5 after the final LAG, indicating that death could have occurred at any point after the last yearly marker was deposited while assigning an age at the midpoint of that final year. Jane required estimation of LAG/periosteum circumferences from LAG spacing and femoral width due to the incomplete cross-section of the femur midshaft. (1) LAG (or periosteum) distance from endosteum is measured at the time of death, (2) femoral width at midshaft (including medullary cavity) is measured at time of death and back estimated using LAG spacing, (3) femoral circumference derived from femoral width at a given year is approximated as a circular cross-section. Data sources: Campione et al. [
<bibRefCitation id="CC294B2DFFE0FF89FA2CF9B9FA13F9E6" author="Campione, N. E &amp; Evans, D. C. &amp; Brown, C. M. &amp; Carrano, M. T." box="[1457,1481,1548,1570]" pageId="35" pageNumber="36" pagination="913 - 923" part="5" refId="ref29438" refString="50. Campione, N. E.; Evans, D. C.; Brown, C. M.; Carrano, M. T. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol. Evol. 2014, 5, 913 - 923. [CrossRef]" title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" volumeTitle="Methods Ecol. Evol." year="2014">50</bibRefCitation>
]; Cullen et al. [
<bibRefCitation id="CC294B2DFFE0FF89FDD7F99BFDB8F987" author="Cullen, T. M. &amp; Canale, J. I &amp; Apesteguia, S. &amp; Smith, N. D. &amp; Hu, D. &amp; Makovicky, P. J." box="[586,610,1582,1603]" journalOrPublisher="Proc. R. Soc. B" pageId="35" pageNumber="36" pagination="20202258" part="287" refId="ref29372" refString="49 Cullen, T. M.; Canale, J. I.; Apesteguia, S.; Smith, N. D.; Hu, D.; Makovicky, P. J. Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution. Proc. R. Soc. B 2020, 287, 20202258. [CrossRef]" title="Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution." type="journal article" year="2020">49</bibRefCitation>
]; Woodward et al. [
<bibRefCitation id="CC294B2DFFE0FF89FCAEF99BFC91F980" author="Woodward, H. N. &amp; Tremaine, K. &amp; Williams, S. A &amp; Zanno, L. E. &amp; Horner, J. R. &amp; Myhrvold, N." box="[819,843,1582,1604]" journalOrPublisher="Sci. Adv." pageId="35" pageNumber="36" pagination="eaax 6250" part="6" refId="ref29210" refString="46. Woodward, H. N.; Tremaine, K.; Williams, S. A.; Zanno, L. E.; Horner, J. R.; Myhrvold, N. Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Sci. Adv. 2020, 6, eaax 6250. [CrossRef]" title="Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">46</bibRefCitation>
].
</paragraph>
</caption>
<paragraph id="A80736DCFFE0FF89FF10F936FA5BF818" pageId="35" pageNumber="36">
<table id="DAB8C47CFFE00055FFEDF9D3FA67F819" box="[112,1469,1638,2013]" colsContinueIn="36.[112,1469,295,2015]" gridcols="7" gridrows="8" pageId="35" pageNumber="36">
<tr id="1688349EFFE00055FFEDF9D3FA67F972" box="[112,1469,1638,1718]" gridrow="0" pageId="35" pageNumber="36">
<th id="55595DE2FFE00055FFEDF9D3FECCF972" box="[112,278,1638,1718]" gridcol="0" gridrow="0" pageId="35" pageNumber="36">
<emphasis id="9ACCEACEFFE0FF89FF10F936FF23F95D" bold="true" box="[141,249,1667,1689]" pageId="35" pageNumber="36">Specimen</emphasis>
</th>
<th id="55595DE2FFE00055FECFF9D3FE19F972" box="[338,451,1638,1718]" gridcol="1" gridrow="0" pageId="35" pageNumber="36">
<emphasis id="9ACCEACEFFE0FF89FECFF936FE19F95D" bold="true" box="[338,451,1667,1689]" pageId="35" pageNumber="36">Nickname</emphasis>
</th>
<th id="55595DE2FFE00055FD95F9D3FD41F972" box="[520,667,1638,1718]" gridcol="2" gridrow="0" pageId="35" pageNumber="36">
<emphasis id="9ACCEACEFFE0FF89FDACF931FDA8F95D" bold="true" box="[561,626,1668,1689]" pageId="35" pageNumber="36">Taxon</emphasis>
</th>
<th id="55595DE2FFE00055FD48F9D3FCEDF972" box="[725,823,1638,1718]" gridcol="3" gridrow="0" pageId="35" pageNumber="36">
<emphasis id="9ACCEACEFFE0FF89FD48F936FCEDF95D" bold="true" box="[725,823,1667,1689]" pageId="35" pageNumber="36">LAG No.</emphasis>
</th>
<th id="55595DE2FFE00055FCF7F9D3FBD1F972" box="[874,1035,1638,1718]" gridcol="4" gridrow="0" pageId="35" pageNumber="36">
<emphasis id="9ACCEACEFFE0FF89FCF7F9C0FC0DF96C" bold="true" box="[874,1035,1653,1704]" pageId="35" pageNumber="36">Circumference (mm)</emphasis>
</th>
<th id="55595DE2FFE00055FBB3F9D3FB21F972" box="[1070,1275,1638,1718]" gridcol="5" gridrow="0" pageId="35" pageNumber="36">
<emphasis id="9ACCEACEFFE0FF89FBFDF9D3FB21F972" bold="true" pageId="35" pageNumber="36">Mass (kg) (Campione et al., 2014), Equation (7))</emphasis>
</th>
<th id="55595DE2FFE00055FABFF9D3FA67F972" box="[1314,1469,1638,1718]" gridcol="6" gridrow="0" pageId="35" pageNumber="36">
<emphasis id="9ACCEACEFFE0FF89FABFF9D3FA78F972" bold="true" pageId="35" pageNumber="36">Mass Increase from Previous LAG (kg)</emphasis>
</th>
</tr>
<tr id="1688349EFFE00055FFEDF97EFA67F924" box="[112,1469,1739,1760]" gridrow="1" pageId="35" pageNumber="36">
<th id="55595DE2FFE00055FFEDF97EFECCF924" box="[112,278,1739,1760]" gridcol="0" gridrow="1" pageId="35" pageNumber="36">FMNH PR 2081</th>
<td id="55595DE2FFE00055FECFF97EFE19F924" box="[338,451,1739,1760]" gridcol="1" gridrow="1" pageId="35" pageNumber="36">Sue</td>
<td id="55595DE2FFE00055FD95F97EFD41F924" box="[520,667,1739,1760]" gridcol="2" gridrow="1" pageId="35" pageNumber="36">
<taxonomicName id="6FB84D5FFFE0FF89FD95F97EFD41F924" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,1739,1760]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="35" pageNumber="36" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE0FF89FD95F97EFD41F924" box="[520,667,1739,1760]" italics="true" pageId="35" pageNumber="36">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE00055FD48F97EFCEDF924" box="[725,823,1739,1760]" gridcol="3" gridrow="1" pageId="35" pageNumber="36">1</td>
<td id="55595DE2FFE00055FCF7F97EFBD1F924" box="[874,1035,1739,1760]" gridcol="4" gridrow="1" pageId="35" pageNumber="36">338.7</td>
<td id="55595DE2FFE00055FBB3F97EFB21F924" box="[1070,1275,1739,1760]" gridcol="5" gridrow="1" pageId="35" pageNumber="36">1924</td>
<td id="55595DE2FFE00055FABFF97EFA67F924" box="[1314,1469,1739,1760]" gridcol="6" gridrow="1" pageId="35" pageNumber="36">NA</td>
</tr>
<tr id="1688349EFFE00055FFEDF940FA67F8CF" box="[112,1469,1781,1803]" gridrow="2" pageId="35" pageNumber="36">
<th id="55595DE2FFE00055FFEDF940FECCF8CF" box="[112,278,1781,1803]" gridcol="0" gridrow="2" pageId="35" pageNumber="36">FMNH PR 2081</th>
<td id="55595DE2FFE00055FECFF940FE19F8CF" box="[338,451,1781,1803]" gridcol="1" gridrow="2" pageId="35" pageNumber="36">Sue</td>
<td id="55595DE2FFE00055FD95F940FD41F8CF" box="[520,667,1781,1803]" gridcol="2" gridrow="2" pageId="35" pageNumber="36">
<taxonomicName id="6FB84D5FFFE0FF89FD95F940FD41F8CE" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,1781,1802]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="35" pageNumber="36" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE0FF89FD95F940FD41F8CE" box="[520,667,1781,1802]" italics="true" pageId="35" pageNumber="36">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE00055FD48F940FCEDF8CF" box="[725,823,1781,1803]" gridcol="3" gridrow="2" pageId="35" pageNumber="36">2</td>
<td id="55595DE2FFE00055FCF7F940FBD1F8CF" box="[874,1035,1781,1803]" gridcol="4" gridrow="2" pageId="35" pageNumber="36">387</td>
<td id="55595DE2FFE00055FBB3F940FB21F8CF" box="[1070,1275,1781,1803]" gridcol="5" gridrow="2" pageId="35" pageNumber="36">2777</td>
<td id="55595DE2FFE00055FABFF940FA67F8CF" box="[1314,1469,1781,1803]" gridcol="6" gridrow="2" pageId="35" pageNumber="36">853</td>
</tr>
<tr id="1688349EFFE00055FFEDF8AAFA67F8F1" box="[112,1469,1823,1845]" gridrow="3" pageId="35" pageNumber="36">
<th id="55595DE2FFE00055FFEDF8AAFECCF8F1" box="[112,278,1823,1845]" gridcol="0" gridrow="3" pageId="35" pageNumber="36">FMNH PR 2081</th>
<td id="55595DE2FFE00055FECFF8AAFE19F8F1" box="[338,451,1823,1845]" gridcol="1" gridrow="3" pageId="35" pageNumber="36">Sue</td>
<td id="55595DE2FFE00055FD95F8AAFD41F8F1" box="[520,667,1823,1845]" gridcol="2" gridrow="3" pageId="35" pageNumber="36">
<taxonomicName id="6FB84D5FFFE0FF89FD95F8AAFD41F8F0" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,1823,1844]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="35" pageNumber="36" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE0FF89FD95F8AAFD41F8F0" box="[520,667,1823,1844]" italics="true" pageId="35" pageNumber="36">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE00055FD48F8AAFCEDF8F1" box="[725,823,1823,1845]" gridcol="3" gridrow="3" pageId="35" pageNumber="36">3</td>
<td id="55595DE2FFE00055FCF7F8AAFBD1F8F1" box="[874,1035,1823,1845]" gridcol="4" gridrow="3" pageId="35" pageNumber="36">415.7</td>
<td id="55595DE2FFE00055FBB3F8AAFB21F8F1" box="[1070,1275,1823,1845]" gridcol="5" gridrow="3" pageId="35" pageNumber="36">3382</td>
<td id="55595DE2FFE00055FABFF8AAFA67F8F1" box="[1314,1469,1823,1845]" gridcol="6" gridrow="3" pageId="35" pageNumber="36">605</td>
</tr>
<tr id="1688349EFFE00055FFEDF8FCFA67F89B" box="[112,1469,1865,1887]" gridrow="4" pageId="35" pageNumber="36">
<th id="55595DE2FFE00055FFEDF8FCFECCF89B" box="[112,278,1865,1887]" gridcol="0" gridrow="4" pageId="35" pageNumber="36">FMNH PR 2081</th>
<td id="55595DE2FFE00055FECFF8FCFE19F89B" box="[338,451,1865,1887]" gridcol="1" gridrow="4" pageId="35" pageNumber="36">Sue</td>
<td id="55595DE2FFE00055FD95F8FCFD41F89B" box="[520,667,1865,1887]" gridcol="2" gridrow="4" pageId="35" pageNumber="36">
<taxonomicName id="6FB84D5FFFE0FF89FD95F8FCFD41F89A" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,1865,1886]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="35" pageNumber="36" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE0FF89FD95F8FCFD41F89A" box="[520,667,1865,1886]" italics="true" pageId="35" pageNumber="36">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE00055FD48F8FCFCEDF89B" box="[725,823,1865,1887]" gridcol="3" gridrow="4" pageId="35" pageNumber="36">4</td>
<td id="55595DE2FFE00055FCF7F8FCFBD1F89B" box="[874,1035,1865,1887]" gridcol="4" gridrow="4" pageId="35" pageNumber="36">440.3</td>
<td id="55595DE2FFE00055FBB3F8FCFB21F89B" box="[1070,1275,1865,1887]" gridcol="5" gridrow="4" pageId="35" pageNumber="36">3962</td>
<td id="55595DE2FFE00055FABFF8FCFA67F89B" box="[1314,1469,1865,1887]" gridcol="6" gridrow="4" pageId="35" pageNumber="36">580</td>
</tr>
<tr id="1688349EFFE00055FFEDF8C6FA67F84D" box="[112,1469,1907,1929]" gridrow="5" pageId="35" pageNumber="36">
<th id="55595DE2FFE00055FFEDF8C6FECCF84D" box="[112,278,1907,1929]" gridcol="0" gridrow="5" pageId="35" pageNumber="36">FMNH PR 2081</th>
<td id="55595DE2FFE00055FECFF8C6FE19F84D" box="[338,451,1907,1929]" gridcol="1" gridrow="5" pageId="35" pageNumber="36">Sue</td>
<td id="55595DE2FFE00055FD95F8C6FD41F84D" box="[520,667,1907,1929]" gridcol="2" gridrow="5" pageId="35" pageNumber="36">
<taxonomicName id="6FB84D5FFFE0FF89FD95F8C6FD41F84C" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,1907,1928]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="35" pageNumber="36" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE0FF89FD95F8C6FD41F84C" box="[520,667,1907,1928]" italics="true" pageId="35" pageNumber="36">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE00055FD48F8C6FCEDF84D" box="[725,823,1907,1929]" gridcol="3" gridrow="5" pageId="35" pageNumber="36">5</td>
<td id="55595DE2FFE00055FCF7F8C6FBD1F84D" box="[874,1035,1907,1929]" gridcol="4" gridrow="5" pageId="35" pageNumber="36">459.7</td>
<td id="55595DE2FFE00055FBB3F8C6FB21F84D" box="[1070,1275,1907,1929]" gridcol="5" gridrow="5" pageId="35" pageNumber="36">4461</td>
<td id="55595DE2FFE00055FABFF8C6FA67F84D" box="[1314,1469,1907,1929]" gridcol="6" gridrow="5" pageId="35" pageNumber="36">500</td>
</tr>
<tr id="1688349EFFE00055FFEDF828FA67F877" box="[112,1469,1949,1971]" gridrow="6" pageId="35" pageNumber="36">
<th id="55595DE2FFE00055FFEDF828FECCF877" box="[112,278,1949,1971]" gridcol="0" gridrow="6" pageId="35" pageNumber="36">FMNH PR 2081</th>
<td id="55595DE2FFE00055FECFF828FE19F877" box="[338,451,1949,1971]" gridcol="1" gridrow="6" pageId="35" pageNumber="36">Sue</td>
<td id="55595DE2FFE00055FD95F828FD41F877" box="[520,667,1949,1971]" gridcol="2" gridrow="6" pageId="35" pageNumber="36">
<taxonomicName id="6FB84D5FFFE0FF89FD95F828FD41F876" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,1949,1970]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="35" pageNumber="36" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE0FF89FD95F828FD41F876" box="[520,667,1949,1970]" italics="true" pageId="35" pageNumber="36">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE00055FD48F828FCEDF877" box="[725,823,1949,1971]" gridcol="3" gridrow="6" pageId="35" pageNumber="36">6</td>
<td id="55595DE2FFE00055FCF7F828FBD1F877" box="[874,1035,1949,1971]" gridcol="4" gridrow="6" pageId="35" pageNumber="36">485.5</td>
<td id="55595DE2FFE00055FBB3F828FB21F877" box="[1070,1275,1949,1971]" gridcol="5" gridrow="6" pageId="35" pageNumber="36">5185</td>
<td id="55595DE2FFE00055FABFF828FA67F877" box="[1314,1469,1949,1971]" gridcol="6" gridrow="6" pageId="35" pageNumber="36">724</td>
</tr>
<tr id="1688349EFFE00055FFEDF872FA67F819" box="[112,1469,1991,2013]" gridrow="7" pageId="35" pageNumber="36">
<th id="55595DE2FFE00055FFEDF872FECCF819" box="[112,278,1991,2013]" gridcol="0" gridrow="7" pageId="35" pageNumber="36">FMNH PR 2081</th>
<td id="55595DE2FFE00055FECFF872FE19F819" box="[338,451,1991,2013]" gridcol="1" gridrow="7" pageId="35" pageNumber="36">Sue</td>
<td id="55595DE2FFE00055FD95F872FD41F819" box="[520,667,1991,2013]" gridcol="2" gridrow="7" pageId="35" pageNumber="36">
<taxonomicName id="6FB84D5FFFE0FF89FD95F87DFD41F819" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,1992,2013]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="35" pageNumber="36" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE0FF89FD95F87DFD41F819" box="[520,667,1992,2013]" italics="true" pageId="35" pageNumber="36">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE00055FD48F872FCEDF819" box="[725,823,1991,2013]" gridcol="3" gridrow="7" pageId="35" pageNumber="36">7</td>
<td id="55595DE2FFE00055FCF7F872FBD1F819" box="[874,1035,1991,2013]" gridcol="4" gridrow="7" pageId="35" pageNumber="36">511.1</td>
<td id="55595DE2FFE00055FBB3F872FB21F819" box="[1070,1275,1991,2013]" gridcol="5" gridrow="7" pageId="35" pageNumber="36">5973</td>
<td id="55595DE2FFE00055FABFF872FA67F819" box="[1314,1469,1991,2013]" gridcol="6" gridrow="7" pageId="35" pageNumber="36">788</td>
</tr>
</table>
</paragraph>
<caption id="FCC76654FFE7FF8EFE26FF5AFD97FEC1" box="[443,589,239,261]" isContinuationCaption="true" pageId="36" pageNumber="37" startId="36.[443,503,239,260]" targetType="table">
<paragraph id="A80736DCFFE7FF8EFE26FF5AFD97FEC1" blockId="36.[443,589,239,261]" box="[443,589,239,261]" pageId="36" pageNumber="37">
<emphasis id="9ACCEACEFFE7FF8EFE26FF5AFDD5FEC0" bold="true" box="[443,527,239,260]" pageId="36" pageNumber="37">Table 5.</emphasis>
Cont.
