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<document ID-DOI="http://doi.org/10.5281/zenodo.3717027" ID-GBIF-Dataset="d73d45db-a61c-481d-9e02-685676e51f73" ID-GBIF-Taxon="162486981" ID-Zenodo-Dep="3717027" approvalRequired="15" approvalRequired_for_document="1" approvalRequired_for_illustrations="11" approvalRequired_for_originalDoi="1" approvalRequired_for_textStreams="2" checkinTime="1582194442820" checkinUser="jeremy" docAuthor="Carpenter, Kenneth" docDate="1990" docId="039987D5DD7C8D380FFAF9A8F0899C30" docLanguage="en" docName="Carpenter1990VariationTyrannosaurus.pdf.imf" docOrigin="Dinosaur Systematics: Perspectives and Approaches, Cambridge: Cambridge University Press" docStyle="DocumentStyle{}" docTitle="Tyrannosaurus rex Osborn 1905" docType="treatment" docVersion="20" lastPageId="3" lastPageNumber="144" masterDocId="FFA0FFADDD7C8D3B0B4DFFC2F65F9F23" masterDocTitle="Variation in Tyrannosaurus rex" masterLastPageNumber="145" masterPageNumber="141" pageId="0" pageNumber="141" updateTime="1635896949400" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Variation in Tyrannosaurus rex</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Carpenter, Kenneth</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:originInfo>
<mods:dateIssued>1990</mods:dateIssued>
<mods:dateOther type="pubDate">1990-12-31</mods:dateOther>
<mods:publisher>Cambridge University Press</mods:publisher>
<mods:place>
<mods:placeTerm>Cambridge</mods:placeTerm>
</mods:place>
</mods:originInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Editor</mods:roleTerm>
</mods:role>
<mods:namePart>Carpenter, Kenneth</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Editor</mods:roleTerm>
</mods:role>
<mods:namePart>Currie, Philip J.</mods:namePart>
</mods:name>
<mods:titleInfo>
<mods:title>Dinosaur Systematics: Perspectives and Approaches</mods:title>
</mods:titleInfo>
<mods:part>
<mods:extent unit="page">
<mods:start>141</mods:start>
<mods:end>145</mods:end>
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<mods:classification>book chapter</mods:classification>
<mods:identifier type="DOI">10.5281/zenodo.3676505</mods:identifier>
<mods:identifier type="GBIF-Dataset">d73d45db-a61c-481d-9e02-685676e51f73</mods:identifier>
<mods:identifier type="Zenodo-Dep">3676505</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.3717027" ID-GBIF-Taxon="162486981" ID-Zenodo-Dep="3717027" LSID="urn:lsid:plazi:treatment:039987D5DD7C8D380FFAF9A8F0899C30" httpUri="http://treatment.plazi.org/id/039987D5DD7C8D380FFAF9A8F0899C30" lastPageId="3" lastPageNumber="144" pageId="0" pageNumber="141">
<subSubSection box="[1207,1314,1642,1673]" pageId="0" pageNumber="141" type="description">
<paragraph blockId="0.[1105,1955,1638,2817]" box="[1207,1314,1642,1673]" pageId="0" pageNumber="141">
<heading bold="true" box="[1207,1314,1642,1673]" fontSize="10" level="2" pageId="0" pageNumber="141" reason="6">
<emphasis bold="true" box="[1207,1314,1642,1673]" pageId="0" pageNumber="141">Skulls</emphasis>
</heading>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="141" type="nomenclature">
<paragraph blockId="0.[1105,1955,1638,2817]" pageId="0" pageNumber="141">
Six partial and complete skulls are now known for
<emphasis box="[1159,1449,1734,1765]" italics="true" pageId="0" pageNumber="141">
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1159,1443,1734,1765]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="141" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
,
</emphasis>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="141" type="materials_examined">
<paragraph blockId="0.[1105,1955,1638,2817]" pageId="0" pageNumber="141">
including the holotype
<materialsCitation ID-GBIF-Occurrence="3396373321" collectionCode="CM" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=52853 " pageId="0" pageNumber="141" specimenCode="CM 9380 (formerly AMNH 973)" typeStatus="holotype">CM 9380 (formerly AMNH 973)</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="3396373302" box="[1493,1704,1778,1810]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="0" pageNumber="141" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="3396373318" box="[1724,1948,1778,1810]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/Display.php?irn=1028446&amp;QueryPage=%2Fdinosaur-institute%2F&amp;BackRef=ResultsList.php " pageId="0" pageNumber="141" specimenCode="LACM 23844">LACM 23844</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="2575874708" box="[1106,1263,1825,1857]" collectionCode="MOR" pageId="0" pageNumber="141" specimenCode="MOR 009">MOR 009</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="2575874703" box="[1284,1498,1825,1857]" collectionCode="SDSM" pageId="0" pageNumber="141" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
