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<document id="01164FBF0D491E1421F1AEBD3D03FFA0" ID-DOI="10.2307/3889334" ID-GBIF-Dataset="6f85aa0a-0368-41a5-83d3-21c33f40845e" ID-Zenodo-Dep="3739929" IM.tables_requiresApprovalFor="existingObjects,jeremy" IM.taxonomicNames_approvedBy="jeremy" approvalRequired="19" approvalRequired_for_bibRefs="2" approvalRequired_for_illustrations="7" approvalRequired_for_taxonomicNames="2" approvalRequired_for_textStreams="8" checkinTime="1585838636713" checkinUser="plazi" docAuthor="Christopher A. Brochu" docDate="2003" docId="9A3A87D00B5E0D67FF2CA9703942F791" docLanguage="en" docName="socVertebratePaleoMemoir_Brochu2003ABBYY_2.pdf" docOrigin="Journal of Vertebrate Paleontology 22" docStyle="DocumentStyle{}" docTitle="Tarbosaurus bataar Maleev 1955" docType="treatment" docVersion="10" lastPageNumber="4" masterDocId="6603FFA80B5D0D64FFF6AB533D39FF9A" masterDocTitle="Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull" masterLastPageNumber="138" masterPageNumber="1" pageNumber="4" updateTime="1689591936236" updateUser="jeremy">
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<mods:title id="C7B8CA05DDFB4D0A5EE82D182D5C8566">Osteology of Tyrannosaurus rex: insights from a nearly complete skeleton and high-resolution computed tomographic analysis of the skull</mods:title>
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<mods:namePart id="2D78F7A9E00B75D037264006503F879D">Christopher A. Brochu</mods:namePart>
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<treatment id="9A3A87D00B5E0D67FF2CA9703942F791" ID-DOI="http://doi.org/10.5281/zenodo.3810815" ID-GBIF-Taxon="190303345" ID-Zenodo-Dep="3810815" LSID="urn:lsid:plazi:treatment:9A3A87D00B5E0D67FF2CA9703942F791" httpUri="http://treatment.plazi.org/id/9A3A87D00B5E0D67FF2CA9703942F791" lastPageNumber="4" pageId="3" pageNumber="4">
<subSubSection id="5A89654D0B5E0D67FF2CA9703E1AFDDE" box="[218,803,545,582]" pageId="3" pageNumber="4" type="nomenclature">
<paragraph id="122C36C60B5E0D67FF2CA9703E1AFDDE" blockId="3.[172,1232,285,3161]" box="[218,803,545,582]" pageId="3" pageNumber="4">
<taxonomicName id="D5934D450B5E0D67FF2CA9703E1AFDDE" authority="Maleev, 1955" authorityName="Maleev" authorityYear="1955" box="[218,803,545,582]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="bataar">
<emphasis id="20E7EAD40B5E0D67FF2CA9703F13FDDF" box="[218,554,546,582]" italics="true" pageId="3" pageNumber="4">Tarbosaurus bataar</emphasis>
<bibRefCitation id="76024B370B5E0D67FDCFA9713E1AFDDE" author="Maleev, E. A." box="[569,803,545,581]" journalOrPublisher="C. R. Acad. Sci. USSR" pageId="3" pageNumber="4" pagination="779 - 783" part="104" refId="ref75585" refString="Maleev, E. A. 1955. [New carnivorous dinosaurs of Mongolia.] C. R. Acad. Sci. USSR 104: 779 - 783. [Russian]" title="New carnivorous dinosaurs of Mongolia." type="journal article" year="1955">
<emphasis id="20E7EAD40B5E0D67FDCFA9713E1AFDDE" bold="true" box="[569,803,545,581]" pageId="3" pageNumber="4">Maleev, 1955</emphasis>