</paragraph>
</caption>
<paragraph id="A80736DCFFE7FF8EFF10FEF1FA5BF81A" pageId="36" pageNumber="37">
<table id="DAB8C47CFFE70055FFEDFE92FA67F81B" box="[112,1469,295,2015]" colsContinueFrom="35.[112,1469,1638,2013]" colsContinueIn="37.[116,1469,295,502]" gridcols="7" gridrows="40" pageId="36" pageNumber="37">
<tr id="1688349EFFE70055FFEDFE92FA67FEBC" box="[112,1469,295,376]" gridrow="0" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFE92FECCFEBC" box="[112,278,295,376]" gridcol="0" gridrow="0" pageId="36" pageNumber="37">
<emphasis id="9ACCEACEFFE7FF8EFF10FEF1FF23FE9E" bold="true" box="[141,249,324,346]" pageId="36" pageNumber="37">Specimen</emphasis>
</th>
<th id="55595DE2FFE70055FEA5FE92FE07FEBC" box="[312,477,295,376]" gridcol="1" gridrow="0" pageId="36" pageNumber="37">
<emphasis id="9ACCEACEFFE7FF8EFECFFEF1FE19FE9E" bold="true" box="[338,451,324,346]" pageId="36" pageNumber="37">Nickname</emphasis>
</th>
<th id="55595DE2FFE70055FD95FE92FD41FEBC" box="[520,667,295,376]" gridcol="2" gridrow="0" pageId="36" pageNumber="37">
<emphasis id="9ACCEACEFFE7FF8EFDACFEF0FDA8FE9E" bold="true" box="[561,626,325,346]" pageId="36" pageNumber="37">Taxon</emphasis>
</th>
<th id="55595DE2FFE70055FD57FE92FC98FEBC" box="[714,834,295,376]" gridcol="3" gridrow="0" pageId="36" pageNumber="37">
<emphasis id="9ACCEACEFFE7FF8EFD48FEF1FCEDFE9E" bold="true" box="[725,823,324,346]" pageId="36" pageNumber="37">LAG No.</emphasis>
</th>
<th id="55595DE2FFE70055FCF7FE92FBD1FEBC" box="[874,1035,295,376]" gridcol="4" gridrow="0" pageId="36" pageNumber="37">
<emphasis id="9ACCEACEFFE7FF8EFCF7FE83FC0DFEAD" bold="true" box="[874,1035,310,361]" pageId="36" pageNumber="37">Circumference (mm)</emphasis>
</th>
<th id="55595DE2FFE70055FBB3FE92FB21FEBC" box="[1070,1275,295,376]" gridcol="5" gridrow="0" pageId="36" pageNumber="37">
<emphasis id="9ACCEACEFFE7FF8EFBFDFE9DFB21FEBC" bold="true" pageId="36" pageNumber="37">Mass (kg) (Campione et al., 2014), Equation (7))</emphasis>
</th>
<th id="55595DE2FFE70055FABFFE92FA67FEBC" box="[1314,1469,295,376]" gridcol="6" gridrow="0" pageId="36" pageNumber="37">
<emphasis id="9ACCEACEFFE7FF8EFABFFE9DFA78FEBC" bold="true" pageId="36" pageNumber="37">Mass Increase from Previous LAG (kg)</emphasis>
</th>
</tr>
<tr id="1688349EFFE70055FFEDFE39FA67FE66" box="[112,1469,396,418]" gridrow="1" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFE39FECCFE66" box="[112,278,396,418]" gridcol="0" gridrow="1" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FE39FE07FE66" box="[312,477,396,418]" gridcol="1" gridrow="1" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FE39FD41FE66" box="[520,667,396,418]" gridcol="2" gridrow="1" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FE38FD41FE66" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,397,418]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FE38FD41FE66" box="[520,667,397,418]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FE39FC98FE66" box="[714,834,396,418]" gridcol="3" gridrow="1" pageId="36" pageNumber="37">8</td>
<td id="55595DE2FFE70055FCF7FE39FBD1FE66" box="[874,1035,396,418]" gridcol="4" gridrow="1" pageId="36" pageNumber="37">527.1</td>
<td id="55595DE2FFE70055FBB3FE39FB21FE66" box="[1070,1275,396,418]" gridcol="5" gridrow="1" pageId="36" pageNumber="37">6503</td>
<td id="55595DE2FFE70055FABFFE39FA67FE66" box="[1314,1469,396,418]" gridcol="6" gridrow="1" pageId="36" pageNumber="37">529</td>
</tr>
<tr id="1688349EFFE70055FFEDFE03FA67FE08" box="[112,1469,438,460]" gridrow="2" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFE03FECCFE08" box="[112,278,438,460]" gridcol="0" gridrow="2" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FE03FE07FE08" box="[312,477,438,460]" gridcol="1" gridrow="2" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FE03FD41FE08" box="[520,667,438,460]" gridcol="2" gridrow="2" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FE02FD41FE08" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,439,460]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FE02FD41FE08" box="[520,667,439,460]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FE03FC98FE08" box="[714,834,438,460]" gridcol="3" gridrow="2" pageId="36" pageNumber="37">9</td>
<td id="55595DE2FFE70055FCF7FE03FBD1FE08" box="[874,1035,438,460]" gridcol="4" gridrow="2" pageId="36" pageNumber="37">542.8</td>
<td id="55595DE2FFE70055FBB3FE03FB21FE08" box="[1070,1275,438,460]" gridcol="5" gridrow="2" pageId="36" pageNumber="37">7050</td>
<td id="55595DE2FFE70055FABFFE03FA67FE08" box="[1314,1469,438,460]" gridcol="6" gridrow="2" pageId="36" pageNumber="37">547</td>
</tr>
<tr id="1688349EFFE70055FFEDFE55FA67FE32" box="[112,1469,480,502]" gridrow="3" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFE55FECCFE32" box="[112,278,480,502]" gridcol="0" gridrow="3" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FE55FE07FE32" box="[312,477,480,502]" gridcol="1" gridrow="3" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FE55FD41FE32" box="[520,667,480,502]" gridcol="2" gridrow="3" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FE54FD41FE32" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,481,502]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FE54FD41FE32" box="[520,667,481,502]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FE55FC98FE32" box="[714,834,480,502]" gridcol="3" gridrow="3" pageId="36" pageNumber="37">10</td>
<td id="55595DE2FFE70055FCF7FE55FBD1FE32" box="[874,1035,480,502]" gridcol="4" gridrow="3" pageId="36" pageNumber="37">551.6</td>
<td id="55595DE2FFE70055FBB3FE55FB21FE32" box="[1070,1275,480,502]" gridcol="5" gridrow="3" pageId="36" pageNumber="37">7369</td>
<td id="55595DE2FFE70055FABFFE55FA67FE32" box="[1314,1469,480,502]" gridcol="6" gridrow="3" pageId="36" pageNumber="37">319</td>
</tr>
<tr id="1688349EFFE70055FFEDFDBFFA67FDE4" box="[112,1469,522,544]" gridrow="4" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFDBFFECCFDE4" box="[112,278,522,544]" gridcol="0" gridrow="4" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FDBFFE07FDE4" box="[312,477,522,544]" gridcol="1" gridrow="4" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FDBFFD41FDE4" box="[520,667,522,544]" gridcol="2" gridrow="4" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FDBEFD41FDE4" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,523,544]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FDBEFD41FDE4" box="[520,667,523,544]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FDBFFC98FDE4" box="[714,834,522,544]" gridcol="3" gridrow="4" pageId="36" pageNumber="37">11</td>
<td id="55595DE2FFE70055FCF7FDBFFBD1FDE4" box="[874,1035,522,544]" gridcol="4" gridrow="4" pageId="36" pageNumber="37">559.3</td>
<td id="55595DE2FFE70055FBB3FDBFFB21FDE4" box="[1070,1275,522,544]" gridcol="5" gridrow="4" pageId="36" pageNumber="37">7656</td>
<td id="55595DE2FFE70055FABFFDBFFA67FDE4" box="[1314,1469,522,544]" gridcol="6" gridrow="4" pageId="36" pageNumber="37">287</td>
</tr>
<tr id="1688349EFFE70055FFEDFD81FA67FD8E" box="[112,1469,564,586]" gridrow="5" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFD81FECCFD8E" box="[112,278,564,586]" gridcol="0" gridrow="5" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FD81FE07FD8E" box="[312,477,564,586]" gridcol="1" gridrow="5" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FD81FD41FD8E" box="[520,667,564,586]" gridcol="2" gridrow="5" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FD80FD41FD8E" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,565,586]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FD80FD41FD8E" box="[520,667,565,586]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FD81FC98FD8E" box="[714,834,564,586]" gridcol="3" gridrow="5" pageId="36" pageNumber="37">12</td>
<td id="55595DE2FFE70055FCF7FD81FBD1FD8E" box="[874,1035,564,586]" gridcol="4" gridrow="5" pageId="36" pageNumber="37">564.9</td>
<td id="55595DE2FFE70055FBB3FD81FB21FD8E" box="[1070,1275,564,586]" gridcol="5" gridrow="5" pageId="36" pageNumber="37">7869</td>
<td id="55595DE2FFE70055FABFFD81FA67FD8E" box="[1314,1469,564,586]" gridcol="6" gridrow="5" pageId="36" pageNumber="37">213</td>
</tr>
<tr id="1688349EFFE70055FFEDFDEBFA67FDB0" box="[112,1469,606,628]" gridrow="6" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFDEBFECCFDB0" box="[112,278,606,628]" gridcol="0" gridrow="6" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FDEBFE07FDB0" box="[312,477,606,628]" gridcol="1" gridrow="6" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FDEBFD41FDB0" box="[520,667,606,628]" gridcol="2" gridrow="6" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FDEAFD41FDB0" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,607,628]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FDEAFD41FDB0" box="[520,667,607,628]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FDEBFC98FDB0" box="[714,834,606,628]" gridcol="3" gridrow="6" pageId="36" pageNumber="37">13</td>
<td id="55595DE2FFE70055FCF7FDEBFBD1FDB0" box="[874,1035,606,628]" gridcol="4" gridrow="6" pageId="36" pageNumber="37">565.7</td>
<td id="55595DE2FFE70055FBB3FDEBFB21FDB0" box="[1070,1275,606,628]" gridcol="5" gridrow="6" pageId="36" pageNumber="37">7900</td>
<td id="55595DE2FFE70055FABFFDEBFA67FDB0" box="[1314,1469,606,628]" gridcol="6" gridrow="6" pageId="36" pageNumber="37">31</td>
</tr>
<tr id="1688349EFFE70055FFEDFD3DFA67FD5A" box="[112,1469,648,670]" gridrow="7" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFD3DFECCFD5A" box="[112,278,648,670]" gridcol="0" gridrow="7" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FD3DFE07FD5A" box="[312,477,648,670]" gridcol="1" gridrow="7" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FD3DFD41FD5A" box="[520,667,648,670]" gridcol="2" gridrow="7" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FD3CFD41FD5A" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,649,670]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FD3CFD41FD5A" box="[520,667,649,670]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FD3DFC98FD5A" box="[714,834,648,670]" gridcol="3" gridrow="7" pageId="36" pageNumber="37">14</td>
<td id="55595DE2FFE70055FCF7FD3DFBD1FD5A" box="[874,1035,648,670]" gridcol="4" gridrow="7" pageId="36" pageNumber="37">566.5</td>
<td id="55595DE2FFE70055FBB3FD3DFB21FD5A" box="[1070,1275,648,670]" gridcol="5" gridrow="7" pageId="36" pageNumber="37">7931</td>
<td id="55595DE2FFE70055FABFFD3DFA67FD5A" box="[1314,1469,648,670]" gridcol="6" gridrow="7" pageId="36" pageNumber="37">31</td>
</tr>
<tr id="1688349EFFE70055FFEDFD07FA67FD0C" box="[112,1469,690,712]" gridrow="8" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFD07FECCFD0C" box="[112,278,690,712]" gridcol="0" gridrow="8" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FD07FE07FD0C" box="[312,477,690,712]" gridcol="1" gridrow="8" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FD07FD41FD0C" box="[520,667,690,712]" gridcol="2" gridrow="8" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FD06FD41FD0C" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,691,712]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FD06FD41FD0C" box="[520,667,691,712]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FD07FC98FD0C" box="[714,834,690,712]" gridcol="3" gridrow="8" pageId="36" pageNumber="37">15</td>
<td id="55595DE2FFE70055FCF7FD07FBD1FD0C" box="[874,1035,690,712]" gridcol="4" gridrow="8" pageId="36" pageNumber="37">567.4</td>
<td id="55595DE2FFE70055FBB3FD07FB21FD0C" box="[1070,1275,690,712]" gridcol="5" gridrow="8" pageId="36" pageNumber="37">7965</td>
<td id="55595DE2FFE70055FABFFD07FA67FD0C" box="[1314,1469,690,712]" gridcol="6" gridrow="8" pageId="36" pageNumber="37">35</td>
</tr>
<tr id="1688349EFFE70055FFEDFD69FA67FD36" box="[112,1469,732,754]" gridrow="9" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFD69FECCFD36" box="[112,278,732,754]" gridcol="0" gridrow="9" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FD69FE07FD36" box="[312,477,732,754]" gridcol="1" gridrow="9" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FD69FD41FD36" box="[520,667,732,754]" gridcol="2" gridrow="9" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FD68FD41FD36" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,733,754]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FD68FD41FD36" box="[520,667,733,754]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FD69FC98FD36" box="[714,834,732,754]" gridcol="3" gridrow="9" pageId="36" pageNumber="37">16</td>
<td id="55595DE2FFE70055FCF7FD69FBD1FD36" box="[874,1035,732,754]" gridcol="4" gridrow="9" pageId="36" pageNumber="37">568.2</td>
<td id="55595DE2FFE70055FBB3FD69FB21FD36" box="[1070,1275,732,754]" gridcol="5" gridrow="9" pageId="36" pageNumber="37">7996</td>
<td id="55595DE2FFE70055FABFFD69FA67FD36" box="[1314,1469,732,754]" gridcol="6" gridrow="9" pageId="36" pageNumber="37">31</td>
</tr>
<tr id="1688349EFFE70055FFEDFCB3FA67FCD8" box="[112,1469,774,796]" gridrow="10" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFCB3FECCFCD8" box="[112,278,774,796]" gridcol="0" gridrow="10" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FCB3FE07FCD8" box="[312,477,774,796]" gridcol="1" gridrow="10" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FCB3FD41FCD8" box="[520,667,774,796]" gridcol="2" gridrow="10" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FCB2FD41FCD8" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,775,796]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FCB2FD41FCD8" box="[520,667,775,796]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FCB3FC98FCD8" box="[714,834,774,796]" gridcol="3" gridrow="10" pageId="36" pageNumber="37">17</td>
<td id="55595DE2FFE70055FCF7FCB3FBD1FCD8" box="[874,1035,774,796]" gridcol="4" gridrow="10" pageId="36" pageNumber="37">569</td>
<td id="55595DE2FFE70055FBB3FCB3FB21FCD8" box="[1070,1275,774,796]" gridcol="5" gridrow="10" pageId="36" pageNumber="37">8027</td>
<td id="55595DE2FFE70055FABFFCB3FA67FCD8" box="[1314,1469,774,796]" gridcol="6" gridrow="10" pageId="36" pageNumber="37">31</td>
</tr>
<tr id="1688349EFFE70055FFEDFC85FA67FC82" box="[112,1469,816,838]" gridrow="11" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFC85FECCFC82" box="[112,278,816,838]" gridcol="0" gridrow="11" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FC85FE07FC82" box="[312,477,816,838]" gridcol="1" gridrow="11" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FC85FD41FC82" box="[520,667,816,838]" gridcol="2" gridrow="11" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FC84FD41FC82" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,817,838]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FC84FD41FC82" box="[520,667,817,838]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FC85FC98FC82" box="[714,834,816,838]" gridcol="3" gridrow="11" pageId="36" pageNumber="37">18</td>
<td id="55595DE2FFE70055FCF7FC85FBD1FC82" box="[874,1035,816,838]" gridcol="4" gridrow="11" pageId="36" pageNumber="37">569.8</td>
<td id="55595DE2FFE70055FBB3FC85FB21FC82" box="[1070,1275,816,838]" gridcol="5" gridrow="11" pageId="36" pageNumber="37">8059</td>
<td id="55595DE2FFE70055FABFFC85FA67FC82" box="[1314,1469,816,838]" gridcol="6" gridrow="11" pageId="36" pageNumber="37">31</td>
</tr>
<tr id="1688349EFFE70055FFEDFCEFFA67FCB4" box="[112,1469,858,880]" gridrow="12" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFCEFFECCFCB4" box="[112,278,858,880]" gridcol="0" gridrow="12" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FCEFFE07FCB4" box="[312,477,858,880]" gridcol="1" gridrow="12" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FCEFFD41FCB4" box="[520,667,858,880]" gridcol="2" gridrow="12" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FCEEFD41FCB4" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,859,880]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FCEEFD41FCB4" box="[520,667,859,880]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FCEFFC98FCB4" box="[714,834,858,880]" gridcol="3" gridrow="12" pageId="36" pageNumber="37">19</td>
<td id="55595DE2FFE70055FCF7FCEFFBD1FCB4" box="[874,1035,858,880]" gridcol="4" gridrow="12" pageId="36" pageNumber="37">570.7</td>
<td id="55595DE2FFE70055FBB3FCEFFB21FCB4" box="[1070,1275,858,880]" gridcol="5" gridrow="12" pageId="36" pageNumber="37">8094</td>
<td id="55595DE2FFE70055FABFFCEFFA67FCB4" box="[1314,1469,858,880]" gridcol="6" gridrow="12" pageId="36" pageNumber="37">35</td>
</tr>