, and
<materialsCitation ID-GBIF-Occurrence="3396373336" box="[1587,1774,1825,1857]" collectionCode="TMP" pageId="0" pageNumber="141" specimenCode="TMP 81.6.1">TMP 81.6.1</materialsCitation>
. Two additional specimens,
<materialsCitation ID-GBIF-Occurrence="3396373335" box="[1391,1597,1870,1902]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47763 " pageId="0" pageNumber="141" specimenCode="AMNH 5029">AMNH 5029</materialsCitation>
and
<materialsCitation ID-GBIF-Occurrence="3396373322" box="[1680,1885,1870,1902]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47861 " pageId="0" pageNumber="141" specimenCode="AMNH 5117">AMNH 5117</materialsCitation>
, are braincases (
<bibRefCitation author="Osborn, H." box="[1284,1489,1916,1948]" journalOrPublisher="American Museum of Natural History Memoirs, new series" pageId="0" pageNumber="141" pagination="1 - 30" part="1" refId="ref2376" refString="1912. Crania of Tyrannosaurus and Allosaurus. American Museum of Natural History Memoirs, new series 1: 1 - 30." title="Crania of Tyrannosaurus and Allosaurus" type="book chapter" year="1912">Osborn 1912</bibRefCitation>
). Five of the skulls are shown in
<figureCitation box="[1148,1325,1962,1994]" captionStart="Figure 10.1" captionText="Figure 10.1. A comparison of four Tyrannosaurus rex skulls. A, CM 9380 (formerly AMNH 973); B, AMNH 5027; C, LACM 23844; D, TMP 81.6.1; E, SDSM 12047. The reconstruction of CM 9380 differs from that given by Osborn (1906) in the orbital and postorbital regions. Osborn had erroneously used the right ectoptygoid as a right postorbital in the reconstructed skull at the Carnegie Museum. As may be seen in the figure, the squamosal no longer extends into the orbit. TMP 81.6.1 is more complete than shown, but preparation was not complete at the time this figure was made. SDSM 12047 is crushed dorsolaterally, which has greatly affected the snout, no attempt has been made to compensate for this. (Scale bar = lOcm.)" httpUri="https://zenodo.org/record/4030834/files/figure.png" pageId="0" pageNumber="141" targetBox="[101,1845,1061,2798]" targetBoxID="1">Figure 10.1</figureCitation>
. As may be seen, there is a considerable amount of variation in the size and shape of all the cranial openings (e.g., orbit and lateral temporal fenestra), as well as all the individual elements, including the lacrimal, postorbital, quadratojugal, jugal, and surangular. In fact, no two specimens are identical.
</paragraph>
</subSubSection>
<subSubSection lastPageId="3" lastPageNumber="144" pageId="0" pageNumber="141" type="description">
<paragraph blockId="0.[1105,1955,1638,2817]" pageId="0" pageNumber="141">
Seven maxillae were available for comparison. Five of these were overlain in
<figureCitation box="[1592,1796,2281,2313]" captionStart="Figure 10.2" captionText="Figure 10.2. A, an overlay of maxillae: CM 9380, AMNH 5027, LACM 23844, TMP 81.6.1, and TMM 41436-1. Teeth not shown. B, right (reversed) and left maxillae of LACM 23844 and C, AMNH 5027." httpUri="https://zenodo.org/record/3676509/files/figure.png" pageId="0" pageNumber="141" targetBox="[161,998,402,2017]" targetBoxID="2">Figure 10.2A</figureCitation>
using the anterior-most margins of the maxilla and maxillary fenestra as standard lengths. Most of the maxillae resemble one another, except for
<materialsCitation ID-GBIF-Occurrence="3396373311" box="[1488,1724,2418,2450]" collectionCode="TMM" pageId="0" pageNumber="141" specimenCode="TMM 41436-1">TMM 41436-1</materialsCitation>
, the specimen reported from the Tornillo Formation by
<bibRefCitation author="Lawson, D." box="[1712,1938,2464,2496]" journalOrPublisher="Journal of Paleontology" pageId="0" pageNumber="141" pagination="158 - 164" part="50" refId="ref2242" refString="Lawson, D. 1976. Tyrannosaurus and Torosaurus, Maestrichtian dinosaurs from Trans-Pecos, Texas. Journal of Paleontology 50: 158 - 164." title="Tyrannosaurus and Torosaurus, Maestrichtian dinosaurs from Trans-Pecos, Texas" type="journal article" year="1976">Lawson (1976)</bibRefCitation>
.