</bibRefCitation>
</taxonomicName>
</paragraph>
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<subSubSection id="5A89654D0B5E0D67FCD4A9723967FCF3" pageId="3" pageNumber="4" type="discussion">
<paragraph id="122C36C60B5E0D67FCD4A9723967FCF3" blockId="3.[172,1232,285,3161]" pageId="3" pageNumber="4">
—This is the best-known non-North American tyrannosaurid, and is known from multiple skeletons from the Nemegt Formation of Mongolia (
<bibRefCitation id="76024B370B5E0D67FBB1A9223C3AFD5E" author="Maleev, E. A." journalOrPublisher="C. R. Acad. Sci. USSR" pageId="3" pageNumber="4" pagination="779 - 783" part="104" refId="ref75585" refString="Maleev, E. A. 1955. [New carnivorous dinosaurs of Mongolia.] C. R. Acad. Sci. USSR 104: 779 - 783. [Russian]" title="New carnivorous dinosaurs of Mongolia." type="journal article" year="1955">Maleev, 1955</bibRefCitation>
, 1974). Most recent estimates place the Nemegt in the Early Maastrichtian. Isolated tyrannosaurid fragments from several Maastrichtian localities in Mongolia and China have been referred to
<taxonomicName id="D5934D450B5E0D67FE85A84D3E87FCDA" authority="(Molnar et al., 1990)" baseAuthorityName="Molnar" baseAuthorityYear="1990" box="[371,958,796,834]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="20E7EAD40B5E0D67FE85A84D3F78FCD8" box="[371,577,798,834]" italics="true" pageId="3" pageNumber="4">Tarbosaurus</emphasis>
(Molnar et
<emphasis id="20E7EAD40B5E0D67FCECA84F3E04FCDA" box="[794,829,796,832]" italics="true" pageId="3" pageNumber="4">al</emphasis>
., 1990)
</taxonomicName>
, but all consideration in this study refers to the Nemegt tyrannosaurid.
</paragraph>
</subSubSection>
<subSubSection id="5A89654D0B5E0D67FF2FA8273942F791" pageId="3" pageNumber="4" type="discussion">
<paragraph id="122C36C60B5E0D67FF2FA8273970F9A0" blockId="3.[172,1232,285,3161]" pageId="3" pageNumber="4">
In its first description,
<taxonomicName id="D5934D450B5E0D67FDA9A8213E14FC0C" box="[607,813,882,918]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="20E7EAD40B5E0D67FDA9A8213E14FC0C" box="[607,813,882,918]" italics="true" pageId="3" pageNumber="4">Tarbosaurus</emphasis>
</taxonomicName>
was called
<taxonomicName id="D5934D450B5E0D67FC08A8233F4EFC5B" authority="(Maleev, 1955)" baseAuthorityName="Maleev" baseAuthorityYear="1955" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="bataar">
<emphasis id="20E7EAD40B5E0D67FC08A8233C5AFC59" italics="true" pageId="3" pageNumber="4">Tyrannosaurus bataar</emphasis>
(
<bibRefCitation id="76024B370B5E0D67FE89A8CD3F52FC5B" author="Maleev, E. A." box="[383,619,925,962]" journalOrPublisher="C. R. Acad. Sci. USSR" pageId="3" pageNumber="4" pagination="779 - 783" part="104" refId="ref75585" refString="Maleev, E. A. 1955. [New carnivorous dinosaurs of Mongolia.] C. R. Acad. Sci. USSR 104: 779 - 783. [Russian]" title="New carnivorous dinosaurs of Mongolia." type="journal article" year="1955">Maleev, 1955</bibRefCitation>
)
</taxonomicName>
.