<tr id="1688349EFFE70055FFEDFC31FA67FC5E" box="[112,1469,900,922]" gridrow="13" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFC31FECCFC5E" box="[112,278,900,922]" gridcol="0" gridrow="13" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FC31FE07FC5E" box="[312,477,900,922]" gridcol="1" gridrow="13" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FC31FD41FC5E" box="[520,667,900,922]" gridcol="2" gridrow="13" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FC30FD41FC5E" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,901,922]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FC30FD41FC5E" box="[520,667,901,922]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FC31FC98FC5E" box="[714,834,900,922]" gridcol="3" gridrow="13" pageId="36" pageNumber="37">20</td>
<td id="55595DE2FFE70055FCF7FC31FBD1FC5E" box="[874,1035,900,922]" gridcol="4" gridrow="13" pageId="36" pageNumber="37">571.5</td>
<td id="55595DE2FFE70055FBB3FC31FB21FC5E" box="[1070,1275,900,922]" gridcol="5" gridrow="13" pageId="36" pageNumber="37">8125</td>
<td id="55595DE2FFE70055FABFFC31FA67FC5E" box="[1314,1469,900,922]" gridcol="6" gridrow="13" pageId="36" pageNumber="37">31</td>
</tr>
<tr id="1688349EFFE70055FFEDFC1BFA67FC00" box="[112,1469,942,964]" gridrow="14" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFC1BFECCFC00" box="[112,278,942,964]" gridcol="0" gridrow="14" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FC1BFE07FC00" box="[312,477,942,964]" gridcol="1" gridrow="14" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FC1BFD41FC00" box="[520,667,942,964]" gridcol="2" gridrow="14" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FC1AFD41FC00" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,943,964]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FC1AFD41FC00" box="[520,667,943,964]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FC1BFC98FC00" box="[714,834,942,964]" gridcol="3" gridrow="14" pageId="36" pageNumber="37">21</td>
<td id="55595DE2FFE70055FCF7FC1BFBD1FC00" box="[874,1035,942,964]" gridcol="4" gridrow="14" pageId="36" pageNumber="37">572.3</td>
<td id="55595DE2FFE70055FBB3FC1BFB21FC00" box="[1070,1275,942,964]" gridcol="5" gridrow="14" pageId="36" pageNumber="37">8156</td>
<td id="55595DE2FFE70055FABFFC1BFA67FC00" box="[1314,1469,942,964]" gridcol="6" gridrow="14" pageId="36" pageNumber="37">31</td>
</tr>
<tr id="1688349EFFE70055FFEDFC6DFA67FC2A" box="[112,1469,984,1006]" gridrow="15" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFC6DFECCFC2A" box="[112,278,984,1006]" gridcol="0" gridrow="15" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FC6DFE07FC2A" box="[312,477,984,1006]" gridcol="1" gridrow="15" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FC6DFD41FC2A" box="[520,667,984,1006]" gridcol="2" gridrow="15" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FC6CFD41FC2A" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,985,1006]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FC6CFD41FC2A" box="[520,667,985,1006]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FC6DFC98FC2A" box="[714,834,984,1006]" gridcol="3" gridrow="15" pageId="36" pageNumber="37">22</td>
<td id="55595DE2FFE70055FCF7FC6DFBD1FC2A" box="[874,1035,984,1006]" gridcol="4" gridrow="15" pageId="36" pageNumber="37">573.1</td>
<td id="55595DE2FFE70055FBB3FC6DFB21FC2A" box="[1070,1275,984,1006]" gridcol="5" gridrow="15" pageId="36" pageNumber="37">8188</td>
<td id="55595DE2FFE70055FABFFC6DFA67FC2A" box="[1314,1469,984,1006]" gridcol="6" gridrow="15" pageId="36" pageNumber="37">31</td>
</tr>
<tr id="1688349EFFE70055FFEDFBB7FA67FBDC" box="[112,1469,1026,1048]" gridrow="16" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFBB7FECCFBDC" box="[112,278,1026,1048]" gridcol="0" gridrow="16" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FBB7FE07FBDC" box="[312,477,1026,1048]" gridcol="1" gridrow="16" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FBB7FD41FBDC" box="[520,667,1026,1048]" gridcol="2" gridrow="16" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FBB6FD41FBDC" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,1027,1048]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FBB6FD41FBDC" box="[520,667,1027,1048]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FBB7FC98FBDC" box="[714,834,1026,1048]" gridcol="3" gridrow="16" pageId="36" pageNumber="37">23</td>
<td id="55595DE2FFE70055FCF7FBB7FBD1FBDC" box="[874,1035,1026,1048]" gridcol="4" gridrow="16" pageId="36" pageNumber="37">574</td>
<td id="55595DE2FFE70055FBB3FBB7FB21FBDC" box="[1070,1275,1026,1048]" gridcol="5" gridrow="16" pageId="36" pageNumber="37">8223</td>
<td id="55595DE2FFE70055FABFFBB7FA67FBDC" box="[1314,1469,1026,1048]" gridcol="6" gridrow="16" pageId="36" pageNumber="37">35</td>
</tr>
<tr id="1688349EFFE70055FFEDFB99FA67FB86" box="[112,1469,1068,1090]" gridrow="17" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFB99FECCFB86" box="[112,278,1068,1090]" gridcol="0" gridrow="17" pageId="36" pageNumber="37">FMNH PR 2081</th>
<td id="55595DE2FFE70055FEA5FB99FE07FB86" box="[312,477,1068,1090]" gridcol="1" gridrow="17" pageId="36" pageNumber="37">Sue</td>
<td id="55595DE2FFE70055FD95FB99FD41FB86" box="[520,667,1068,1090]" gridcol="2" gridrow="17" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD95FB98FD41FB86" authorityName=", Sue FMNH PR" authorityYear="2081" box="[520,667,1069,1090]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD95FB98FD41FB86" box="[520,667,1069,1090]" italics="true" pageId="36" pageNumber="37">Tyrannosaurus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FB99FC98FB86" box="[714,834,1068,1090]" gridcol="3" gridrow="17" pageId="36" pageNumber="37">Periosteum</td>
<td id="55595DE2FFE70055FCF7FB99FBD1FB86" box="[874,1035,1068,1090]" gridcol="4" gridrow="17" pageId="36" pageNumber="37">579</td>
<td id="55595DE2FFE70055FBB3FB99FB21FB86" box="[1070,1275,1068,1090]" gridcol="5" gridrow="17" pageId="36" pageNumber="37">8422</td>
<td id="55595DE2FFE70055FABFFB99FA67FB86" box="[1314,1469,1068,1090]" gridcol="6" gridrow="17" pageId="36" pageNumber="37">199</td>
</tr>
<tr id="1688349EFFE70055FFEDFBE3FA67FBA8" box="[112,1469,1110,1132]" gridrow="18" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFBE3FECCFBA8" box="[112,278,1110,1132]" gridcol="0" gridrow="18" pageId="36" pageNumber="37">BMRP 2006.4.4</th>
<td id="55595DE2FFE70055FEA5FBE3FE07FBA8" box="[312,477,1110,1132]" gridcol="1" gridrow="18" pageId="36" pageNumber="37">Petey</td>
<td id="55595DE2FFE70055FD95FBE3FD41FBA8" box="[520,667,1110,1132]" gridcol="2" gridrow="18" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96FBE2FD43FBA8" box="[523,665,1111,1132]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96FBE2FD43FBA8" box="[523,665,1111,1132]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FBE3FC98FBA8" box="[714,834,1110,1132]" gridcol="3" gridrow="18" pageId="36" pageNumber="37">1</td>
<td id="55595DE2FFE70055FCF7FBE3FBD1FBA8" box="[874,1035,1110,1132]" gridcol="4" gridrow="18" pageId="36" pageNumber="37">148.4</td>
<td id="55595DE2FFE70055FBB3FBE3FB21FBA8" box="[1070,1275,1110,1132]" gridcol="5" gridrow="18" pageId="36" pageNumber="37">198</td>
<td id="55595DE2FFE70055FABFFBE3FA67FBA8" box="[1314,1469,1110,1132]" gridcol="6" gridrow="18" pageId="36" pageNumber="37">NA</td>
</tr>
<tr id="1688349EFFE70055FFEDFB35FA67FB52" box="[112,1469,1152,1174]" gridrow="19" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFB35FECCFB52" box="[112,278,1152,1174]" gridcol="0" gridrow="19" pageId="36" pageNumber="37">BMRP 2006.4.4</th>
<td id="55595DE2FFE70055FEA5FB35FE07FB52" box="[312,477,1152,1174]" gridcol="1" gridrow="19" pageId="36" pageNumber="37">Petey</td>
<td id="55595DE2FFE70055FD95FB35FD41FB52" box="[520,667,1152,1174]" gridcol="2" gridrow="19" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96FB34FD43FB52" box="[523,665,1153,1174]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96FB34FD43FB52" box="[523,665,1153,1174]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FB35FC98FB52" box="[714,834,1152,1174]" gridcol="3" gridrow="19" pageId="36" pageNumber="37">2</td>
<td id="55595DE2FFE70055FCF7FB35FBD1FB52" box="[874,1035,1152,1174]" gridcol="4" gridrow="19" pageId="36" pageNumber="37">165.9</td>
<td id="55595DE2FFE70055FBB3FB35FB21FB52" box="[1070,1275,1152,1174]" gridcol="5" gridrow="19" pageId="36" pageNumber="37">269</td>
<td id="55595DE2FFE70055FABFFB35FA67FB52" box="[1314,1469,1152,1174]" gridcol="6" gridrow="19" pageId="36" pageNumber="37">71</td>
</tr>
<tr id="1688349EFFE70055FFEDFB1FFA67FB04" box="[112,1469,1194,1216]" gridrow="20" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFB1FFECCFB04" box="[112,278,1194,1216]" gridcol="0" gridrow="20" pageId="36" pageNumber="37">BMRP 2006.4.4</th>
<td id="55595DE2FFE70055FEA5FB1FFE07FB04" box="[312,477,1194,1216]" gridcol="1" gridrow="20" pageId="36" pageNumber="37">Petey</td>
<td id="55595DE2FFE70055FD95FB1FFD41FB04" box="[520,667,1194,1216]" gridcol="2" gridrow="20" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96FB1EFD43FB04" box="[523,665,1195,1216]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96FB1EFD43FB04" box="[523,665,1195,1216]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FB1FFC98FB04" box="[714,834,1194,1216]" gridcol="3" gridrow="20" pageId="36" pageNumber="37">3</td>
<td id="55595DE2FFE70055FCF7FB1FFBD1FB04" box="[874,1035,1194,1216]" gridcol="4" gridrow="20" pageId="36" pageNumber="37">194.7</td>
<td id="55595DE2FFE70055FBB3FB1FFB21FB04" box="[1070,1275,1194,1216]" gridcol="5" gridrow="20" pageId="36" pageNumber="37">419</td>
<td id="55595DE2FFE70055FABFFB1FFA67FB04" box="[1314,1469,1194,1216]" gridcol="6" gridrow="20" pageId="36" pageNumber="37">149</td>
</tr>
<tr id="1688349EFFE70055FFEDFB61FA67FB2E" box="[112,1469,1236,1258]" gridrow="21" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFB61FECCFB2E" box="[112,278,1236,1258]" gridcol="0" gridrow="21" pageId="36" pageNumber="37">BMRP 2006.4.4</th>
<td id="55595DE2FFE70055FEA5FB61FE07FB2E" box="[312,477,1236,1258]" gridcol="1" gridrow="21" pageId="36" pageNumber="37">Petey</td>
<td id="55595DE2FFE70055FD95FB61FD41FB2E" box="[520,667,1236,1258]" gridcol="2" gridrow="21" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96FB60FD43FB2E" box="[523,665,1237,1258]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96FB60FD43FB2E" box="[523,665,1237,1258]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FB61FC98FB2E" box="[714,834,1236,1258]" gridcol="3" gridrow="21" pageId="36" pageNumber="37">4</td>
<td id="55595DE2FFE70055FCF7FB61FBD1FB2E" box="[874,1035,1236,1258]" gridcol="4" gridrow="21" pageId="36" pageNumber="37">214.5</td>
<td id="55595DE2FFE70055FBB3FB61FB21FB2E" box="[1070,1275,1236,1258]" gridcol="5" gridrow="21" pageId="36" pageNumber="37">547</td>
<td id="55595DE2FFE70055FABFFB61FA67FB2E" box="[1314,1469,1236,1258]" gridcol="6" gridrow="21" pageId="36" pageNumber="37">128</td>
</tr>
<tr id="1688349EFFE70055FFEDFB4BFA67FAD0" box="[112,1469,1278,1300]" gridrow="22" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFB4BFECCFAD0" box="[112,278,1278,1300]" gridcol="0" gridrow="22" pageId="36" pageNumber="37">BMRP 2006.4.4</th>
<td id="55595DE2FFE70055FEA5FB4BFE07FAD0" box="[312,477,1278,1300]" gridcol="1" gridrow="22" pageId="36" pageNumber="37">Petey</td>
<td id="55595DE2FFE70055FD95FB4BFD41FAD0" box="[520,667,1278,1300]" gridcol="2" gridrow="22" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96FB4AFD43FAD0" box="[523,665,1279,1300]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96FB4AFD43FAD0" box="[523,665,1279,1300]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FB4BFC98FAD0" box="[714,834,1278,1300]" gridcol="3" gridrow="22" pageId="36" pageNumber="37">5</td>
<td id="55595DE2FFE70055FCF7FB4BFBD1FAD0" box="[874,1035,1278,1300]" gridcol="4" gridrow="22" pageId="36" pageNumber="37">223.6</td>
<td id="55595DE2FFE70055FBB3FB4BFB21FAD0" box="[1070,1275,1278,1300]" gridcol="5" gridrow="22" pageId="36" pageNumber="37">613</td>
<td id="55595DE2FFE70055FABFFB4BFA67FAD0" box="[1314,1469,1278,1300]" gridcol="6" gridrow="22" pageId="36" pageNumber="37">66</td>
</tr>
<tr id="1688349EFFE70055FFEDFA9DFA67FAFB" box="[112,1469,1320,1343]" gridrow="23" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFA9DFECCFAFB" box="[112,278,1320,1343]" gridcol="0" gridrow="23" pageId="36" pageNumber="37">BMRP 2006.4.4</th>
<td id="55595DE2FFE70055FEA5FA9DFE07FAFB" box="[312,477,1320,1343]" gridcol="1" gridrow="23" pageId="36" pageNumber="37">Petey</td>
<td id="55595DE2FFE70055FD95FA9DFD41FAFB" box="[520,667,1320,1343]" gridcol="2" gridrow="23" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96FA9CFD43FAFA" box="[523,665,1321,1342]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96FA9CFD43FAFA" box="[523,665,1321,1342]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FA9DFC98FAFB" box="[714,834,1320,1343]" gridcol="3" gridrow="23" pageId="36" pageNumber="37">6</td>
<td id="55595DE2FFE70055FCF7FA9DFBD1FAFB" box="[874,1035,1320,1343]" gridcol="4" gridrow="23" pageId="36" pageNumber="37">229.4</td>
<td id="55595DE2FFE70055FBB3FA9DFB21FAFB" box="[1070,1275,1320,1343]" gridcol="5" gridrow="23" pageId="36" pageNumber="37">658</td>
<td id="55595DE2FFE70055FABFFA9DFA67FAFB" box="[1314,1469,1320,1343]" gridcol="6" gridrow="23" pageId="36" pageNumber="37">45</td>
</tr>
<tr id="1688349EFFE70055FFEDFAE6FA67FAAD" box="[112,1469,1363,1385]" gridrow="24" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFAE6FECCFAAD" box="[112,278,1363,1385]" gridcol="0" gridrow="24" pageId="36" pageNumber="37">BMRP 2006.4.4</th>
<td id="55595DE2FFE70055FEA5FAE6FE07FAAD" box="[312,477,1363,1385]" gridcol="1" gridrow="24" pageId="36" pageNumber="37">Petey</td>
<td id="55595DE2FFE70055FD95FAE6FD41FAAD" box="[520,667,1363,1385]" gridcol="2" gridrow="24" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96FAE6FD43FAAC" box="[523,665,1363,1384]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96FAE6FD43FAAC" box="[523,665,1363,1384]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FAE6FC98FAAD" box="[714,834,1363,1385]" gridcol="3" gridrow="24" pageId="36" pageNumber="37">7</td>
<td id="55595DE2FFE70055FCF7FAE6FBD1FAAD" box="[874,1035,1363,1385]" gridcol="4" gridrow="24" pageId="36" pageNumber="37">234.8</td>
<td id="55595DE2FFE70055FBB3FAE6FB21FAAD" box="[1070,1275,1363,1385]" gridcol="5" gridrow="24" pageId="36" pageNumber="37">701</td>
<td id="55595DE2FFE70055FABFFAE6FA67FAAD" box="[1314,1469,1363,1385]" gridcol="6" gridrow="24" pageId="36" pageNumber="37">44</td>
</tr>
<tr id="1688349EFFE70055FFEDFAC8FA67FA57" box="[112,1469,1405,1427]" gridrow="25" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFAC8FECCFA57" box="[112,278,1405,1427]" gridcol="0" gridrow="25" pageId="36" pageNumber="37">BMRP 2006.4.4</th>
<td id="55595DE2FFE70055FEA5FAC8FE07FA57" box="[312,477,1405,1427]" gridcol="1" gridrow="25" pageId="36" pageNumber="37">Petey</td>
<td id="55595DE2FFE70055FD95FAC8FD41FA57" box="[520,667,1405,1427]" gridcol="2" gridrow="25" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96FAC8FD43FA56" box="[523,665,1405,1426]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96FAC8FD43FA56" box="[523,665,1405,1426]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FAC8FC98FA57" box="[714,834,1405,1427]" gridcol="3" gridrow="25" pageId="36" pageNumber="37">Periosteum</td>
<td id="55595DE2FFE70055FCF7FAC8FBD1FA57" box="[874,1035,1405,1427]" gridcol="4" gridrow="25" pageId="36" pageNumber="37">244.7</td>
<td id="55595DE2FFE70055FBB3FAC8FB21FA57" box="[1070,1275,1405,1427]" gridcol="5" gridrow="25" pageId="36" pageNumber="37">786</td>
<td id="55595DE2FFE70055FABFFAC8FA67FA57" box="[1314,1469,1405,1427]" gridcol="6" gridrow="25" pageId="36" pageNumber="37">84</td>
</tr>
<tr id="1688349EFFE70055FFEDFA12FA67FA79" box="[112,1469,1447,1469]" gridrow="26" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFA12FECCFA79" box="[112,278,1447,1469]" gridcol="0" gridrow="26" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5FA12FE07FA79" box="[312,477,1447,1469]" gridcol="1" gridrow="26" pageId="36" pageNumber="37">Jane</td>
<td id="55595DE2FFE70055FD95FA12FD41FA79" box="[520,667,1447,1469]" gridcol="2" gridrow="26" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96FA12FD43FA78" box="[523,665,1447,1468]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96FA12FD43FA78" box="[523,665,1447,1468]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FA12FC98FA79" box="[714,834,1447,1469]" gridcol="3" gridrow="26" pageId="36" pageNumber="37">1</td>