</paragraph>
<paragraph blockId="0.[1105,1955,1638,2817]" lastBlockId="1.[102,954,190,681]" lastPageId="1" lastPageNumber="142" pageId="0" pageNumber="141">
Variation can be seen in the depth of the maxilla, size and shape of the maxillary and antorbital fenestrae, position of the lacrimal processes, and the position and shape of the jugal process (
<figureCitation box="[1534,1705,2647,2679]" captionStart="Figure 10.2" captionText="Figure 10.2. A, an overlay of maxillae: CM 9380, AMNH 5027, LACM 23844, TMP 81.6.1, and TMM 41436-1. Teeth not shown. B, right (reversed) and left maxillae of LACM 23844 and C, AMNH 5027." httpUri="https://zenodo.org/record/3676509/files/figure.png" pageId="0" pageNumber="141" targetBox="[161,998,402,2017]" targetBoxID="2">Fig. 10.2A</figureCitation>
). Differences in the depth of the maxilla affect the position of the lacrimal process, which in tum influences the height and shape of the antorbital fenestra. There is also a considerable amount of variation in position of the jugal process that forms the lower rim of the antorbital fenestra, but this does not seem to correlate with the size of the antorbital process. The shape of the maxillary fenestra ranges from almost oval (
<materialsCitation ID-GBIF-Occurrence="2575874710" box="[396,546,375,407]" collectionCode="CM" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=52853 " pageId="1" pageNumber="142" specimenCode="CM 9380" typeStatus="holotype">CM 9380</materialsCitation>
) to subtriangular (
<materialsCitation ID-GBIF-Occurrence="3396373330" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/Display.php?irn=1028446&amp;QueryPage=%2Fdinosaur-institute%2F&amp;BackRef=ResultsList.php " pageId="1" pageNumber="142" specimenCode="LACM 23844">LACM 23844</materialsCitation>
and
<materialsCitation ID-GBIF-Occurrence="3396373307" box="[294,495,420,452]" collectionCode="TMP" pageId="1" pageNumber="142" specimenCode="TMP 81.6.1">TMP 81.6.1</materialsCitation>
) to almost square (
<materialsCitation ID-GBIF-Occurrence="3396373312" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47745 " pageId="1" pageNumber="142" specimenCode="AMNH 5012">AMNH 5012</materialsCitation>
). There is little or no difference in the position of the largest teeth and therefore this feature is not affected by the depth of the maxilla or the shape of the antorbital fenestra. Even the left and right maxilla of the same skull show variation (
<figureCitation box="[426,638,648,680]" captionStart="Figure 10.2" captionText="Figure 10.2. A, an overlay of maxillae: CM 9380, AMNH 5027, LACM 23844, TMP 81.6.1, and TMM 41436-1. Teeth not shown. B, right (reversed) and left maxillae of LACM 23844 and C, AMNH 5027." httpUri="https://zenodo.org/record/3676509/files/figure.png" pageId="1" pageNumber="142" targetBox="[161,998,402,2017]" targetBoxID="2">Figs. 10.2B,C</figureCitation>
).
