207 lines
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207 lines
21 KiB
XML
<document ID-DOI="10.5281/zenodo.3245327" ID-GBIF-Dataset="9bf3a9f5-cd36-4be6-bba9-49607815bcd0" ID-Zenodo-Dep="3245327" approvalRequired="3" approvalRequired_for_bibRefs="3" checkinTime="1559900965784" checkinUser="jeremy" docAuthor="Holtz, T. R." docDate="2001" docId="03EE87AEFFA7FF8AFD27F7F489400061" docLanguage="en" docName="Holtz2001TyrannosauridaeGS.pdf.imf" docOrigin="Mesozoic Vertebrate Life, Bloomington: Indiana University Press" docStyle="DocumentStyle{}" docTitle="Tyrannosauridae" docType="treatment" docVersion="16" lastPageNumber="67" masterDocId="FFD7FFD6FFA5FF89FFE1FFFB8A660428" masterDocTitle="The phylogeny and taxonomy of the Tyrannosauridae" masterLastPageNumber="83" masterPageNumber="64" pageNumber="66" updateTime="1645527105248" updateUser="jeremy">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>The phylogeny and taxonomy of the Tyrannosauridae</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Holtz, T. R.</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:originInfo>
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<mods:dateIssued>2001</mods:dateIssued>
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<mods:dateOther type="pubDate">2001-12-31</mods:dateOther>
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<mods:publisher>Indiana University Press</mods:publisher>
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<mods:place>
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<mods:placeTerm>Bloomington</mods:placeTerm>
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</mods:originInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Editor</mods:roleTerm>
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</mods:role>
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<mods:namePart>Tanke D. H.</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Editor</mods:roleTerm>
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</mods:role>
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<mods:namePart>Carpenter K.</mods:namePart>
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</mods:name>
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<mods:titleInfo>
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<mods:title>Mesozoic Vertebrate Life</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:extent unit="page">
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<mods:start>64</mods:start>
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<mods:end>83</mods:end>
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<mods:classification>book chapter</mods:classification>
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<mods:identifier type="DOI">10.5281/zenodo.3245327</mods:identifier>
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<mods:identifier type="GBIF-Dataset">9bf3a9f5-cd36-4be6-bba9-49607815bcd0</mods:identifier>
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<mods:identifier type="Zenodo-Dep">3245327</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.3483016" ID-GBIF-Taxon="157533265" ID-Zenodo-Dep="3483016" LSID="urn:lsid:plazi:treatment:03EE87AEFFA7FF8AFD27F7F489400061" httpUri="http://treatment.plazi.org/id/03EE87AEFFA7FF8AFD27F7F489400061" lastPageId="3" lastPageNumber="67" pageId="2" pageNumber="66">
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<subSubSection pageId="2" pageNumber="66" type="nomenclature">
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<paragraph blockId="2.[634,1840,1157,2650]" pageId="2" pageNumber="66">
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<emphasis box="[710,749,2063,2094]" italics="true" pageId="2" pageNumber="66">As</emphasis>
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in some previous studies (
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<bibRefCitation author="Matthew and Brown" box="[1191,1636,2058,2098]" firstAuthor="Matthew" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="2" pageNumber="66" pagination="367 - 385" part="46" refId="ref5721" refString="Matthew, V. D., and B. Brown. 1922. The family Deinodontidae, with notice of a new genus from the Cretaceous of Alberta. Bulletin of the American Museum of Natural History 46: 367 - 385." title="The family Deinodontidae, with notice of a new genus from the Cretaceous of Alberta" type="journal article" year="1922">Matthew and Brown 1922</bibRefCitation>
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;
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<bibRefCitation author="Olshevsky, G. & T. L. Ford & S. Yamamoto" journalOrPublisher="Kyoryugaku saizensen" pageId="2" pageNumber="66" pagination="92 - 119" part="9" refId="ref5959" refString="Olshevsky, G., T. L. Ford, and S. Yamamoto. 1995 a. [The origin and evolution of the tyrannosaurids, part 1]. Kyoryugaku saizensen 9: 92 - 119. (In Japanese.)" title="The origin and evolution of the tyrannosaurids, part 1" type="journal article" year="1995">Olshevsky et al.. 1995a</bibRefCitation>
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,
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<bibRefCitation author="Olshevsky G. & T. L. Ford & S. Yamamaoto" box="[857,878,2106,2144]" journalOrPublisher="Kyoryugaku saizensen" pageId="2" pageNumber="75" pagination="75 - 99" part="10" refId="ref6004" title="The origin and evolution of the tyrannosaurids, part 2" type="journal article" year="1995">b</bibRefCitation>
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;
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<bibRefCitation author="Currie, P. J." box="[904,1165,2108,2147]" editor="M. Benton & E. Kurochkin & M. Shiskin & D. Unwin" journalOrPublisher="Cambridge Univer- sity Press" pageId="2" pageNumber="66" refId="ref4795" refString="Currie, P. J. in press. Theropods from the Cretaceous of Mongolia. In M. Benton, E. Kurochkin, M. Shiskin, and D. Unwin (eds.), The Age of Dinosaurs in Russia and Mongolia. Cambridge: Cambridge Univer- sity Press." title="Theropods from the Cretaceous of Mongolia" type="book chapter" volumeTitle="The Age of Dinosaurs in Russia and Mongolia" year="in press">Currie in press</bibRefCitation>