<td id="55595DE2FFE70055FCF7FA12FBD1FA79" box="[874,1035,1447,1469]" gridcol="4" gridrow="26" pageId="36" pageNumber="37">195.9</td>
<td id="55595DE2FFE70055FBB3FA12FB21FA79" box="[1070,1275,1447,1469]" gridcol="5" gridrow="26" pageId="36" pageNumber="37">426</td>
<td id="55595DE2FFE70055FABFFA12FA67FA79" box="[1314,1469,1447,1469]" gridcol="6" gridrow="26" pageId="36" pageNumber="37">NA</td>
</tr>
<tr id="1688349EFFE70055FFEDFA64FA67FA23" box="[112,1469,1489,1511]" gridrow="27" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFA64FECCFA23" box="[112,278,1489,1511]" gridcol="0" gridrow="27" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5FA64FE07FA23" box="[312,477,1489,1511]" gridcol="1" gridrow="27" pageId="36" pageNumber="37">Jane</td>
<td id="55595DE2FFE70055FD95FA64FD41FA23" box="[520,667,1489,1511]" gridcol="2" gridrow="27" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96FA64FD43FA22" box="[523,665,1489,1510]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96FA64FD43FA22" box="[523,665,1489,1510]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FA64FC98FA23" box="[714,834,1489,1511]" gridcol="3" gridrow="27" pageId="36" pageNumber="37">2</td>
<td id="55595DE2FFE70055FCF7FA64FBD1FA23" box="[874,1035,1489,1511]" gridcol="4" gridrow="27" pageId="36" pageNumber="37">218.4</td>
<td id="55595DE2FFE70055FBB3FA64FB21FA23" box="[1070,1275,1489,1511]" gridcol="5" gridrow="27" pageId="36" pageNumber="37">575</td>
<td id="55595DE2FFE70055FABFFA64FA67FA23" box="[1314,1469,1489,1511]" gridcol="6" gridrow="27" pageId="36" pageNumber="37">149</td>
</tr>
<tr id="1688349EFFE70055FFEDFA4EFA67F9D5" box="[112,1469,1531,1553]" gridrow="28" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDFA4EFECCF9D5" box="[112,278,1531,1553]" gridcol="0" gridrow="28" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5FA4EFE07F9D5" box="[312,477,1531,1553]" gridcol="1" gridrow="28" pageId="36" pageNumber="37">Jane</td>
<td id="55595DE2FFE70055FD95FA4EFD41F9D5" box="[520,667,1531,1553]" gridcol="2" gridrow="28" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96FA4EFD43F9D4" box="[523,665,1531,1552]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96FA4EFD43F9D4" box="[523,665,1531,1552]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57FA4EFC98F9D5" box="[714,834,1531,1553]" gridcol="3" gridrow="28" pageId="36" pageNumber="37">3</td>
<td id="55595DE2FFE70055FCF7FA4EFBD1F9D5" box="[874,1035,1531,1553]" gridcol="4" gridrow="28" pageId="36" pageNumber="37">231.8</td>
<td id="55595DE2FFE70055FBB3FA4EFB21F9D5" box="[1070,1275,1531,1553]" gridcol="5" gridrow="28" pageId="36" pageNumber="37">677</td>
<td id="55595DE2FFE70055FABFFA4EFA67F9D5" box="[1314,1469,1531,1553]" gridcol="6" gridrow="28" pageId="36" pageNumber="37">102</td>
</tr>
<tr id="1688349EFFE70055FFEDF990FA67F9FE" box="[112,1469,1573,1594]" gridrow="29" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDF990FECCF9FE" box="[112,278,1573,1594]" gridcol="0" gridrow="29" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5F990FE07F9FE" box="[312,477,1573,1594]" gridcol="1" gridrow="29" pageId="36" pageNumber="37">Jane</td>
<td id="55595DE2FFE70055FD95F990FD41F9FE" box="[520,667,1573,1594]" gridcol="2" gridrow="29" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96F990FD43F9FE" box="[523,665,1573,1594]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96F990FD43F9FE" box="[523,665,1573,1594]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57F990FC98F9FE" box="[714,834,1573,1594]" gridcol="3" gridrow="29" pageId="36" pageNumber="37">4</td>
<td id="55595DE2FFE70055FCF7F990FBD1F9FE" box="[874,1035,1573,1594]" gridcol="4" gridrow="29" pageId="36" pageNumber="37">247.3</td>
<td id="55595DE2FFE70055FBB3F990FB21F9FE" box="[1070,1275,1573,1594]" gridcol="5" gridrow="29" pageId="36" pageNumber="37">809</td>
<td id="55595DE2FFE70055FABFF990FA67F9FE" box="[1314,1469,1573,1594]" gridcol="6" gridrow="29" pageId="36" pageNumber="37">132</td>
</tr>
<tr id="1688349EFFE70055FFEDF9FAFA67F9A1" box="[112,1469,1615,1637]" gridrow="30" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDF9FAFECCF9A1" box="[112,278,1615,1637]" gridcol="0" gridrow="30" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5F9FAFE07F9A1" box="[312,477,1615,1637]" gridcol="1" gridrow="30" pageId="36" pageNumber="37">Jane</td>
<td id="55595DE2FFE70055FD95F9FAFD41F9A1" box="[520,667,1615,1637]" gridcol="2" gridrow="30" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96F9FAFD43F9A0" box="[523,665,1615,1636]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96F9FAFD43F9A0" box="[523,665,1615,1636]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57F9FAFC98F9A1" box="[714,834,1615,1637]" gridcol="3" gridrow="30" pageId="36" pageNumber="37">5</td>
<td id="55595DE2FFE70055FCF7F9FAFBD1F9A1" box="[874,1035,1615,1637]" gridcol="4" gridrow="30" pageId="36" pageNumber="37">248.6</td>
<td id="55595DE2FFE70055FBB3F9FAFB21F9A1" box="[1070,1275,1615,1637]" gridcol="5" gridrow="30" pageId="36" pageNumber="37">820</td>
<td id="55595DE2FFE70055FABFF9FAFA67F9A1" box="[1314,1469,1615,1637]" gridcol="6" gridrow="30" pageId="36" pageNumber="37">11</td>
</tr>
<tr id="1688349EFFE70055FFEDF9CCFA67F94B" box="[112,1469,1657,1679]" gridrow="31" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDF9CCFECCF94B" box="[112,278,1657,1679]" gridcol="0" gridrow="31" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5F9CCFE07F94B" box="[312,477,1657,1679]" gridcol="1" gridrow="31" pageId="36" pageNumber="37">Jane</td>
<td id="55595DE2FFE70055FD95F9CCFD41F94B" box="[520,667,1657,1679]" gridcol="2" gridrow="31" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96F9CCFD43F94A" box="[523,665,1657,1678]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96F9CCFD43F94A" box="[523,665,1657,1678]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57F9CCFC98F94B" box="[714,834,1657,1679]" gridcol="3" gridrow="31" pageId="36" pageNumber="37">6</td>
<td id="55595DE2FFE70055FCF7F9CCFBD1F94B" box="[874,1035,1657,1679]" gridcol="4" gridrow="31" pageId="36" pageNumber="37">255.3</td>
<td id="55595DE2FFE70055FBB3F9CCFB21F94B" box="[1070,1275,1657,1679]" gridcol="5" gridrow="31" pageId="36" pageNumber="37">884</td>
<td id="55595DE2FFE70055FABFF9CCFA67F94B" box="[1314,1469,1657,1679]" gridcol="6" gridrow="31" pageId="36" pageNumber="37">63</td>
</tr>
<tr id="1688349EFFE70055FFEDF916FA67F97D" box="[112,1469,1699,1721]" gridrow="32" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDF916FECCF97D" box="[112,278,1699,1721]" gridcol="0" gridrow="32" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5F916FE07F97D" box="[312,477,1699,1721]" gridcol="1" gridrow="32" pageId="36" pageNumber="37">Jane</td>
<td id="55595DE2FFE70055FD95F916FD41F97D" box="[520,667,1699,1721]" gridcol="2" gridrow="32" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96F911FD43F97D" box="[523,665,1700,1721]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96F911FD43F97D" box="[523,665,1700,1721]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57F916FC98F97D" box="[714,834,1699,1721]" gridcol="3" gridrow="32" pageId="36" pageNumber="37">7</td>
<td id="55595DE2FFE70055FCF7F916FBD1F97D" box="[874,1035,1699,1721]" gridcol="4" gridrow="32" pageId="36" pageNumber="37">257.7</td>
<td id="55595DE2FFE70055FBB3F916FB21F97D" box="[1070,1275,1699,1721]" gridcol="5" gridrow="32" pageId="36" pageNumber="37">906</td>
<td id="55595DE2FFE70055FABFF916FA67F97D" box="[1314,1469,1699,1721]" gridcol="6" gridrow="32" pageId="36" pageNumber="37">23</td>
</tr>
<tr id="1688349EFFE70055FFEDF978FA67F927" box="[112,1469,1741,1763]" gridrow="33" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDF978FECCF927" box="[112,278,1741,1763]" gridcol="0" gridrow="33" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5F978FE07F927" box="[312,477,1741,1763]" gridcol="1" gridrow="33" pageId="36" pageNumber="37">Jane</td>
<td id="55595DE2FFE70055FD95F978FD41F927" box="[520,667,1741,1763]" gridcol="2" gridrow="33" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96F97BFD43F927" box="[523,665,1742,1763]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96F97BFD43F927" box="[523,665,1742,1763]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57F978FC98F927" box="[714,834,1741,1763]" gridcol="3" gridrow="33" pageId="36" pageNumber="37">8</td>
<td id="55595DE2FFE70055FCF7F978FBD1F927" box="[874,1035,1741,1763]" gridcol="4" gridrow="33" pageId="36" pageNumber="37">265.7</td>
<td id="55595DE2FFE70055FBB3F978FB21F927" box="[1070,1275,1741,1763]" gridcol="5" gridrow="33" pageId="36" pageNumber="37">986</td>
<td id="55595DE2FFE70055FABFF978FA67F927" box="[1314,1469,1741,1763]" gridcol="6" gridrow="33" pageId="36" pageNumber="37">79</td>
</tr>
<tr id="1688349EFFE70055FFEDF942FA67F8C9" box="[112,1469,1783,1805]" gridrow="34" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDF942FECCF8C9" box="[112,278,1783,1805]" gridcol="0" gridrow="34" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5F942FE07F8C9" box="[312,477,1783,1805]" gridcol="1" gridrow="34" pageId="36" pageNumber="37">Jane</td>
<td id="55595DE2FFE70055FD95F942FD41F8C9" box="[520,667,1783,1805]" gridcol="2" gridrow="34" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96F94DFD43F8C9" box="[523,665,1784,1805]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96F94DFD43F8C9" box="[523,665,1784,1805]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57F942FC98F8C9" box="[714,834,1783,1805]" gridcol="3" gridrow="34" pageId="36" pageNumber="37">9</td>
<td id="55595DE2FFE70055FCF7F942FBD1F8C9" box="[874,1035,1783,1805]" gridcol="4" gridrow="34" pageId="36" pageNumber="37">266.6</td>
<td id="55595DE2FFE70055FBB3F942FB21F8C9" box="[1070,1275,1783,1805]" gridcol="5" gridrow="34" pageId="36" pageNumber="37">995</td>
<td id="55595DE2FFE70055FABFF942FA67F8C9" box="[1314,1469,1783,1805]" gridcol="6" gridrow="34" pageId="36" pageNumber="37">9</td>
</tr>
<tr id="1688349EFFE70055FFEDF894FA67F8F3" box="[112,1469,1825,1847]" gridrow="35" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDF894FECCF8F3" box="[112,278,1825,1847]" gridcol="0" gridrow="35" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5F894FE07F8F3" box="[312,477,1825,1847]" gridcol="1" gridrow="35" pageId="36" pageNumber="37">Jane</td>
<td id="55595DE2FFE70055FD95F894FD41F8F3" box="[520,667,1825,1847]" gridcol="2" gridrow="35" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96F897FD43F8F3" box="[523,665,1826,1847]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96F897FD43F8F3" box="[523,665,1826,1847]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57F894FC98F8F3" box="[714,834,1825,1847]" gridcol="3" gridrow="35" pageId="36" pageNumber="37">Periosteum</td>
<td id="55595DE2FFE70055FCF7F894FBD1F8F3" box="[874,1035,1825,1847]" gridcol="4" gridrow="35" pageId="36" pageNumber="37">273.2</td>
<td id="55595DE2FFE70055FBB3F894FB21F8F3" box="[1070,1275,1825,1847]" gridcol="5" gridrow="35" pageId="36" pageNumber="37">1064</td>
<td id="55595DE2FFE70055FABFF894FA67F8F3" box="[1314,1469,1825,1847]" gridcol="6" gridrow="35" pageId="36" pageNumber="37">69</td>
</tr>
<tr id="1688349EFFE70055FFEDF8FEFA67F8A5" box="[112,1469,1867,1889]" gridrow="36" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDF8FEFECCF8A5" box="[112,278,1867,1889]" gridcol="0" gridrow="36" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5F8FEFE07F8A5" box="[312,477,1867,1889]" gridcol="1" gridrow="36" pageId="36" pageNumber="37">Jane (corrected)</td>
<td id="55595DE2FFE70055FD95F8FEFD41F8A5" box="[520,667,1867,1889]" gridcol="2" gridrow="36" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96F8F9FD43F8A5" box="[523,665,1868,1889]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96F8F9FD43F8A5" box="[523,665,1868,1889]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57F8FEFC98F8A5" box="[714,834,1867,1889]" gridcol="3" gridrow="36" pageId="36" pageNumber="37">1</td>
<td id="55595DE2FFE70055FCF7F8FEFBD1F8A5" box="[874,1035,1867,1889]" gridcol="4" gridrow="36" pageId="36" pageNumber="37">195.9</td>
<td id="55595DE2FFE70055FBB3F8FEFB21F8A5" box="[1070,1275,1867,1889]" gridcol="5" gridrow="36" pageId="36" pageNumber="37">426</td>
<td id="55595DE2FFE70055FABFF8FEFA67F8A5" box="[1314,1469,1867,1889]" gridcol="6" gridrow="36" pageId="36" pageNumber="37">NA</td>
</tr>
<tr id="1688349EFFE70055FFEDF8C0FA67F84F" box="[112,1469,1909,1931]" gridrow="37" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDF8C0FECCF84F" box="[112,278,1909,1931]" gridcol="0" gridrow="37" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5F8C0FE07F84F" box="[312,477,1909,1931]" gridcol="1" gridrow="37" pageId="36" pageNumber="37">Jane (corrected)</td>
<td id="55595DE2FFE70055FD95F8C0FD41F84F" box="[520,667,1909,1931]" gridcol="2" gridrow="37" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96F8C3FD43F84F" box="[523,665,1910,1931]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96F8C3FD43F84F" box="[523,665,1910,1931]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57F8C0FC98F84F" box="[714,834,1909,1931]" gridcol="3" gridrow="37" pageId="36" pageNumber="37">2</td>
<td id="55595DE2FFE70055FCF7F8C0FBD1F84F" box="[874,1035,1909,1931]" gridcol="4" gridrow="37" pageId="36" pageNumber="37">218.4</td>
<td id="55595DE2FFE70055FBB3F8C0FB21F84F" box="[1070,1275,1909,1931]" gridcol="5" gridrow="37" pageId="36" pageNumber="37">575</td>
<td id="55595DE2FFE70055FABFF8C0FA67F84F" box="[1314,1469,1909,1931]" gridcol="6" gridrow="37" pageId="36" pageNumber="37">149</td>
</tr>
<tr id="1688349EFFE70055FFEDF82AFA67F871" box="[112,1469,1951,1973]" gridrow="38" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDF82AFECCF871" box="[112,278,1951,1973]" gridcol="0" gridrow="38" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5F82AFE07F871" box="[312,477,1951,1973]" gridcol="1" gridrow="38" pageId="36" pageNumber="37">Jane (corrected)</td>
<td id="55595DE2FFE70055FD95F82AFD41F871" box="[520,667,1951,1973]" gridcol="2" gridrow="38" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96F815FD43F871" box="[523,665,1952,1973]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96F815FD43F871" box="[523,665,1952,1973]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57F82AFC98F871" box="[714,834,1951,1973]" gridcol="3" gridrow="38" pageId="36" pageNumber="37">3</td>
<td id="55595DE2FFE70055FCF7F82AFBD1F871" box="[874,1035,1951,1973]" gridcol="4" gridrow="38" pageId="36" pageNumber="37">231.8</td>
<td id="55595DE2FFE70055FBB3F82AFB21F871" box="[1070,1275,1951,1973]" gridcol="5" gridrow="38" pageId="36" pageNumber="37">677</td>
<td id="55595DE2FFE70055FABFF82AFA67F871" box="[1314,1469,1951,1973]" gridcol="6" gridrow="38" pageId="36" pageNumber="37">102</td>
</tr>
<tr id="1688349EFFE70055FFEDF87CFA67F81B" box="[112,1469,1993,2015]" gridrow="39" pageId="36" pageNumber="37">
<th id="55595DE2FFE70055FFEDF87CFECCF81B" box="[112,278,1993,2015]" gridcol="0" gridrow="39" pageId="36" pageNumber="37">BMRP 2002.4.4</th>
<td id="55595DE2FFE70055FEA5F87CFE07F81B" box="[312,477,1993,2015]" gridcol="1" gridrow="39" pageId="36" pageNumber="37">Jane (corrected)</td>
<td id="55595DE2FFE70055FD95F87CFD41F81B" box="[520,667,1993,2015]" gridcol="2" gridrow="39" pageId="36" pageNumber="37">
<taxonomicName id="6FB84D5FFFE7FF8EFD96F87FFD43F81B" box="[523,665,1994,2015]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="36" pageNumber="37" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE7FF8EFD96F87FFD43F81B" box="[523,665,1994,2015]" italics="true" pageId="36" pageNumber="37">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE70055FD57F87CFC98F81B" box="[714,834,1993,2015]" gridcol="3" gridrow="39" pageId="36" pageNumber="37">4</td>
<td id="55595DE2FFE70055FCF7F87CFBD1F81B" box="[874,1035,1993,2015]" gridcol="4" gridrow="39" pageId="36" pageNumber="37">248.6</td>
<td id="55595DE2FFE70055FBB3F87CFB21F81B" box="[1070,1275,1993,2015]" gridcol="5" gridrow="39" pageId="36" pageNumber="37">820</td>
<td id="55595DE2FFE70055FABFF87CFA67F81B" box="[1314,1469,1993,2015]" gridcol="6" gridrow="39" pageId="36" pageNumber="37">144</td>
</tr>
</table>
</paragraph>
<caption id="FCC76654FFE6FF8FFE26FF5AFD97FEC1" box="[443,589,239,261]" isContinuationCaption="true" pageId="37" pageNumber="38" startId="37.[443,503,239,260]" targetType="table">
<paragraph id="A80736DCFFE6FF8FFE26FF5AFD97FEC1" blockId="37.[443,589,239,261]" box="[443,589,239,261]" pageId="37" pageNumber="38">
<emphasis id="9ACCEACEFFE6FF8FFE26FF5AFDD5FEC0" bold="true" box="[443,527,239,260]" pageId="37" pageNumber="38">Table 5.</emphasis>
Cont.