</paragraph>
<paragraph blockId="1.[1001,1851,190,681]" lastBlockId="2.[1057,1910,196,1188]" lastPageId="2" lastPageNumber="143" pageId="1" pageNumber="142">
The maxilla from the Tornillo Formation lacks the anterior-most margin and was scaled to the other maxilla using height and the anterior margin of the maxillary fenestra (
<figureCitation box="[1231,1397,328,360]" captionStart="Figure 10.2" captionText="Figure 10.2. A, an overlay of maxillae: CM 9380, AMNH 5027, LACM 23844, TMP 81.6.1, and TMM 41436-1. Teeth not shown. B, right (reversed) and left maxillae of LACM 23844 and C, AMNH 5027." httpUri="https://zenodo.org/record/3676509/files/figure.png" pageId="1" pageNumber="142" targetBox="[161,998,402,2017]" targetBoxID="2">Fig. 10.2A</figureCitation>
). The nasal, or dorsal, margin of the maxilla arcs sharply down suggesting a face considerably shorter than in
<emphasis box="[1370,1459,421,452]" italics="true" pageId="1" pageNumber="142">
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1370,1455,421,452]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="142" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
.
</emphasis>
Other differences include a slightly larger maxillary fenestra in proportion to the size of the maxilla and a much deeper jugal process. The antorbital fenestra appears to be smaller, but the lacrimal process is too incomplete to be certain. These differences are great enough to suggest that
<materialsCitation ID-GBIF-Occurrence="3396373309" box="[1481,1697,648,680]" collectionCode="TMM" pageId="1" pageNumber="142" specimenCode="TMM 41436-1">TMM 41436-1</materialsCitation>
falls outside the range of individual variation for
<emphasis italics="true" pageId="2" pageNumber="143">
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="143" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
.
</emphasis>
It may belong to a new genus, but no name should be proposed until additional material is available. Because it is unusual, it is excluded from discussions on variation.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.4030834" ID-Zenodo-Dep="4030834" httpUri="https://zenodo.org/record/4030834/files/figure.png" pageId="1" pageNumber="142" targetBox="[101,1845,1061,2798]" targetPageId="1">
<paragraph blockId="1.[201,1834,729,1008]" pageId="1" pageNumber="142">
Figure 10.1. A comparison of four
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[675,925,731,759]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="142" phylum="Chordata" rank="species" species="rex">
<emphasis box="[675,925,731,759]" italics="true" pageId="1" pageNumber="142">Tyrannosaurus rex</emphasis>
</taxonomicName>
skulls.
<emphasis bold="true" box="[1028,1054,731,759]" pageId="1" pageNumber="142">A</emphasis>
,
<materialsCitation ID-GBIF-Occurrence="3396373325" box="[1069,1512,731,759]" collectionCode="CM" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=52853 " pageId="1" pageNumber="142" specimenCode="CM 9380" typeStatus="holotype">CM 9380 (formerly AMNH 973)</materialsCitation>
;
<emphasis bold="true" box="[1526,1555,730,759]" pageId="1" pageNumber="142">B,</emphasis>
<materialsCitation ID-GBIF-Occurrence="3396373308" box="[1566,1744,731,759]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="1" pageNumber="142" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
;
<emphasis bold="true" box="[1760,1786,731,759]" pageId="1" pageNumber="142">C</emphasis>
,
<materialsCitation ID-GBIF-Occurrence="3396373342" box="[201,392,773,801]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/Display.php?irn=1028446&amp;QueryPage=%2Fdinosaur-institute%2F&amp;BackRef=ResultsList.php " pageId="1" pageNumber="142" specimenCode="LACM 23844">LACM 23844</materialsCitation>
;
<emphasis box="[407,432,773,801]" italics="true" pageId="1" pageNumber="142">D</emphasis>
,
<materialsCitation ID-GBIF-Occurrence="3396373333" box="[447,608,773,801]" collectionCode="TMP" pageId="1" pageNumber="142" specimenCode="TMP 81.6.1">TMP 81.6.1</materialsCitation>
; E,
<materialsCitation ID-GBIF-Occurrence="3396373306" box="[663,848,773,801]" collectionCode="SDSM" pageId="1" pageNumber="142" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
. The reconstruction of
<materialsCitation ID-GBIF-Occurrence="2575874711" box="[1157,1283,773,801]" collectionCode="CM" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=52853 " pageId="1" pageNumber="142" specimenCode="CM 9380" typeStatus="holotype">CM 9380</materialsCitation>
differs from that given by
<bibRefCitation author="Osborn, H." box="[1642,1834,773,801]" journalOrPublisher="American Museum of Natural History Bulletin" pageId="1" pageNumber="142" pagination="281 - 296" part="22" refId="ref2355" refString="1906. Tyrannosaurus, Upper Cretaceous carnivorous dinosaur. American Museum of Natural History Bulletin 22: 281 - 296." title="Tyrannosaurus, Upper Cretaceous carnivorous dinosaur" type="journal article" year="1906">Osborn (1906)</bibRefCitation>
in the orbital and postorbital regions. Osborn had erroneously used the right ectoptygoid as a right postorbital in the reconstructed skull at the Carnegie Museum. As may be seen in the figure, the squamosal no longer extends into the orbit.