<taxonomicName id="D5934D450B5E0D67FD66A8CE3E67FC5B" box="[656,862,925,961]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="20E7EAD40B5E0D67FD66A8CE3E67FC5B" box="[656,862,925,961]" italics="true" pageId="3" pageNumber="4">Tarbosaurus</emphasis>
</taxonomicName>
was later erected for a smaller skeleton from the same unit
<emphasis id="20E7EAD40B5E0D67FCCDA89439F3FC70" box="[827,1226,966,1003]" italics="true" pageId="3" pageNumber="4">
(
<taxonomicName id="D5934D450B5E0D67FCB1A8943920FC71" box="[839,1049,967,1003]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Tarbosaurus</taxonomicName>
efremovi),
</emphasis>
and the two were later regarded as synonymous, with
<taxonomicName id="D5934D450B5E0D67FBA8A8BC3C27FBD8" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="20E7EAD40B5E0D67FBA8A8BC3C27FBD8" italics="true" pageId="3" pageNumber="4">Tarbosaurus</emphasis>
</taxonomicName>
being used as the generic name (Maleev, 1974). Maleevs extensive posthumous descriptive work on this animal (1974) applied the name
<emphasis id="20E7EAD40B5E0D67FDA1AF223FD7FB0F" box="[599,750,1137,1173]" italics="true" pageId="3" pageNumber="4">efremovi,</emphasis>
even though
<taxonomicName id="D5934D450B5E0D67FC20AF3C397DFB09" baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[982,1092,1135,1171]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="bataar">
<emphasis id="20E7EAD40B5E0D67FC20AF3C397DFB09" box="[982,1092,1135,1171]" italics="true" pageId="3" pageNumber="4">bataar</emphasis>
</taxonomicName>
has priority. Maleev also described several small tyrannosaurid skeletons, including two he referred to
<emphasis id="20E7EAD40B5E0D67FCF6AF9639F6FB72" box="[768,1231,1220,1257]" italics="true" pageId="3" pageNumber="4">
<taxonomicName id="D5934D450B5E0D67FCF6AF963EECFB73" box="[768,981,1221,1257]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
(G. lancinator
</emphasis>
and
<emphasis id="20E7EAD40B5E0D67FF0FAFA23CEEFA8F" box="[249,471,1265,1301]" italics="true" pageId="3" pageNumber="4">G. novojilovi,</emphasis>
later referred by Carpenter [1992] to the new genus
<emphasis id="20E7EAD40B5E0D67FED7AE483F0AFAA5" box="[289,563,1307,1343]" italics="true" pageId="3" pageNumber="4">
<taxonomicName id="D5934D450B5E0D67FED7AE483F18FAA5" authorityName="Carpenter" authorityYear="1992" box="[289,545,1307,1343]" class="Reptilia" family="Tyrannosauridae" genus="Maleevosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Maleevosaurus</taxonomicName>
),
</emphasis>
but these are now regarded as earlier immature specimens of
<taxonomicName id="D5934D450B5E0D67FDA4AE173EEBFAFD" authority="(Carr, 1999)" baseAuthorityName="Carr" baseAuthorityYear="1999" box="[594,978,1347,1384]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="bataar">
<emphasis id="20E7EAD40B5E0D67FDA4AE173FC9FAF2" box="[594,752,1348,1384]" italics="true" pageId="3" pageNumber="4">T. bataar</emphasis>
(
<bibRefCitation id="76024B370B5E0D67FCFAAE103EFEFAFD" author="Carr, T. D." box="[780,967,1347,1383]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="3" pageNumber="4" pagination="497 - 520" part="19" refId="ref71985" refString="Carr, T. D. 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Theropoda). Journal of Vertebrate Paleontology 19: 497 - 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Theropoda)" type="journal article" year="1999">Carr, 1999</bibRefCitation>
)
</taxonomicName>
. The smallestknown tyrannosaurid skeleton, the type of
<taxonomicName id="D5934D450B5E0D67FC59AE3F3FA0FA26" authority="Dong, 1992" authorityName="Dong" authorityYear="1992" class="Reptilia" family="Tyrannosauridae" genus="Shanshanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="houyanshanensis">
<emphasis id="20E7EAD40B5E0D67FC59AE3F3CF0FA24" italics="true" pageId="3" pageNumber="4">Shanshanosaurus houyanshanensis</emphasis>
Dong, 1992
</taxonomicName>
from western China, may also be an immature example of
<taxonomicName id="D5934D450B5E0D67FD62AE913C1EF98E" authority="(Currie and Dong, 2001 a)" baseAuthorityName="Currie and Dong" baseAuthorityYear="2001" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="20E7EAD40B5E0D67FD62AE913E5BFA7C" box="[660,866,1474,1510]" italics="true" pageId="3" pageNumber="4">Tarbosaurus</emphasis>
(Currie and Dong, 2001a)
</taxonomicName>
. All specimens of
<taxonomicName id="D5934D450B5E0D67FD86AEBD3E06F988" baseAuthorityName="Currie and Dong" baseAuthorityYear="2001" box="[624,831,1518,1554]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="20E7EAD40B5E0D67FD86AEBD3E06F988" box="[624,831,1518,1554]" italics="true" pageId="3" pageNumber="4">Tarbosaurus</emphasis>
</taxonomicName>
examined in this study were mature, and comments will be restricted to them.