</paragraph>
</caption>
<paragraph id="A80736DCFFE6FF8FFF10FEF1FAA1FE32" pageId="37" pageNumber="38">
<table id="DAB8C47CFFE60055FFE9FE92FA67FE32" box="[116,1469,295,502]" colsContinueFrom="36.[112,1469,295,2015]" gridcols="7" gridrows="4" pageId="37" pageNumber="38">
<tr id="1688349EFFE60055FFE9FE92FA67FEBC" box="[116,1469,295,376]" gridrow="0" pageId="37" pageNumber="38">
<th id="55595DE2FFE60055FFE9FE92FEC8FEBC" box="[116,274,295,376]" gridcol="0" gridrow="0" pageId="37" pageNumber="38">
<emphasis id="9ACCEACEFFE6FF8FFF10FEF1FF23FE9E" bold="true" box="[141,249,324,346]" pageId="37" pageNumber="38">Specimen</emphasis>
</th>
<th id="55595DE2FFE60055FEA5FE92FE07FEBC" box="[312,477,295,376]" gridcol="1" gridrow="0" pageId="37" pageNumber="38">
<emphasis id="9ACCEACEFFE6FF8FFECFFEF1FE19FE9E" bold="true" box="[338,451,324,346]" pageId="37" pageNumber="38">Nickname</emphasis>
</th>
<th id="55595DE2FFE60055FD96FE92FD43FEBC" box="[523,665,295,376]" gridcol="2" gridrow="0" pageId="37" pageNumber="38">
<emphasis id="9ACCEACEFFE6FF8FFDACFEF0FDA8FE9E" bold="true" box="[561,626,325,346]" pageId="37" pageNumber="38">Taxon</emphasis>
</th>
<th id="55595DE2FFE60055FD57FE92FC98FEBC" box="[714,834,295,376]" gridcol="3" gridrow="0" pageId="37" pageNumber="38">
<emphasis id="9ACCEACEFFE6FF8FFD48FEF1FCEDFE9E" bold="true" box="[725,823,324,346]" pageId="37" pageNumber="38">LAG No.</emphasis>
</th>
<th id="55595DE2FFE60055FCF7FE92FBD1FEBC" box="[874,1035,295,376]" gridcol="4" gridrow="0" pageId="37" pageNumber="38">
<emphasis id="9ACCEACEFFE6FF8FFCF7FE83FC0DFEAD" bold="true" box="[874,1035,310,361]" pageId="37" pageNumber="38">Circumference (mm)</emphasis>
</th>
<th id="55595DE2FFE60055FBB3FE92FB21FEBC" box="[1070,1275,295,376]" gridcol="5" gridrow="0" pageId="37" pageNumber="38">
<emphasis id="9ACCEACEFFE6FF8FFBFDFE9DFB21FEBC" bold="true" pageId="37" pageNumber="38">Mass (kg) (Campione et al., 2014), Equation (7))</emphasis>
</th>
<th id="55595DE2FFE60055FABFFE92FA67FEBC" box="[1314,1469,295,376]" gridcol="6" gridrow="0" pageId="37" pageNumber="38">
<emphasis id="9ACCEACEFFE6FF8FFABFFE9DFA78FEBC" bold="true" pageId="37" pageNumber="38">Mass Increase from Previous LAG (kg)</emphasis>
</th>
</tr>
<tr id="1688349EFFE60055FFE9FE39FA67FE66" box="[116,1469,396,418]" gridrow="1" pageId="37" pageNumber="38">
<th id="55595DE2FFE60055FFE9FE39FEC8FE66" box="[116,274,396,418]" gridcol="0" gridrow="1" pageId="37" pageNumber="38">BMRP 2002.4.4</th>
<td id="55595DE2FFE60055FEA5FE39FE07FE66" box="[312,477,396,418]" gridcol="1" gridrow="1" pageId="37" pageNumber="38">Jane (corrected)</td>
<td id="55595DE2FFE60055FD96FE39FD43FE66" box="[523,665,396,418]" gridcol="2" gridrow="1" pageId="37" pageNumber="38">
<taxonomicName id="6FB84D5FFFE6FF8FFD96FE38FD43FE66" box="[523,665,397,418]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE6FF8FFD96FE38FD43FE66" box="[523,665,397,418]" italics="true" pageId="37" pageNumber="38">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE60055FD57FE39FC98FE66" box="[714,834,396,418]" gridcol="3" gridrow="1" pageId="37" pageNumber="38">5</td>
<td id="55595DE2FFE60055FCF7FE39FBD1FE66" box="[874,1035,396,418]" gridcol="4" gridrow="1" pageId="37" pageNumber="38">257.7</td>
<td id="55595DE2FFE60055FBB3FE39FB21FE66" box="[1070,1275,396,418]" gridcol="5" gridrow="1" pageId="37" pageNumber="38">906</td>
<td id="55595DE2FFE60055FABFFE39FA67FE66" box="[1314,1469,396,418]" gridcol="6" gridrow="1" pageId="37" pageNumber="38">86</td>
</tr>
<tr id="1688349EFFE60055FFE9FE03FA67FE08" box="[116,1469,438,460]" gridrow="2" pageId="37" pageNumber="38">
<th id="55595DE2FFE60055FFE9FE03FEC8FE08" box="[116,274,438,460]" gridcol="0" gridrow="2" pageId="37" pageNumber="38">BMRP 2002.4.4</th>
<td id="55595DE2FFE60055FEA5FE03FE07FE08" box="[312,477,438,460]" gridcol="1" gridrow="2" pageId="37" pageNumber="38">Jane (corrected)</td>
<td id="55595DE2FFE60055FD96FE03FD43FE08" box="[523,665,438,460]" gridcol="2" gridrow="2" pageId="37" pageNumber="38">
<taxonomicName id="6FB84D5FFFE6FF8FFD96FE02FD43FE08" box="[523,665,439,460]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE6FF8FFD96FE02FD43FE08" box="[523,665,439,460]" italics="true" pageId="37" pageNumber="38">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE60055FD57FE03FC98FE08" box="[714,834,438,460]" gridcol="3" gridrow="2" pageId="37" pageNumber="38">6</td>
<td id="55595DE2FFE60055FCF7FE03FBD1FE08" box="[874,1035,438,460]" gridcol="4" gridrow="2" pageId="37" pageNumber="38">266.6</td>
<td id="55595DE2FFE60055FBB3FE03FB21FE08" box="[1070,1275,438,460]" gridcol="5" gridrow="2" pageId="37" pageNumber="38">995</td>
<td id="55595DE2FFE60055FABFFE03FA67FE08" box="[1314,1469,438,460]" gridcol="6" gridrow="2" pageId="37" pageNumber="38">88</td>
</tr>
<tr id="1688349EFFE60055FFE9FE55FA67FE32" box="[116,1469,480,502]" gridrow="3" pageId="37" pageNumber="38">
<th id="55595DE2FFE60055FFE9FE55FEC8FE32" box="[116,274,480,502]" gridcol="0" gridrow="3" pageId="37" pageNumber="38">BMRP 2002.4.4</th>
<td id="55595DE2FFE60055FEA5FE55FE07FE32" box="[312,477,480,502]" gridcol="1" gridrow="3" pageId="37" pageNumber="38">Jane (corrected)</td>
<td id="55595DE2FFE60055FD96FE55FD43FE32" box="[523,665,480,502]" gridcol="2" gridrow="3" pageId="37" pageNumber="38">
<taxonomicName id="6FB84D5FFFE6FF8FFD96FE54FD43FE32" box="[523,665,481,502]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE6FF8FFD96FE54FD43FE32" box="[523,665,481,502]" italics="true" pageId="37" pageNumber="38">Nanotyrannus</emphasis>
</taxonomicName>
</td>
<td id="55595DE2FFE60055FD57FE55FC98FE32" box="[714,834,480,502]" gridcol="3" gridrow="3" pageId="37" pageNumber="38">Periosteum</td>
<td id="55595DE2FFE60055FCF7FE55FBD1FE32" box="[874,1035,480,502]" gridcol="4" gridrow="3" pageId="37" pageNumber="38">273.2</td>
<td id="55595DE2FFE60055FBB3FE55FB21FE32" box="[1070,1275,480,502]" gridcol="5" gridrow="3" pageId="37" pageNumber="38">1064</td>
<td id="55595DE2FFE60055FABFFE55FA67FE32" box="[1314,1469,480,502]" gridcol="6" gridrow="3" pageId="37" pageNumber="38">69</td>
</tr>
</table>
</paragraph>
<paragraph id="A80736DCFFE6FF8FFE69FD9AFD08FC96" blockId="37.[441,1496,559,1018]" pageId="37" pageNumber="38">
Juvenile tyrannosaurs have high maximal growth rates, approaching [
<bibRefCitation id="CC294B2DFFE6FF8FFAA2FD9AFA96FD83" author="Erickson, G. M. &amp; Makovicky, P. J. &amp; Currie, P. J. &amp; Norell, M. A. &amp; Yerby, S. A. &amp; Brochu, C. A." box="[1343,1356,559,583]" journalOrPublisher="Nature" pageId="37" pageNumber="38" pagination="772 - 775" part="430" refId="ref27355" refString="8. Erickson, G. M.; Makovicky, P. J.; Currie, P. J.; Norell, M. A.; Yerby, S. A.; Brochu, C. A. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 2004, 430, 772 - 775. [CrossRef]" title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">8</bibRefCitation>
] or exceeding
<quantity id="6F409B39FFE6FF8FFE74FDE4FDE0FDAC" box="[489,570,592,617]" metricMagnitude="2" metricUnit="kg" metricValue="8.0" pageId="37" pageNumber="38" unit="kg" value="800.0">800 kg</quantity>
/y in
<taxonomicName id="6FB84D5FFFE6FF8FFDE7FDE4FCFAFDAC" authorityName=", Sue FMNH PR" authorityYear="2081" box="[634,800,593,616]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE6FF8FFDE7FDE4FCFAFDAC" box="[634,800,593,616]" italics="true" pageId="37" pageNumber="38">Tyrannosaurus</emphasis>
</taxonomicName>
. Immature
<emphasis id="9ACCEACEFFE6FF8FFC31FDE4FB82FDAC" box="[940,1112,593,616]" italics="true" pageId="37" pageNumber="38">
<taxonomicName id="6FB84D5FFFE6FF8FFC31FDE4FB8EFDAC" authorityName=", Sue FMNH PR" authorityYear="2081" box="[940,1108,593,616]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
,
</emphasis>
particularly juveniles weighing ~
<quantity id="6F409B39FFE6FF8FFE5BFDC7FDBFFD4E" box="[454,613,626,650]" metricMagnitude="3" metricUnit="kg" metricValue="1.5" metricValueMax="2.0" metricValueMin="1.0" pageId="37" pageNumber="38" unit="kg" value="1500.0" valueMax="2000.0" valueMin="1000.0">10002000 kg</quantity>
, are predicted to have high growth rates as they enter their exponential growth phase. If
<taxonomicName id="6FB84D5FFFE6FF8FFD16FD21FC9DFD6F" box="[651,839,659,683]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="37" pageNumber="46" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE6FF8FFD16FD21FCF6FD6F" box="[651,812,660,683]" italics="true" pageId="37" pageNumber="38">Nanotyrannus</emphasis>
is
</taxonomicName>
a distinct, small-bodied tyrannosaur, then it will have much lower growth rates at this size, comparable to growth curves modeled for small-bodied tyrannosaurids such as
<taxonomicName id="6FB84D5FFFE6FF8FFCB3FD62FC60FD2A" authorityName="Lambe" authorityYear="1914" box="[814,954,727,750]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE6FF8FFCB3FD62FC60FD2A" box="[814,954,727,750]" italics="true" pageId="37" pageNumber="38">Gorgosaurus</emphasis>
</taxonomicName>
or
<taxonomicName id="6FB84D5FFFE6FF8FFC7CFD63FBA1FD2A" authorityName="Osborn" authorityYear="1905" box="[993,1147,726,750]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE6FF8FFC7CFD63FBA1FD2A" box="[993,1147,726,750]" italics="true" pageId="37" pageNumber="38">Albertosaurus</emphasis>
</taxonomicName>
[
<bibRefCitation id="CC294B2DFFE6FF8FFB17FD62FB4DFD2B" author="Erickson, G. M. &amp; Makovicky, P. J. &amp; Currie, P. J. &amp; Norell, M. A. &amp; Yerby, S. A. &amp; Brochu, C. A." box="[1162,1175,727,751]" journalOrPublisher="Nature" pageId="37" pageNumber="38" pagination="772 - 775" part="430" refId="ref27355" refString="8. Erickson, G. M.; Makovicky, P. J.; Currie, P. J.; Norell, M. A.; Yerby, S. A.; Brochu, C. A. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 2004, 430, 772 - 775. [CrossRef]" title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">8</bibRefCitation>
]. However, if the two taxa are synonymous, then specimens the size of
<collectionCode id="CEA9AE19FFE6FF8FFC5EFD4DFBD4FCCB" box="[963,1038,760,783]" collectionName="BMRP" pageId="37" pageNumber="38">BMRP</collectionCode>
<date id="DC06101CFFE6FF8FFB88FD4DFBABFCD4" box="[1045,1137,760,784]" pageId="37" pageNumber="38" value="2002-04-01">2002.4.1</date>
and
<collectionCode id="CEA9AE19FFE6FF8FFB36FD4DFB2CFCCB" box="[1195,1270,760,783]" collectionName="BMRP" pageId="37" pageNumber="38">BMRP</collectionCode>
<date id="DC06101CFFE6FF8FFB60FD4DFA80FCD4" box="[1277,1370,760,784]" pageId="37" pageNumber="38" value="2006-04-04">2006.4.4</date>
should be in their rapid, exponential growth phase—especially if one assumes that they have entered their teenage years [
<bibRefCitation id="CC294B2DFFE6FF8FFD35FC8EFD19FC97" author="Woodward, H. N. &amp; Tremaine, K. &amp; Williams, S. A &amp; Zanno, L. E. &amp; Horner, J. R. &amp; Myhrvold, N." box="[680,707,827,851]" journalOrPublisher="Sci. Adv." pageId="37" pageNumber="38" pagination="eaax 6250" part="6" refId="ref29210" refString="46. Woodward, H. N.; Tremaine, K.; Williams, S. A.; Zanno, L. E.; Horner, J. R.; Myhrvold, N. Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Sci. Adv. 2020, 6, eaax 6250. [CrossRef]" title="Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">46</bibRefCitation>
].