<materialsCitation ID-GBIF-Occurrence="3396373310" collectionCode="TMP" pageId="1" pageNumber="142" specimenCode="TMP 81.6.1">TMP 81.6.1</materialsCitation>
is more complete than shown, but preparation was not complete at the time this figure was made.
<materialsCitation ID-GBIF-Occurrence="3396373304" box="[1593,1774,898,926]" collectionCode="SDSM" pageId="1" pageNumber="142" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
is crushed dorsolaterally, which has greatly affected the snout, no attempt has been made to compensate for this. (Scale bar = lOcm.)
</paragraph>
</caption>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3676509" ID-Zenodo-Dep="3676509" httpUri="https://zenodo.org/record/3676509/files/figure.png" pageId="2" pageNumber="143" targetBox="[161,998,402,2017]" targetPageId="2">
<paragraph blockId="2.[259,977,198,354]" pageId="2" pageNumber="143">
Figure 10.2.
<emphasis bold="true" box="[438,462,198,230]" pageId="2" pageNumber="143">A</emphasis>
, an overlay of maxillae:
<materialsCitation ID-GBIF-Occurrence="2575874712" box="[795,924,198,230]" collectionCode="CM" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=52853 " pageId="2" pageNumber="143" specimenCode="CM 9380" typeStatus="holotype">CM 9380</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="3396373314" box="[259,438,240,272]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="2" pageNumber="143" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="3396373323" box="[452,642,240,272]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/Display.php?irn=1028446&amp;QueryPage=%2Fdinosaur-institute%2F&amp;BackRef=ResultsList.php " pageId="2" pageNumber="143" specimenCode="LACM 23844">LACM 23844</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="3396373334" box="[657,819,240,272]" collectionCode="TMP" pageId="2" pageNumber="143" specimenCode="TMP 81.6.1">TMP 81.6.1</materialsCitation>
, and
<materialsCitation ID-GBIF-Occurrence="3396373316" collectionCode="TMM" pageId="2" pageNumber="143" specimenCode="TMM 41436-1">TMM 41436-1</materialsCitation>
. Teeth not shown.
<emphasis bold="true" box="[623,652,283,312]" pageId="2" pageNumber="143">B,</emphasis>
right (reversed) and left maxillae of
<materialsCitation ID-GBIF-Occurrence="3396373313" box="[420,605,322,354]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/Display.php?irn=1028446&amp;QueryPage=%2Fdinosaur-institute%2F&amp;BackRef=ResultsList.php " pageId="2" pageNumber="143" specimenCode="LACM 23844">LACM 23844</materialsCitation>
and
<emphasis bold="true" box="[672,695,322,354]" pageId="2" pageNumber="143">C</emphasis>
,
<materialsCitation ID-GBIF-Occurrence="3396373338" box="[711,890,322,354]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="2" pageNumber="143" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
.