</paragraph>
<paragraph id="122C36C60B5E0D67FF2CAD163942F791" blockId="3.[172,1232,285,3161]" pageId="3" pageNumber="4">
This animal is today frequently called
<taxonomicName id="D5934D450B5E0D67FC97AD1239E9F9FE" baseAuthorityName="Carr" baseAuthorityYear="1999" box="[865,1232,1600,1637]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="bataar">
<emphasis id="20E7EAD40B5E0D67FC97AD1239E9F9FE" box="[865,1232,1600,1637]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus bataar</emphasis>
</taxonomicName>
(e.g.,
<bibRefCitation id="76024B370B5E0D67FEEEAD3C3CEBF908" author="Paul, G. S." bookContentInfo="464 pp." box="[280,466,1646,1683]" journalOrPublisher="Simon and Schuster, New York" pageId="3" pageNumber="4" refId="ref76596" refString="Paul, G. S. 1988. Predatory Dinosaurs of the World. Simon and Schuster, New York, 464 pp." title="Predatory Dinosaurs of the World" type="book" year="1988">Paul, 1988</bibRefCitation>
; Carpenter, 1992; Holtz, 1997, 2001a;
<bibRefCitation id="76024B370B5E0D67FB77AD393C30F927" author="Carr, T. D." journalOrPublisher="Journal of Vertebrate Paleontology" pageId="3" pageNumber="4" pagination="497 - 520" part="19" refId="ref71985" refString="Carr, T. D. 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Theropoda). Journal of Vertebrate Paleontology 19: 497 - 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Theropoda)" type="journal article" year="1999">Carr, 1999</bibRefCitation>
), and similarities between the two large Maastrichtian tyrannosaurids are striking. Both have similar postorbital protuberances, and in both the lacrymal “horn” is barely a protuberance at all.
<taxonomicName id="D5934D450B5E0D67FE48AC443FB4F8A1" baseAuthorityName="Currie and Dong" baseAuthorityYear="2001" box="[446,653,1815,1851]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="20E7EAD40B5E0D67FE48AC443FB4F8A1" box="[446,653,1815,1851]" italics="true" pageId="3" pageNumber="4">Tarbosaurus</emphasis>
</taxonomicName>
has more maxillary and dentary alveoli at maturity (13 maxillary alveoli in
<emphasis id="20E7EAD40B5E0D67FC5AAC6C396BF8F8" box="[940,1106,1854,1891]" italics="true" pageId="3" pageNumber="4">
<taxonomicName id="D5934D450B5E0D67FC5AAC6C3975F8F8" baseAuthorityName="Carr" baseAuthorityYear="1999" box="[940,1100,1854,1891]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="bataar">T. bataar</taxonomicName>
,
</emphasis>
as opposed to 12 in
<emphasis id="20E7EAD40B5E0D67FE41AC383F17F815" box="[439,558,1899,1935]" italics="true" pageId="3" pageNumber="4">
<taxonomicName id="D5934D450B5E0D67FE41AC383F25F815" baseAuthorityName="Larson" baseAuthorityYear="1999" box="[439,540,1899,1935]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
),
</emphasis>
but the number of alveoli is smaller in these two than in other mature tyrannosaurids. A close relationship has been presumed in the past (
<figureCitation id="8AA82A430B5E0D67FC98ACEE3EEAF87A" box="[878,979,1980,2017]" captionStart="FIGURE 1" captionStartId="4.[174,306,2624,2656]" captionTargetBox="[268,2234,288,2594]" captionTargetPageId="4" captionText="FIGURE 1. Phylogenetic relationships among theropods and (inset) within Tyrannosauridae as applied herein. Theropod phylogeny based largely on Sereno (1997, 2001) and Norell et al. (2001); uncertain placement of Herrerasauridae reflects information from Langer (2001), and unresolved “Ceratosauria” is based on Carrano and Sampson (1999). Tyrannosauroid phylogeny based on Currie (2000) and Holtz (2001a)." figureDoi="http://doi.org/10.5281/zenodo.3739931" httpUri="https://zenodo.org/record/3739931/files/figure.png" pageId="3" pageNumber="4">Fig. 1</figureCitation>
), a conclusion supported by Holtz (1997, 2001a) phylogenetic analyses.
</paragraph>
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