</paragraph>
<paragraph id="A80736DCFFE6FF8FFE69FCE8FA6DFC3E" blockId="37.[441,1496,559,1018]" pageId="37" pageNumber="38">
Narrow spacing of LAGs (
<figureCitation id="30832A59FFE6FF8FFCB1FCE9FC46FCB0" box="[812,924,860,884]" captionStart="Figure 24" captionStartId="35.[444,513,940,962]" captionTargetId="figure-210@35.[157,1431,-431,904]" captionTargetPageId="35" captionText="Figure 24. Thin sections of the femur of (A), BMRP 2002.4.1 (“Jane”) and (B), BMRP 2006.4.4 (“Petey”). An external fundamental system (EFS) is absent, but lines of arrested growth or LAGs (blue) become closer toward the outer edge of the bone, showing decelerating growth. Modified from [46]. Note that some LAGs are likely split (multi-LAGs) and do not represent a full year of growth." figureDoi="http://doi.org/10.5281/zenodo.10535319" httpUri="https://zenodo.org/record/10535319/files/figure.png" pageId="37" pageNumber="38">Figure 24</figureCitation>
), especially towards the periosteum, shows low growth rates in putative
<taxonomicName id="6FB84D5FFFE6FF8FFD75FCCAFC53FC52" box="[744,905,895,918]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="37" pageNumber="38" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE6FF8FFD75FCCAFC53FC52" box="[744,905,895,918]" italics="true" pageId="37" pageNumber="38">Nanotyrannus</emphasis>
</taxonomicName>
specimens (
<figureCitation id="30832A59FFE6FF8FFB80FCCBFB54FC52" box="[1053,1166,894,918]" captionStart="Figure 25" captionStartId="37.[444,516,1769,1791]" captionTargetBox="[124,1466,1061,1723]" captionTargetId="figure-758@37.[103,1485,1036,1735]" captionTargetPageId="37" captionText="Figure 25. Changes in estimated body mass from previous LAG as preserved in the femurs of Sue (red, circles), Petey (green, triangles), and Jane BMRP 2002.4.1, corrected for possible split multi-LAGs as in Cullen et al. [49] (dark blue, solid squares) and uncorrected as in Woodward et al. [46] (light blue, open squares), at the time of death. Periosteum included here as the final half LAG. LAG 1 excluded here because (1) it is a partial record of growth due to medullary cavity remodeling and bone resorption and (2) because Janes incomplete femur and Sues core sampling (rather than a complete transverse section) exclude easy measurement/estimation of endosteal circumference. An EFS is readily apparent in the last decade or so of Sues life. See the Supplemental Material for plots that exclude the periosteum." figureDoi="http://doi.org/10.5281/zenodo.10535321" httpUri="https://zenodo.org/record/10535321/files/figure.png" pageId="37" pageNumber="38">Figure 25</figureCitation>
,
<tableCitation id="E53A0367FFE6FF8FFB00FCCBFB29FC52" box="[1181,1267,894,918]" captionStart="Table 5" captionStartId="35.[443,502,1103,1124]" captionTargetPageId="35" captionText="Table 5. LAG data, body mass, and growth rate estimates rounded to the nearest kg for Sue (FMNH PR 2081), Petey (BMRP 2006.4.4), and Jane (BMRP 2002.4.4) based on femoral circumferences, including corrected values for Jane based on a split multi-LAG interpretation [49]. Campione et al. [50] estimate Sues mass at death (i.e., after LAG #23) as 7377 kg after applying a correction for non-circular femoral cross-section. Hutchinson et al. [104] estimate Janes mass at death (i.e., after LAG #9) as 954 kg based on 3D modeling. See Supplemental File for equations used in calculations. Note that LAG #1 and the periosteum do not record a full year of growth. Periosteum is assigned a LAG pseudo-number of +0.5 after the final LAG, indicating that death could have occurred at any point after the last yearly marker was deposited while assigning an age at the midpoint of that final year. Jane required estimation of LAG/periosteum circumferences from LAG spacing and femoral width due to the incomplete cross-section of the femur midshaft. (1) LAG (or periosteum) distance from endosteum is measured at the time of death, (2) femoral width at midshaft (including medullary cavity) is measured at time of death and back estimated using LAG spacing, (3) femoral circumference derived from femoral width at a given year is approximated as a circular cross-section. Data sources: Campione et al. [50]; Cullen et al. [49]; Woodward et al. [46]." httpUri="http://table.plazi.org/id/FCC76654FFE0FF89FE26FBFAFC83F987" pageId="37" pageNumber="38" tableUuid="FCC76654FFE0FF89FE26FBFAFC83F987">Table 5</tableCitation>
). Growth rates do not exceed
<quantity id="6F409B39FFE6FF8FFDDDFC2AFD55FC73" box="[576,655,927,951]" metricMagnitude="2" metricUnit="kg" metricValue="1.5" pageId="37" pageNumber="38" unit="kg" value="150.0">150 kg</quantity>
/year for the last few years of life and can be less than
<quantity id="6F409B39FFE6FF8FFA8CFC2AFA88FC73" box="[1297,1362,927,951]" metricMagnitude="1" metricUnit="kg" metricValue="5.0" pageId="37" pageNumber="38" unit="kg" value="50.0">50 kg</quantity>
/year. This rejects the hypothesis that these are young, rapidly growing
<taxonomicName id="6FB84D5FFFE6FF8FFB16FC74FB11FC1C" box="[1163,1227,961,984]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="37" pageNumber="38" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFE6FF8FFB16FC74FB46FC1C" box="[1163,1180,961,984]" italics="true" pageId="37" pageNumber="38">T</emphasis>
.
<emphasis id="9ACCEACEFFE6FF8FFB34FC74FB11FC1C" box="[1193,1227,961,984]" italics="true" pageId="37" pageNumber="38">rex</emphasis>
</taxonomicName>
, which achieved peak growth rates exceeding
<quantity id="6F409B39FFE6FF8FFD49FC57FCFEFC3E" box="[724,804,994,1018]" metricMagnitude="2" metricUnit="kg" metricValue="8.0" pageId="37" pageNumber="38" unit="kg" value="800.0">800 kg</quantity>
/y based on the estimates from
<collectionCode id="CEA9AE19FFE6FF8FFB09FC56FB32FC3E" box="[1172,1256,995,1018]" collectionName="USA, Illinois, Chicago, Field Museum of Natural History (also used by Finnish Museum of Natural History)" pageId="37" pageNumber="38">FMNH</collectionCode>
PR2081 (
<tableCitation id="E53A0367FFE6FF8FFAC9FC57FA7DFC3E" box="[1364,1447,994,1018]" captionStart="Table 5" captionStartId="35.[443,502,1103,1124]" captionTargetPageId="35" captionText="Table 5. LAG data, body mass, and growth rate estimates rounded to the nearest kg for Sue (FMNH PR 2081), Petey (BMRP 2006.4.4), and Jane (BMRP 2002.4.4) based on femoral circumferences, including corrected values for Jane based on a split multi-LAG interpretation [49]. Campione et al. [50] estimate Sues mass at death (i.e., after LAG #23) as 7377 kg after applying a correction for non-circular femoral cross-section. Hutchinson et al. [104] estimate Janes mass at death (i.e., after LAG #9) as 954 kg based on 3D modeling. See Supplemental File for equations used in calculations. Note that LAG #1 and the periosteum do not record a full year of growth. Periosteum is assigned a LAG pseudo-number of +0.5 after the final LAG, indicating that death could have occurred at any point after the last yearly marker was deposited while assigning an age at the midpoint of that final year. Jane required estimation of LAG/periosteum circumferences from LAG spacing and femoral width due to the incomplete cross-section of the femur midshaft. (1) LAG (or periosteum) distance from endosteum is measured at the time of death, (2) femoral width at midshaft (including medullary cavity) is measured at time of death and back estimated using LAG spacing, (3) femoral circumference derived from femoral width at a given year is approximated as a circular cross-section. Data sources: Campione et al. [50]; Cullen et al. [49]; Woodward et al. [46]." httpUri="http://table.plazi.org/id/FCC76654FFE0FF89FE26FBFAFC83F987" pageId="37" pageNumber="38" tableUuid="FCC76654FFE0FF89FE26FBFAFC83F987">Table 5</tableCitation>
).
</paragraph>
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<paragraph id="A80736DCFFE6FF8FFE21F95CFD33F7D5" blockId="37.[443,1492,1769,2065]" pageId="37" pageNumber="38">
<emphasis id="9ACCEACEFFE6FF8FFE21F95CFDF2F93A" bold="true" box="[444,552,1769,1791]" pageId="37" pageNumber="38">Figure 25.</emphasis>
Changes in estimated body mass from previous LAG as preserved in the femurs of Sue (red, circles), Petey (green, triangles), and Jane BMRP 2002.4.1, corrected for possible split multi-LAGs as in Cullen et al. [
<bibRefCitation id="CC294B2DFFE6FF8FFD18F89BFD47F887" author="Cullen, T. M. &amp; Canale, J. I &amp; Apesteguia, S. &amp; Smith, N. D. &amp; Hu, D. &amp; Makovicky, P. J." box="[645,669,1838,1859]" journalOrPublisher="Proc. R. Soc. B" pageId="37" pageNumber="38" pagination="20202258" part="287" refId="ref29372" refString="49 Cullen, T. M.; Canale, J. I.; Apesteguia, S.; Smith, N. D.; Hu, D.; Makovicky, P. J. Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution. Proc. R. Soc. B 2020, 287, 20202258. [CrossRef]" title="Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution." type="journal article" year="2020">49</bibRefCitation>
] (dark blue, solid squares) and uncorrected as in Woodward et al. [
<bibRefCitation id="CC294B2DFFE6FF8FFAEEF89BFA51F880" author="Woodward, H. N. &amp; Tremaine, K. &amp; Williams, S. A &amp; Zanno, L. E. &amp; Horner, J. R. &amp; Myhrvold, N." box="[1395,1419,1838,1860]" journalOrPublisher="Sci. Adv." pageId="37" pageNumber="38" pagination="eaax 6250" part="6" refId="ref29210" refString="46. Woodward, H. N.; Tremaine, K.; Williams, S. A.; Zanno, L. E.; Horner, J. R.; Myhrvold, N. Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Sci. Adv. 2020, 6, eaax 6250. [CrossRef]" title="Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">46</bibRefCitation>
] (light blue, open squares), at the time of death. Periosteum included here as the final half LAG. LAG 1 excluded here because (1) it is a partial record of growth due to medullary cavity remodeling and bone resorption and (2) because Janes incomplete femur and Sues core sampling (rather than a complete transverse section) exclude easy measurement/estimation of endosteal circumference. An EFS is readily apparent in the last decade or so of Sues life. See the Supplemental Material for plots that exclude the periosteum.
</paragraph>
</caption>
<paragraph id="A80736DCFFE5FF8CFE21FF45FD6BFECC" blockId="38.[443,1496,240,1778]" box="[444,689,240,264]" pageId="38" pageNumber="39">3.4.6. Growth Trends</paragraph>
<paragraph id="A80736DCFFE5FF8CFE69FEAFFADDFDF9" blockId="38.[443,1496,240,1778]" pageId="38" pageNumber="39">
Growth rates change over time. Growth accelerates until roughly the middle of life in symmetric logistic and asymmetric Gompertz growth models as commonly applied to dinosaurs [
<bibRefCitation id="CC294B2DFFE5FF8CFDFFFEE8FDB5FEB1" author="Erickson, G. M. &amp; Makovicky, P. J. &amp; Currie, P. J. &amp; Norell, M. A. &amp; Yerby, S. A. &amp; Brochu, C. A." box="[610,623,349,373]" journalOrPublisher="Nature" pageId="38" pageNumber="39" pagination="772 - 775" part="430" refId="ref27355" refString="8. Erickson, G. M.; Makovicky, P. J.; Currie, P. J.; Norell, M. A.; Yerby, S. A.; Brochu, C. A. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 2004, 430, 772 - 775. [CrossRef]" title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">8</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE5FF8CFDEBFEE8FD44FEB1" author="Erickson, G. M. &amp; Tumanova, T. A." box="[630,670,349,373]" journalOrPublisher="Zool. J. Linn. Soc." pageId="38" pageNumber="39" pagination="551 - 566" part="130" refId="ref34073" refString="131. Erickson, G. M.; Tumanova, T. A. Growth curve of Psittacosaurus mongoliensis Osborn (Ceratopsia: Psittacosauridae) inferred from long bone histology. Zool. J. Linn. Soc. 2000, 130, 551 - 566. [CrossRef]" title="Growth curve of Psittacosaurus mongoliensis Osborn (Ceratopsia: Psittacosauridae) inferred from long bone histology" type="journal article" year="2000">131</bibRefCitation>
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] then decelerates. Growth finally slows and almost stops in the last few years of life, reaching asymptotic growth. Based on their size (and even their approximated age from prior studies [
<bibRefCitation id="CC294B2DFFE5FF8CFC11FE2AFC7DFE73" author="Woodward, H. N. &amp; Tremaine, K. &amp; Williams, S. A &amp; Zanno, L. E. &amp; Horner, J. R. &amp; Myhrvold, N." box="[908,935,415,439]" journalOrPublisher="Sci. Adv." pageId="38" pageNumber="39" pagination="eaax 6250" part="6" refId="ref29210" refString="46. Woodward, H. N.; Tremaine, K.; Williams, S. A.; Zanno, L. E.; Horner, J. R.; Myhrvold, N. Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Sci. Adv. 2020, 6, eaax 6250. [CrossRef]" title="Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">46</bibRefCitation>
]), if the animals assigned to
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<emphasis id="9ACCEACEFFE5FF8CFA98FE15FA7CFE73" box="[1285,1446,416,439]" italics="true" pageId="38" pageNumber="39">Nanotyrannus</emphasis>
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are juvenile
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<emphasis id="9ACCEACEFFE5FF8CFDBFFE77FDE9FE1D" box="[546,563,450,473]" italics="true" pageId="38" pageNumber="39">T</emphasis>
.
<emphasis id="9ACCEACEFFE5FF8CFDDDFE77FDB8FE1D" box="[576,610,450,473]" italics="true" pageId="38" pageNumber="39">rex</emphasis>
</taxonomicName>
, then they should show increasing growth rates (i.e., exponential growth) as they approach the rapid, protracted growth spurt at the middle of the tyrannosaur life cycle at around
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[
<bibRefCitation id="CC294B2DFFE5FF8CFC83FDB1FCF1FDD8" author="Erickson, G. M. &amp; Makovicky, P. J. &amp; Currie, P. J. &amp; Norell, M. A. &amp; Yerby, S. A. &amp; Brochu, C. A." box="[798,811,516,540]" journalOrPublisher="Nature" pageId="38" pageNumber="39" pagination="772 - 775" part="430" refId="ref27355" refString="8. Erickson, G. M.; Makovicky, P. J.; Currie, P. J.; Norell, M. A.; Yerby, S. A.; Brochu, C. A. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 2004, 430, 772 - 775. [CrossRef]" title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">8</bibRefCitation>
]. If they are subadults or adults of a distinct, small-bodied species, they would be expected to show decelerating or ceased growth.
</paragraph>
<paragraph id="A80736DCFFE5FF8CFE69FDF2FAA6FD07" blockId="38.[443,1496,240,1778]" pageId="38" pageNumber="39">
Growth rates in
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<emphasis id="9ACCEACEFFE5FF8CFD2CFDFDFC97FD9B" box="[689,845,584,607]" italics="true" pageId="38" pageNumber="39">Nanotyrannus</emphasis>
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show a general trend of deceleration in their final years of life. These trends resemble those seen in mature
<taxonomicName id="6FB84D5FFFE5FF8CFB8BFDDCFB8CFD44" box="[1046,1110,617,640]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="38" pageNumber="39" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFE5FF8CFB8BFDDCFBFCFD44" box="[1046,1062,617,640]" italics="true" pageId="38" pageNumber="39">T</emphasis>
.
<emphasis id="9ACCEACEFFE5FF8CFBA9FDDCFB8CFD44" box="[1076,1110,617,640]" italics="true" pageId="38" pageNumber="39">rex</emphasis>
</taxonomicName>
as growth begins to plateau just before it establishes an EFS. These patterns are strongly suggestive of relatively mature animals, either late-stage subadults or early adults, not rapidly growing juveniles.