</paragraph>
</caption>
<paragraph blockId="2.[1057,1910,196,1188]" pageId="2" pageNumber="143">
Variation in the dentary is demonstrated by six specimens (
<figureCitation box="[1235,1374,472,504]" captionStart="Figure 10.3" captionText="Figure 10.3. An overlay of the dentaries of CM 9380, AMNH 5027, BM(NH) R7994 (formerly AMNH 5866), LACM 23844, SDSM 12047, and TMP 81.6.1. Teeth not shown." httpUri="https://zenodo.org/record/3724120/files/figure.png" pageId="2" pageNumber="143" targetBox="[161,998,402,2017]" targetBoxID="2">Fig. 10.3</figureCitation>
). The distance between the anterior tip and the highest part of the dentary were used for the standard length. The greatest difference is the depth of the posteroventral margin of the dentary. It is relatively shallowest in
<materialsCitation ID-GBIF-Occurrence="3396373339" box="[1265,1471,655,687]" collectionCode="SDSM" pageId="2" pageNumber="143" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
and deepest in
<materialsCitation ID-GBIF-Occurrence="3396373301" box="[1706,1905,655,687]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="2" pageNumber="143" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
(
<figureCitation box="[1070,1210,701,733]" captionStart="Figure 10.3" captionText="Figure 10.3. An overlay of the dentaries of CM 9380, AMNH 5027, BM(NH) R7994 (formerly AMNH 5866), LACM 23844, SDSM 12047, and TMP 81.6.1. Teeth not shown." httpUri="https://zenodo.org/record/3724120/files/figure.png" pageId="2" pageNumber="143" targetBox="[161,998,402,2017]" targetBoxID="2">Fig. 10.3</figureCitation>
). However,
<materialsCitation ID-GBIF-Occurrence="3396373324" box="[1392,1590,701,733]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="2" pageNumber="143" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
also has a tooth row that curves more ventrally than the others and this may have had an influence on the posteroventral margin. The posterior margin of the dentary is thin and easily damaged, although three of the dentaries have complete margins (
<materialsCitation ID-GBIF-Occurrence="2575874704" box="[1209,1365,928,960]" collectionCode="CM" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=52853 " pageId="2" pageNumber="143" specimenCode="CM 9380" typeStatus="holotype">CM 9380</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="3396373320" box="[1386,1599,928,960]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="2" pageNumber="143" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
, and
<materialsCitation ID-GBIF-Occurrence="3396373305" box="[1691,1888,928,960]" collectionCode="TMP" pageId="2" pageNumber="143" specimenCode="TMP 81.6.1">TMP 81.6.1</materialsCitation>
). The shape of the anterior portion of the external mandibular fossa (expressed as a notch at the lower edge of the posterior margin, see
<figureCitation box="[1476,1624,1065,1097]" captionStart="Figure 10.1" captionText="Figure 10.1. A comparison of four Tyrannosaurus rex skulls. A, CM 9380 (formerly AMNH 973); B, AMNH 5027; C, LACM 23844; D, TMP 81.6.1; E, SDSM 12047. The reconstruction of CM 9380 differs from that given by Osborn (1906) in the orbital and postorbital regions. Osborn had erroneously used the right ectoptygoid as a right postorbital in the reconstructed skull at the Carnegie Museum. As may be seen in the figure, the squamosal no longer extends into the orbit. TMP 81.6.1 is more complete than shown, but preparation was not complete at the time this figure was made. SDSM 12047 is crushed dorsolaterally, which has greatly affected the snout, no attempt has been made to compensate for this. (Scale bar = lOcm.)" httpUri="https://zenodo.org/record/4030834/files/figure.png" pageId="2" pageNumber="143" targetBox="[101,1845,1061,2798]" targetBoxID="1">Fig. 10.1</figureCitation>
) is variable. The teeth seem to show more variation in positional size, but this may be due to how the dentaries were overlain.