</paragraph>
<paragraph id="A80736DCFFE5FF8CFE69FD78FB24FC6A" blockId="38.[443,1496,240,1778]" pageId="38" pageNumber="39">
<emphasis id="9ACCEACEFFE5FF8CFE69FD78FD26FD20" box="[500,764,717,741]" italics="true" pageId="38" pageNumber="39">Estimated maximum size</emphasis>
. It is possible to fit various kinds of growth curves to estimated masses [
<bibRefCitation id="CC294B2DFFE5FF8CFD86FD5BFD99FCC2" author="Erickson, G. M." box="[539,579,750,774]" journalOrPublisher="Annu. Rev. Earth Planet. Sci." pageId="38" pageNumber="39" pagination="675 - 697" part="42" refId="ref34183" refString="133. Erickson, G. M. On dinosaur growth. Annu. Rev. Earth Planet. Sci. 2014, 42, 675 - 697. [CrossRef]" title="On dinosaur growth" type="journal article" year="2014">133</bibRefCitation>
], which can be extrapolated and used to predict the mass that a given individual would have achieved at full size. This approach can be used to test whether
<taxonomicName id="6FB84D5FFFE5FF8CFAABFCA5FA09FCE3" box="[1334,1491,784,807]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE5FF8CFAABFCA5FA09FCE3" box="[1334,1491,784,807]" italics="true" pageId="38" pageNumber="39">Nanotyrannus</emphasis>
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specimens would have grown to the enormous sizes (~
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) seen in
<taxonomicName id="6FB84D5FFFE5FF8CFB7AFC87FAFEFC8D" box="[1255,1316,818,841]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="38" pageNumber="39" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFE5FF8CFB7AFC87FB2DFC8D" box="[1255,1271,818,841]" italics="true" pageId="38" pageNumber="39">T</emphasis>
.
<emphasis id="9ACCEACEFFE5FF8CFA9EFC87FAFEFC8D" box="[1283,1316,818,841]" italics="true" pageId="38" pageNumber="39">rex</emphasis>
</taxonomicName>
. If the putative
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<emphasis id="9ACCEACEFFE5FF8CFE21FCE6FD86FCAE" box="[444,604,851,874]" italics="true" pageId="38" pageNumber="39">Nanotyrannus</emphasis>
</taxonomicName>
were juveniles of
<taxonomicName id="6FB84D5FFFE5FF8CFCAFFCE6FCB5FCAE" box="[818,879,851,874]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="38" pageNumber="39" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFE5FF8CFCAFFCE6FCB5FCAE" box="[818,879,851,874]" italics="true" pageId="38" pageNumber="39">T. rex</emphasis>
</taxonomicName>
, then their predicted adult masses should be on the order of
<quantity id="6F409B39FFE5FF8CFDBDFCC1FD0AFC48" box="[544,720,884,908]" metricMagnitude="3" metricUnit="kg" metricValue="7.5" metricValueMax="10.0" metricValueMin="5.0" pageId="38" pageNumber="39" unit="kg" value="7500.0" valueMax="10000.0" valueMin="5000.0">500010,000 kg</quantity>
, as in
<taxonomicName id="6FB84D5FFFE5FF8CFC84FCC0FC8FFC48" box="[793,853,885,908]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="38" pageNumber="39" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFE5FF8CFC84FCC0FC8FFC48" box="[793,853,885,908]" italics="true" pageId="38" pageNumber="39">T. rex</emphasis>
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. If they are subadults or young adults of small-bodied tyrannosaurs, then their predicted adult masses should be much lower.
</paragraph>
<paragraph id="A80736DCFFE5FF8CFE69FC02FCF6FAA5" blockId="38.[443,1496,240,1778]" pageId="38" pageNumber="39">
A caveat is that, when fitting a growth model to a year-by-year growth record from a single individual rather than to mass-age data from separate individuals, the assumption of independence of data is violated, making these pseudo-regression analyses (and making the calculation of confidence/prediction intervals moot). These models are nevertheless useful in extrapolating adult masses when individuals die prior to reaching full size. This is because, although extrapolation is always highly uncertain in science, we are limited to the growth record preserved in the femora; speculation that growth rates could have exponentially increased had these putative
<taxonomicName id="6FB84D5FFFE5FF8CFB8CFB17FB68FB7D" box="[1041,1202,1186,1209]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="38" pageNumber="39" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE5FF8CFB8CFB17FB68FB7D" box="[1041,1202,1186,1209]" italics="true" pageId="38" pageNumber="39">Nanotyrannus</emphasis>
</taxonomicName>
specimens lived longer is, therefore, a weaker argument than the use of adult size estimates from these pseudoregressions. We prefer the admittedly high uncertainty of extrapolation modeled on empirical evidence to speculation (i.e., one could speculate that any number of changes in growth rate or morphology might have occurred post-mortem since such speculation is unbounded by fossil evidence).
</paragraph>
<paragraph id="A80736DCFFE5FF8CFE69FADFFDD5F936" blockId="38.[443,1496,240,1778]" pageId="38" pageNumber="39">
Growth curves using asymptotic logistic, Gompertz, and von Bertalanffy models predict fully adult masses (
<figureCitation id="30832A59FFE5FF8CFC98FA39FCADFA60" box="[773,887,1420,1444]" captionStart="Figure 26" captionStartId="39.[444,514,1756,1778]" captionTargetBox="[172,1419,247,1707]" captionTargetId="figure-145@39.[167,1505,225,1750]" captionTargetPageId="39" captionText="Figure 26. Age-independent growth curves for Nanotyrannus BMRP 2006.4.4 Petey (green, triangles) and Jane, the latter both corrected (dark blue, solid squares) for split multi-LAGs as in Cullen et al. [49] and uncorrected (light blue, open squares) as in Woodward et al. [46]. LAG #1 and the periosteum (arbitrarily assigned a half-year value) are included in the regressions. Four models are fit to each specimen: logistic (solid), Gompertz (dashed), logarithmic (dotted), and von Bertalanffy (dotdash). Horizontal line is the mass estimate at the time of death for Jane from Hutchinson et al. [104] using 3D modeling. Results are similar to those deriving from regressions that exclude LAG #1 and the periosteum (Supplemental Material)." figureDoi="http://doi.org/10.5281/zenodo.10535323" httpUri="https://zenodo.org/record/10535323/files/figure.png" pageId="38" pageNumber="39">Figure 26</figureCitation>
) on the order of perhaps ~
<quantity id="6F409B39FFE5FF8CFB23FA39FA89FA60" box="[1214,1363,1420,1444]" metricMagnitude="2" metricUnit="kg" metricValue="9.0" metricValueMax="11.0" metricValueMin="7.0" pageId="38" pageNumber="39" unit="kg" value="900.0" valueMax="1100.0" valueMin="700.0">7001100 kg</quantity>
for
<collectionCode id="CEA9AE19FFE5FF8CFA1AFA39FA0EFA67" box="[1415,1492,1420,1443]" collectionName="BMRP" pageId="38" pageNumber="39">BMRP</collectionCode>
<date id="DC06101CFFE5FF8CFE21FA18FDC2FA01" box="[444,536,1453,1477]" pageId="38" pageNumber="39" value="2006-04-04">2006.4.4</date>
and ~
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for
<collectionCode id="CEA9AE19FFE5FF8CFCB0FA1BFCA2FA01" box="[813,888,1454,1477]" collectionName="BMRP" pageId="38" pageNumber="39">BMRP</collectionCode>
<date id="DC06101CFFE5FF8CFCE2FA18FC06FA01" box="[895,988,1453,1477]" pageId="38" pageNumber="39" value="2002-04-01">2002.4.1</date>
(when corrected for split multi-LAGs [
<bibRefCitation id="CC294B2DFFE5FF8CFA38FA18FA1AFA01" author="Cullen, T. M. &amp; Canale, J. I &amp; Apesteguia, S. &amp; Smith, N. D. &amp; Hu, D. &amp; Makovicky, P. J." box="[1445,1472,1453,1477]" journalOrPublisher="Proc. R. Soc. B" pageId="38" pageNumber="39" pagination="20202258" part="287" refId="ref29372" refString="49 Cullen, T. M.; Canale, J. I.; Apesteguia, S.; Smith, N. D.; Hu, D.; Makovicky, P. J. Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution. Proc. R. Soc. B 2020, 287, 20202258. [CrossRef]" title="Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution." type="journal article" year="2020">49</bibRefCitation>
]). Non-asymptotic logarithmic models can achieve higher masses since they have no upper limit, but these predicted masses still fall far short of
<taxonomicName id="6FB84D5FFFE5FF8CFBA0FA44FBA1F9CC" box="[1085,1147,1521,1544]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="38" pageNumber="39" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFE5FF8CFBA0FA44FBA1F9CC" box="[1085,1147,1521,1544]" italics="true" pageId="38" pageNumber="39">T. rex</emphasis>
</taxonomicName>
(
<figureCitation id="30832A59FFE5FF8CFB16FA45FB26F9CC" box="[1163,1276,1520,1544]" captionStart="Figure 27" captionStartId="40.[444,515,1665,1687]" captionTargetBox="[213,1374,254,1612]" captionTargetId="figure-181@40.[209,1414,220,1633]" captionTargetPageId="40" captionText="Figure 27. Age-independent growth curves for a large, old Tyrannosaurus, Sue FMNH PR 2081 (red, circles), and two Nanotyrannus, Petey BMRP 2006.4.4 (green, triangles) and Jane BMRP 2002.4.1 (dark blue, squares), corrected for split multi-LAGs as in Cullen et al. [49]. LAG #1 and the periosteum are not included in the regressions shown here. Four different growth models are fit to each specimen: logistic (solid), Gompertz (dashed), logarithmic (dotted), and von Bertalanffy (dot—dash). Horizontal black lines are asymptotic masses derived from logistic regressions of multiple individuals (i.e., multiple specimens used in the regression, with each specimen assigned a single mass and age at death) from Longrich et al. (in review) and are presented in decreasing order as follows: Tyrannosaurus, Gorgosaurus, Albertosaurus. Results are similar to those deriving from regressions that include LAG #1 and the periosteum (Supplemental Material)." figureDoi="http://doi.org/10.5281/zenodo.10535327" httpUri="https://zenodo.org/record/10535327/files/figure.png" pageId="38" pageNumber="39">Figure 27</figureCitation>
) and are closer to that of
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<emphasis id="9ACCEACEFFE5FF8CFD8FF9A7FD74F9EE" box="[530,686,1554,1578]" italics="true" pageId="38" pageNumber="39">Albertosaurus</emphasis>
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. These estimates are, in the context of comparison with
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<emphasis id="9ACCEACEFFE5FF8CFACBF9A7FA4FF9ED" box="[1366,1429,1554,1577]" italics="true" pageId="38" pageNumber="39">T. rex</emphasis>
</taxonomicName>
, also roughly consistent with mass estimates at the time of death for
<collectionCode id="CEA9AE19FFE5FF8CFB11F981FB0CF98F" box="[1164,1238,1588,1611]" collectionName="BMRP" pageId="38" pageNumber="39">BMRP</collectionCode>
<date id="DC06101CFFE5FF8CFB46F986FAECF98F" box="[1243,1334,1587,1611]" pageId="38" pageNumber="39" value="2002-04-01">2002.4.1</date>
derived from 3D modeling [
<bibRefCitation id="CC294B2DFFE5FF8CFDFAF9E0FD55F9A9" author="Hutchinson, J. R. &amp; Bates, K. T. &amp; Molnar, J. &amp; Allen, V. &amp; Makovicky, P. J." box="[615,655,1621,1645]" journalOrPublisher="PLoS ONE" pageId="38" pageNumber="39" pagination="e 26037" part="6" refId="ref32548" refString="104. Hutchinson, J. R.; Bates, K. T.; Molnar, J.; Allen, V.; Makovicky, P. J. A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth. PLoS ONE 2011, 6, e 26037. [CrossRef]" title="A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth" type="journal article" year="2011">104</bibRefCitation>
]. Mass estimates are not available for the Zuri specimen (
<collectionCode id="CEA9AE19FFE5FF8CFAA0F9E0FAAFF9A9" box="[1341,1397,1621,1645]" collectionName="HRS" pageId="38" pageNumber="39">HRS</collectionCode>
081514) because the pubis was sectioned rather than the femur. However, plotting the growth of this specimen using data from Griffin [
<bibRefCitation id="CC294B2DFFE5FF8CFC12F922FC70F96B" author="Griffin, C." bookContentInfo="Cedarville, OH, USA," box="[911,938,1687,1711]" pageId="38" pageNumber="39" refId="ref29498" refString="51. Griffin, C. Using osteohistology to determine the taxonomic validity of the Late Cretaceous dinosaur Nanotyrannus lancensis Bakker et al, 1988 (Theropoda: Tyrannosauridae). In Proceedings of the The Research and Scholarship Symposium, Cedarville, OH, USA, 16 April 2014." title="Using osteohistology to determine the taxonomic validity of the Late Cretaceous dinosaur Nanotyrannus lancensis Bakker et al, 1988 (Theropoda: Tyrannosauridae)" type="proceedings paper" volumeTitle="Proceedings of the The Research and Scholarship Symposium" year="2014">51</bibRefCitation>
] shows slow growth and growth deceleration rather than rapid, accelerating growth (
<figureCitation id="30832A59FFE5FF8CFC0FF90CFBD8F915" box="[914,1026,1721,1745]" captionStart="Figure 28" captionStartId="41.[444,516,1409,1431]" captionTargetBox="[133,1460,248,1363]" captionTargetId="figure-295@41.[116,1484,218,1373]" captionTargetPageId="41" captionText="Figure 28. Growth record from Zuri (HRS 081514) pubis, with radius back-estimated from LAG spacing [51]. Models including the periosteum and LAG 1 are in purple; excluding the periosteum and LAG 1 are in orange. Four different growth models are fit under both conditions: logistic (solid), Gompertz (dashed), logarithmic (dotted), and von Bertalanffy (dotdash)." figureDoi="http://doi.org/10.5281/zenodo.10535329" httpUri="https://zenodo.org/record/10535329/files/figure.png" pageId="38" pageNumber="39">Figure 28</figureCitation>
); it was apparently near full size when it died.
</paragraph>
<caption id="FCC76654FFE4FF8DFE21F969FC98F825" ID-DOI="http://doi.org/10.5281/zenodo.10535323" ID-Zenodo-Dep="10535323" httpUri="https://zenodo.org/record/10535323/files/figure.png" pageId="39" pageNumber="40" startId="39.[444,514,1756,1778]" targetBox="[172,1419,247,1707]" targetPageId="39" targetType="figure">
<paragraph id="A80736DCFFE4FF8DFE21F969FC98F825" blockId="39.[443,1496,1756,2017]" pageId="39" pageNumber="40">
<emphasis id="9ACCEACEFFE4FF8DFE21F969FDFFF935" bold="true" box="[444,549,1756,1778]" pageId="39" pageNumber="40">Figure 26.</emphasis>
Age-independent growth curves for
<taxonomicName id="6FB84D5FFFE4FF8DFC33F969FB5EF935" box="[942,1156,1756,1777]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE4FF8DFC33F969FBE1F935" box="[942,1083,1756,1777]" italics="true" pageId="39" pageNumber="40">Nanotyrannus</emphasis>
BMRP
</taxonomicName>
2006.4.4 Petey (green, triangles) and Jane, the latter both corrected (dark blue, solid squares) for split multi-LAGs as in Cullen et al. [
<bibRefCitation id="CC294B2DFFE4FF8DFA29F94BFA16F8D7" author="Cullen, T. M. &amp; Canale, J. I &amp; Apesteguia, S. &amp; Smith, N. D. &amp; Hu, D. &amp; Makovicky, P. J." box="[1460,1484,1790,1811]" journalOrPublisher="Proc. R. Soc. B" pageId="39" pageNumber="40" pagination="20202258" part="287" refId="ref29372" refString="49 Cullen, T. M.; Canale, J. I.; Apesteguia, S.; Smith, N. D.; Hu, D.; Makovicky, P. J. Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution. Proc. R. Soc. B 2020, 287, 20202258. [CrossRef]" title="Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution." type="journal article" year="2020">49</bibRefCitation>
] and uncorrected (light blue, open squares) as in Woodward et al. [
<bibRefCitation id="CC294B2DFFE4FF8DFBE0F895FB4FF8F2" author="Woodward, H. N. &amp; Tremaine, K. &amp; Williams, S. A &amp; Zanno, L. E. &amp; Horner, J. R. &amp; Myhrvold, N." box="[1149,1173,1824,1846]" journalOrPublisher="Sci. Adv." pageId="39" pageNumber="40" pagination="eaax 6250" part="6" refId="ref29210" refString="46. Woodward, H. N.; Tremaine, K.; Williams, S. A.; Zanno, L. E.; Horner, J. R.; Myhrvold, N. Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus. Sci. Adv. 2020, 6, eaax 6250. [CrossRef]" title="Growing up Tyrannosaurus rex: Osteohistology refutes the pygmy &quot; Nanotyrannus &quot; and supports ontogenetic niche partitioning in juvenile Tyrannosaurus" type="journal article" year="2020">46</bibRefCitation>
]. LAG #1 and the periosteum (arbitrarily assigned a half-year value) are included in the regressions. Four models are fit to each specimen: logistic (solid), Gompertz (dashed), logarithmic (dotted), and von Bertalanffy (dotdash). Horizontal line is the mass estimate at the time of death for Jane from Hutchinson et al. [
<bibRefCitation id="CC294B2DFFE4FF8DFAFBF832FA50F858" author="Hutchinson, J. R. &amp; Bates, K. T. &amp; Molnar, J. &amp; Allen, V. &amp; Makovicky, P. J." box="[1382,1418,1927,1948]" journalOrPublisher="PLoS ONE" pageId="39" pageNumber="40" pagination="e 26037" part="6" refId="ref32548" refString="104. Hutchinson, J. R.; Bates, K. T.; Molnar, J.; Allen, V.; Makovicky, P. J. A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth. PLoS ONE 2011, 6, e 26037. [CrossRef]" title="A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth" type="journal article" year="2011">104</bibRefCitation>
] using 3D modeling. Results are similar to those deriving from regressions that exclude LAG #1 and the periosteum (Supplemental Material).