</paragraph>
<paragraph blockId="2.[1056,1909,1244,1920]" box="[1158,1318,1247,1279]" pageId="2" pageNumber="143">
<heading bold="true" box="[1158,1318,1247,1279]" fontSize="10" level="2" pageId="2" pageNumber="143" reason="6">
<emphasis bold="true" box="[1158,1318,1247,1279]" pageId="2" pageNumber="143">Cervicals</emphasis>
</heading>
</paragraph>
<paragraph blockId="2.[1056,1909,1244,1920]" pageId="2" pageNumber="143">
Complete cervical series are known for only two specimens,
<materialsCitation ID-GBIF-Occurrence="3396373317" box="[1234,1434,1339,1371]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="2" pageNumber="143" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
and
<materialsCitation ID-GBIF-Occurrence="3396373332" box="[1512,1762,1339,1371]" collectionCode="BMNH" httpUri="https://data.nhm.ac.uk/object/5bda6356-b77a-41dc-9e6f-12b4fbe54cce/1568678400000 " pageId="2" pageNumber="143" specimenCode="BMNH R7994">BM(NH) R7994</materialsCitation>
. There is a considerable amount of variation in the shape of the neural spines. Similar variation appears to be present in the tyrannosaur
<emphasis box="[1292,1631,1477,1508]" italics="true" pageId="2" pageNumber="143">
<taxonomicName authorityName="Lambe" authorityYear="1914" box="[1292,1626,1477,1508]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="143" phylum="Chordata" rank="species" species="libratus">Albertosaurus libratus</taxonomicName>
,
</emphasis>
so the use of neural spine shape in the diagnosis of the tyrannosaur
<taxonomicName authorityName="Russell" authorityYear="1970" box="[1056,1409,1568,1599]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="143" phylum="Chordata" rank="species" species="torosus">
<emphasis box="[1056,1409,1568,1599]" italics="true" pageId="2" pageNumber="143">Daspletosaurus torosus</emphasis>
</taxonomicName>
(
<bibRefCitation author="Russell, D." box="[1429,1627,1567,1599]" journalOrPublisher="National Museums of Canada, National Museum of Natural History, Publications in Palaeontology" pageId="2" pageNumber="143" pagination="1 - 34" part="1" refId="ref2489" refString="Russell, D. 1970. Tyrannosaurs from the Late Cretaceous of western Canada. National Museums of Canada, National Museum of Natural History, Publications in Palaeontology 1: 1 - 34." title="Tyrannosaurs from the Late Cretaceous of western Canada" type="journal article" year="1970">Russell 1970</bibRefCitation>
) is suspect.
</paragraph>
<paragraph blockId="2.[1056,1909,1244,1920]" pageId="2" pageNumber="143">
The neck is more robust in
<materialsCitation ID-GBIF-Occurrence="3396373319" box="[1580,1827,1612,1644]" collectionCode="BMNH" httpUri="https://data.nhm.ac.uk/object/5bda6356-b77a-41dc-9e6f-12b4fbe54cce/1568678400000 " pageId="2" pageNumber="143" specimenCode="BMNH R7994">BM(NH) R7994</materialsCitation>
than in
<materialsCitation ID-GBIF-Occurrence="3396373328" box="[1095,1295,1658,1690]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="2" pageNumber="143" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
(
<figureCitation box="[1316,1457,1658,1690]" captionStart="Figure 10.4" captionText="Figure 10.4. A comparison of the cervicals of A, AMNH 5027 and B, BM(NH) R7994. Scale = 10 cm." httpUri="https://zenodo.org/record/3676511/files/figure.png" pageId="2" pageNumber="143" targetBox="[139,979,638,1857]" targetBoxID="3">Fig. 10.4</figureCitation>
). This is especially evident in the atlas and its intercentrum, and in the neural spines of cervicals two and three.
</paragraph>
<paragraph blockId="2.[1056,1909,1244,1920]" pageId="2" pageNumber="143">
Asymmetrical co-ossification of the last cervical and first dorsal in
<materialsCitation ID-GBIF-Occurrence="3396373340" box="[1344,1545,1841,1873]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="2" pageNumber="143" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
has not been observed in any other specimen, and is probably pathological.
</paragraph>
<caption ID-DOI="http://doi.org/10.5281/zenodo.3724120" ID-Zenodo-Dep="3724120" httpUri="https://zenodo.org/record/3724120/files/figure.png" pageId="2" pageNumber="143" startId="2.[257,343,2100,2132]" targetBox="[161,998,402,2017]" targetPageId="2">
<paragraph blockId="2.[257,1841,2100,2793]" pageId="2" pageNumber="143">
Figure 10.3. An overlay of the dentaries of
<materialsCitation ID-GBIF-Occurrence="3396373327" box="[843,972,2100,2132]" collectionCode="CM" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=52853 " pageId="2" pageNumber="143" specimenCode="CM 9380" typeStatus="holotype">CM 9380</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="3396373303" box="[986,1166,2100,2132]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="2" pageNumber="143" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="3396373329" box="[1179,1731,2100,2132]" collectionCode="BMNH" httpUri="https://data.nhm.ac.uk/object/5bda6356-b77a-41dc-9e6f-12b4fbe54cce/1568678400000 " pageId="2" pageNumber="143" specimenCode="BMNH R7994">BM(NH) R7994 (formerly AMNH 5866)</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="3396373341" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/Display.php?irn=1028446&amp;QueryPage=%2Fdinosaur-institute%2F&amp;BackRef=ResultsList.php " pageId="2" pageNumber="143" specimenCode="LACM 23844">LACM 23844</materialsCitation>
,
<materialsCitation ID-GBIF-Occurrence="3396373337" box="[358,542,2142,2174]" collectionCode="SDSM" pageId="2" pageNumber="143" specimenCode="SDSM 12047">SDSM 12047</materialsCitation>
, and
<materialsCitation ID-GBIF-Occurrence="3396373331" box="[612,774,2142,2174]" collectionCode="TMP" pageId="2" pageNumber="143" specimenCode="TMP 81.6.1">TMP 81.6.1</materialsCitation>
. Teeth not shown
</paragraph>
</caption>
<paragraph blockId="3.[123,974,204,468]" box="[225,348,206,238]" pageId="3" pageNumber="144">
<heading bold="true" box="[225,348,206,238]" fontSize="11" level="1" pageId="3" pageNumber="144" reason="0">
<emphasis bold="true" box="[225,348,206,238]" pageId="3" pageNumber="144">Ischiae</emphasis>
</heading>
</paragraph>
<paragraph blockId="3.[123,974,204,468]" lastBlockId="3.[1022,1872,205,788]" pageId="3" pageNumber="144">
Three ischia were available for comparison. Length was standardized and the articular surface of the iliac peduncle was used to determine orientation in
<figureCitation box="[828,967,345,376]" captionStart="Figure 10.5" captionText="Figure 10.5. An overlay of the ischia of A, CM 9380; B, TMP 81.61; and C, AMNH 5027." httpUri="https://zenodo.org/record/3676513/files/figure.png" pageId="3" pageNumber="144" targetBox="[139,979,638,1857]" targetBoxID="3">Fig. 10.5</figureCitation>
. Points of variation include the relative size of the iliac peduncle, the relative size and position of the pubic peduncle, the position and size of the obturator process, and the development of the insertion scar for the M. flexor tibialis internus part 3. This scar
<emphasis box="[1614,1636,299,330]" italics="true" pageId="3" pageNumber="144">is</emphasis>
best developed in
<materialsCitation ID-GBIF-Occurrence="2575874714" box="[1062,1205,344,375]" collectionCode="CM" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=52853 " pageId="3" pageNumber="144" specimenCode="CM 9380" typeStatus="holotype">CM 9380</materialsCitation>
where it forms a prominent ridge.
</paragraph>
<paragraph blockId="3.[1022,1872,205,788]" pageId="3" pageNumber="144">
Of the two types of ischia, one (
<figureCitation box="[1647,1819,390,421]" captionStart="Figure 10.5" captionText="Figure 10.5. An overlay of the ischia of A, CM 9380; B, TMP 81.61; and C, AMNH 5027." httpUri="https://zenodo.org/record/3676513/files/figure.png" pageId="3" pageNumber="144" targetBox="[139,979,638,1857]" targetBoxID="3">Fig. 10.5A</figureCitation>
) is oriented more ventrally from the horizontal articular surface of the iliac peduncle. I suspect that
<materialsCitation ID-GBIF-Occurrence="2575874709" box="[1722,1871,482,513]" collectionCode="CM" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=52853 " pageId="3" pageNumber="144" specimenCode="CM 9380" typeStatus="holotype">CM 9380</materialsCitation>
and
<materialsCitation ID-GBIF-Occurrence="3396373326" box="[1090,1271,527,558]" collectionCode="TMP" pageId="3" pageNumber="144" specimenCode="TMP 81.6.1">TMP 81.6.1</materialsCitation>
are females because the greater angle between the sacral vertebrae and distal end of the ischium would permit the passage of eggs (or live young) more readily than that of
<materialsCitation ID-GBIF-Occurrence="3396373315" box="[1335,1541,665,696]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&amp;specimen_id=47761 " pageId="3" pageNumber="144" specimenCode="AMNH 5027">AMNH 5027</materialsCitation>
. The more divergent ischia are associated with robust skeletons, whereas the less divergent ischium is from a gracile skeleton.
</paragraph>
</subSubSection>
</treatment>
</document>
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