</paragraph>
</caption>
<caption id="FCC76654FFEBFF82FE21F934FC43F80F" ID-DOI="http://doi.org/10.5281/zenodo.10535327" ID-Zenodo-Dep="10535327" httpUri="https://zenodo.org/record/10535327/files/figure.png" pageId="40" pageNumber="41" startId="40.[444,515,1665,1687]" targetBox="[213,1374,254,1612]" targetPageId="40" targetType="figure">
<paragraph id="A80736DCFFEBFF82FE21F934FC43F80F" blockId="40.[443,1495,1665,1995]" pageId="40" pageNumber="41">
<emphasis id="9ACCEACEFFEBFF82FE21F934FDFCF952" bold="true" box="[444,550,1665,1687]" pageId="40" pageNumber="41">Figure 27.</emphasis>
Age-independent growth curves for a large, old
<taxonomicName id="6FB84D5FFFEBFF82FBB0F937FA45F952" box="[1069,1439,1665,1687]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFEBFF82FBB0F937FB1AF953" box="[1069,1216,1666,1687]" italics="true" pageId="40" pageNumber="41">Tyrannosaurus</emphasis>
, Sue FMNH PR 2081
</taxonomicName>
(red, circles), and two
<taxonomicName id="6FB84D5FFFEBFF82FDF1F911FC5CF97D" box="[620,902,1700,1721]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFEBFF82FDF1F911FD22F97D" box="[620,760,1700,1721]" italics="true" pageId="40" pageNumber="41">Nanotyrannus</emphasis>
, Petey BMRP
</taxonomicName>
2006.4.4 (green, triangles) and Jane BMRP 2002.4.1 (dark blue, squares), corrected for split multi-LAGs as in Cullen et al. [
<bibRefCitation id="CC294B2DFFEBFF82FBC3F973FBACF91F" author="Cullen, T. M. &amp; Canale, J. I &amp; Apesteguia, S. &amp; Smith, N. D. &amp; Hu, D. &amp; Makovicky, P. J." box="[1118,1142,1734,1755]" journalOrPublisher="Proc. R. Soc. B" pageId="40" pageNumber="41" pagination="20202258" part="287" refId="ref29372" refString="49 Cullen, T. M.; Canale, J. I.; Apesteguia, S.; Smith, N. D.; Hu, D.; Makovicky, P. J. Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution. Proc. R. Soc. B 2020, 287, 20202258. [CrossRef]" title="Osteohistological analyses reveal diverse strategies of theropod dinosaur body-size evolution." type="journal article" year="2020">49</bibRefCitation>
]. LAG #1 and the periosteum are not included in the regressions shown here. Four different growth models are fit to each specimen: logistic (solid), Gompertz (dashed), logarithmic (dotted), and von Bertalanffy (dot—dash). Horizontal black lines are asymptotic masses derived from logistic regressions of multiple individuals (i.e., multiple specimens used in the regression, with each specimen assigned a single mass and age at death) from Longrich et al. (in review) and are presented in decreasing order as follows:
<taxonomicName id="6FB84D5FFFEBFF82FADCF8C4FA0BF842" authorityName=", Sue FMNH PR" authorityYear="2081" box="[1345,1489,1905,1926]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFEBFF82FADCF8C4FA0BF842" box="[1345,1489,1905,1926]" italics="true" pageId="40" pageNumber="41">Tyrannosaurus</emphasis>
</taxonomicName>
,
<taxonomicName id="6FB84D5FFFEBFF82FE26F826FDEFF86C" authorityName="Lambe" authorityYear="1914" box="[443,565,1939,1960]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFEBFF82FE26F826FDEFF86C" box="[443,565,1939,1960]" italics="true" pageId="40" pageNumber="41">Gorgosaurus</emphasis>
</taxonomicName>
,
<taxonomicName id="6FB84D5FFFEBFF82FDDCF826FD12F86C" authorityName="Osborn" authorityYear="1905" box="[577,712,1939,1960]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="40" pageNumber="41" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFEBFF82FDDCF826FD12F86C" box="[577,712,1939,1960]" italics="true" pageId="40" pageNumber="41">Albertosaurus</emphasis>
</taxonomicName>
. Results are similar to those deriving from regressions that include LAG #1 and the periosteum (Supplemental Material).
</paragraph>
</caption>
<caption id="FCC76654FFEAFF83FE21FA34FB1DFA3A" ID-DOI="http://doi.org/10.5281/zenodo.10535329" ID-Zenodo-Dep="10535329" httpUri="https://zenodo.org/record/10535329/files/figure.png" pageId="41" pageNumber="42" startId="41.[444,516,1409,1431]" targetBox="[133,1460,248,1363]" targetPageId="41" targetType="figure">
<paragraph id="A80736DCFFEAFF83FE21FA34FB1DFA3A" blockId="41.[444,1494,1409,1534]" pageId="41" pageNumber="42">
<emphasis id="9ACCEACEFFEAFF83FE21FA34FDF0FA52" bold="true" box="[444,554,1409,1431]" pageId="41" pageNumber="42">Figure 28.</emphasis>
Growth record from Zuri (HRS 081514) pubis, with radius back-estimated from LAG spacing [
<bibRefCitation id="CC294B2DFFEAFF83FD81FA16FDEEFA7D" author="Griffin, C." bookContentInfo="Cedarville, OH, USA," box="[540,564,1443,1465]" pageId="41" pageNumber="42" refId="ref29498" refString="51. Griffin, C. Using osteohistology to determine the taxonomic validity of the Late Cretaceous dinosaur Nanotyrannus lancensis Bakker et al, 1988 (Theropoda: Tyrannosauridae). In Proceedings of the The Research and Scholarship Symposium, Cedarville, OH, USA, 16 April 2014." title="Using osteohistology to determine the taxonomic validity of the Late Cretaceous dinosaur Nanotyrannus lancensis Bakker et al, 1988 (Theropoda: Tyrannosauridae)" type="proceedings paper" volumeTitle="Proceedings of the The Research and Scholarship Symposium" year="2014">51</bibRefCitation>
]. Models including the periosteum and LAG 1 are in purple; excluding the periosteum and LAG 1 are in orange. Four different growth models are fit under both conditions: logistic (solid), Gompertz (dashed), logarithmic (dotted), and von Bertalanffy (dotdash).
</paragraph>
</caption>
<paragraph id="A80736DCFFEAFF83FE69F994FD70F8A2" blockId="41.[443,1496,1569,2062]" pageId="41" pageNumber="42">
All mass estimates for adult
<taxonomicName id="6FB84D5FFFEAFF83FCC9F997FC2FF9FD" box="[852,1013,1570,1593]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFEAFF83FCC9F997FC2FF9FD" box="[852,1013,1570,1593]" italics="true" pageId="41" pageNumber="42">Nanotyrannus</emphasis>
</taxonomicName>
are far below those expected for
<taxonomicName id="6FB84D5FFFEAFF83FA0CF997FA09F9FD" box="[1425,1491,1570,1593]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFEAFF83FA0CF997FA78F9FD" box="[1425,1442,1570,1593]" italics="true" pageId="41" pageNumber="42">T</emphasis>
.
<emphasis id="9ACCEACEFFEAFF83FA2CF997FA09F9FD" box="[1457,1491,1570,1593]" italics="true" pageId="41" pageNumber="42">rex</emphasis>
</taxonomicName>
(
<figureCitation id="30832A59FFEAFF83FE5FF9F6FDF4F99F" box="[450,558,1603,1627]" captionStart="Figure 27" captionStartId="40.[444,515,1665,1687]" captionTargetBox="[213,1374,254,1612]" captionTargetId="figure-181@40.[209,1414,220,1633]" captionTargetPageId="40" captionText="Figure 27. Age-independent growth curves for a large, old Tyrannosaurus, Sue FMNH PR 2081 (red, circles), and two Nanotyrannus, Petey BMRP 2006.4.4 (green, triangles) and Jane BMRP 2002.4.1 (dark blue, squares), corrected for split multi-LAGs as in Cullen et al. [49]. LAG #1 and the periosteum are not included in the regressions shown here. Four different growth models are fit to each specimen: logistic (solid), Gompertz (dashed), logarithmic (dotted), and von Bertalanffy (dot—dash). Horizontal black lines are asymptotic masses derived from logistic regressions of multiple individuals (i.e., multiple specimens used in the regression, with each specimen assigned a single mass and age at death) from Longrich et al. (in review) and are presented in decreasing order as follows: Tyrannosaurus, Gorgosaurus, Albertosaurus. Results are similar to those deriving from regressions that include LAG #1 and the periosteum (Supplemental Material)." figureDoi="http://doi.org/10.5281/zenodo.10535327" httpUri="https://zenodo.org/record/10535327/files/figure.png" pageId="41" pageNumber="42">Figure 27</figureCitation>
), which is predicted to hit ~
<quantity id="6F409B39FFEAFF83FCF6F9F6FC1EF99F" box="[875,964,1603,1627]" metricMagnitude="3" metricUnit="kg" metricValue="8.0" pageId="41" pageNumber="42" unit="kg" value="8000.0">8000 kg</quantity>
or more depending upon the model and mass estimates used. Growth trajectories of
<collectionCode id="CEA9AE19FFEAFF83FC14F9D0FC0CF9B8" box="[905,982,1637,1660]" collectionName="BMRP" pageId="41" pageNumber="42">BMRP</collectionCode>
<date id="DC06101CFFEAFF83FC40F9D0FBE6F9B9" box="[989,1084,1637,1661]" pageId="41" pageNumber="42" value="2002-04-01">2002.4.1</date>
and
<collectionCode id="CEA9AE19FFEAFF83FBE5F9D0FB1FF9B8" box="[1144,1221,1637,1660]" collectionName="BMRP" pageId="41" pageNumber="42">BMRP</collectionCode>
<date id="DC06101CFFEAFF83FB51F9D1FAF0F9B8" box="[1228,1322,1636,1660]" pageId="41" pageNumber="42" value="2006-04-04">2006.4.4</date>
are, therefore, inconsistent with their identification as juvenile
<taxonomicName id="6FB84D5FFFEAFF83FC70F932FB6CF95A" box="[1005,1206,1671,1694]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFEAFF83FC70F932FB6CF95A" box="[1005,1206,1671,1694]" italics="true" pageId="41" pageNumber="42">Tyrannosaurus rex</emphasis>
</taxonomicName>
, even under a variety of growth models and initial conditions during curve fitting (Supplemental Material). Our estimates instead suggest that they represent a distinct, small-bodied taxon. Although it is conceivable that young
<taxonomicName id="6FB84D5FFFEAFF83FC90F95EFC69F8C6" authorityName=", Sue FMNH PR" authorityYear="2081" box="[781,947,1771,1794]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFEAFF83FC90F95EFC69F8C6" box="[781,947,1771,1794]" italics="true" pageId="41" pageNumber="42">Tyrannosaurus</emphasis>
</taxonomicName>
sometimes showed slow growth rates due to sickness, lack of food, or other stresses, it is unlikely that all three individuals sectioned would exhibit similar growth anomalies; it is more likely that they exhibit typical growth rates for their taxon.
</paragraph>
<paragraph id="A80736DCFFEAFF80FE69F8C5FDA8FE74" blockId="41.[443,1496,1569,2062]" lastBlockId="42.[444,1496,240,432]" lastPageId="42" lastPageNumber="43" pageId="41" pageNumber="42">
Another alternative hypothesis for this variation in growth trajectories, while assuming taxonomic synonymy, would be that the putative
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<emphasis id="9ACCEACEFFEAFF83FBBDF827FB61F86D" box="[1056,1211,1938,1961]" italics="true" pageId="41" pageNumber="42">Nanotyrannus</emphasis>
</taxonomicName>
specimens are members of the smaller sex in
<taxonomicName id="6FB84D5FFFEAFF83FD34F801FD3FF80F" box="[681,741,1972,1995]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFEAFF83FD34F801FD3FF80F" box="[681,741,1972,1995]" italics="true" pageId="41" pageNumber="42">T. rex</emphasis>
</taxonomicName>
. While it is reasonable to assume that Sue is fairly representative of average adult
<taxonomicName id="6FB84D5FFFEAFF83FD1FF860FD1AF828" box="[642,704,2005,2028]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFEAFF83FD1FF860FD1AF828" box="[642,704,2005,2028]" italics="true" pageId="41" pageNumber="42">T. rex</emphasis>
</taxonomicName>
size for its sex (i.e., as far as fossil discovery approximates random sampling of the population of
<taxonomicName id="6FB84D5FFFEAFF83FC88F842FC8AF7CA" box="[789,848,2039,2062]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="rex">
<emphasis id="9ACCEACEFFEAFF83FC88F842FC8AF7CA" box="[789,848,2039,2062]" italics="true" pageId="41" pageNumber="42">T. rex</emphasis>
</taxonomicName>
), the magnitude of hypothetical body mass dimorphism between Sue and the
<taxonomicName id="6FB84D5FFFE9FF80FD2FFF44FC97FECC" box="[690,845,241,264]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="42" pageNumber="43" phylum="Chordata" rank="genus">
<emphasis id="9ACCEACEFFE9FF80FD2FFF44FC97FECC" box="[690,845,241,264]" italics="true" pageId="42" pageNumber="43">Nanotyrannus</emphasis>
</taxonomicName>
specimens from at least the asymptotic models (Supplemental Material) would be implausible. This hypothetical dimorphism would exceed those estimated or observed in other non-avian [
<bibRefCitation id="CC294B2DFFE9FF80FC28FE86FC07FE8F" author="Saitta, E. T. &amp; Stockdale, M. T. &amp; Longrich, N. R. &amp; Bonhomme, V. &amp; Benton, M. J. &amp; Cuthill, I. C. &amp; Makovicky, P. J." box="[949,989,307,331]" journalOrPublisher="Biol. J. Linn. Soc." pageId="42" pageNumber="43" pagination="231 - 273" part="131" refId="ref34215" refString="134. Saitta, E. T.; Stockdale, M. T.; Longrich, N. R.; Bonhomme, V.; Benton, M. J.; Cuthill, I. C.; Makovicky, P. J. An effect size statistical framework for investigating sexual dimorphism in non-avian dinosaurs and other extinct taxa. Biol. J. Linn. Soc. 2020, 131, 231 - 273. [CrossRef]" title="An effect size statistical framework for investigating sexual dimorphism in non-avian dinosaurs and other extinct taxa" type="journal article" year="2020">134</bibRefCitation>
] and avian [
<bibRefCitation id="CC294B2DFFE9FF80FBEFFE86FB57FE8F" author="Dunning, J. B." bookContentInfo="2 nd ed.; CRC Press: Boca Raton, FL, USA" box="[1138,1165,307,331]" pageId="42" pageNumber="43" pagination="672" refId="ref31992" refString="93. Dunning, J. B. Handbook of Avian Body Masses, 2 nd ed.; CRC Press: Boca Raton, FL, USA, 2007; p. 672." title="Handbook of Avian Body Masses" type="book" year="2007">93</bibRefCitation>
] dinosaurs, highly sexually dimorphic mammals such as sperm whales [
<bibRefCitation id="CC294B2DFFE9FF80FC4EFEE1FC21FEA8" author="Ralls, K. &amp; Mesnick, S." bookContentInfo="Elsevier: Amsterdam, The Netherlands" box="[979,1019,340,364]" pageId="42" pageNumber="43" pagination="1005 - 1011" refId="ref34299" refString="135. Ralls, K.; Mesnick, S. Sexual Dimorphism. In Encyclopedia of Marine Mammals; Elsevier: Amsterdam, The Netherlands, 2009; pp. 1005 - 1011." title="Sexual Dimorphism" type="book chapter" volumeTitle="Encyclopedia of Marine Mammals" year="2009">135</bibRefCitation>
], and would only be comparable to the most extreme examples of sexual dimorphism in extant tetrapods (e.g., southern elephant seals [
<bibRefCitation id="CC294B2DFFE9FF80FD9EFE22FDF1FE6B" author="Laws, R. M." bookContentInfo="HMSO: London, UK," box="[515,555,407,431]" pageId="42" pageNumber="43" pagination="62" refId="ref34334" refString="136. Laws, R. M. The elephant seal (Mirounga leonina, Linn.): I. Growth and age; HMSO: London, UK, 1953; p. 62." title="The elephant seal (Mirounga leonina, Linn.): I. Growth and age" type="book chapter" year="1953">136</bibRefCitation>
,
<bibRefCitation id="CC294B2DFFE9FF80FDAFFE2DFD80FE74" author="McCann, T. S. &amp; Fedak, M. A. &amp; Harwood, J." box="[562,602,408,432]" journalOrPublisher="Behav. Ecol. Sociobiol." pageId="42" pageNumber="43" pagination="81 - 87" part="25" refId="ref34371" refString="137. McCann, T. S.; Fedak, M. A.; Harwood, J. Parental investment in southern elephant seals, Mirounga leonina. Behav. Ecol. Sociobiol. 1989, 25, 81 - 87. [CrossRef]" title="Parental investment in southern elephant seals, Mirounga leonina" type="journal article" year="1989">137</bibRefCitation>
]).
</paragraph>
</subSubSection>
</treatment>
</document>