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),
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<taxonomicName authority="Osborn, 1906" box="[1200,1481,2112,2147]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="3" lastPageNumber="67" pageId="2" pageNumber="66" type="discussion">
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<paragraph blockId="2.[634,1840,1157,2650]" lastBlockId="3.[191,1394,157,2652]" lastPageId="3" lastPageNumber="67" pageId="2" pageNumber="66">
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was found to comprise two clades:
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[961,1254,2160,2197]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="subFamily" subFamily="Aublysodontinae">Aublysodontinae</taxonomicName>
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and
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<taxonomicName authorityName="Matthew & Brown" authorityYear="1922" box="[1359,1662,2161,2198]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
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. Potential phylogenetic taxonomic definitions for the clades in question might be:
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<taxonomicName box="[635,921,2260,2295]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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, all descendants of the most recent common ancestor of
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[680,930,2311,2346]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="genus">
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<emphasis box="[680,930,2311,2346]" italics="true" pageId="2" pageNumber="66">Tyrannosaurus</emphasis>
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</taxonomicName>
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and
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[1014,1213,2312,2347]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="genus">
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<emphasis box="[1014,1213,2312,2347]" italics="true" pageId="2" pageNumber="66">Aublysodon</emphasis>
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</taxonomicName>
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;
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[1242,1524,2312,2347]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="subFamily" subFamily="Aublysodontinae">Aublysodontinae</taxonomicName>
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,
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[1554,1764,2313,2350]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="genus">
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<emphasis box="[1554,1764,2313,2350]" italics="true" pageId="2" pageNumber="66">Aublysodon</emphasis>
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</taxonomicName>
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and all taxa sharing a more recent common ancestor with it than with
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[637,879,2413,2446]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="genus">
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<emphasis box="[637,879,2413,2446]" italics="true" pageId="2" pageNumber="66">Tyrannosaurus</emphasis>
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</taxonomicName>
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; and
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<taxonomicName authorityName="Matthew & Brown" authorityYear="1922" box="[980,1270,2409,2447]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
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,
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1288,1537,2414,2448]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="genus">
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<emphasis box="[1288,1537,2414,2448]" italics="true" pageId="2" pageNumber="66">Tyrannosaurus</emphasis>
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</taxonomicName>
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and all taxa sharing a more recent common ancestor with it than with
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[1614,1827,2461,2498]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="genus">
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<emphasis box="[1614,1827,2461,2498]" italics="true" pageId="2" pageNumber="66">Aublysodon</emphasis>
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</taxonomicName>
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. However, these definitions must be provisional, as the type species of
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[634,846,2561,2598]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="genus">
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<emphasis box="[634,846,2561,2598]" italics="true" pageId="2" pageNumber="66">Aublysodon</emphasis>
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</taxonomicName>
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,
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[868,1089,2559,2598]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="species" species="mirandus">
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<emphasis box="[868,902,2559,2598]" italics="true" pageId="2" pageNumber="66">A</emphasis>
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.
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<emphasis box="[922,1089,2561,2598]" italics="true" pageId="2" pageNumber="66">mirandus</emphasis>
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</taxonomicName>
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, is known only from isolated premaxillary teeth, while the somewhat more complete
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<taxonomicName authorityName="Paul" authorityYear="1988" box="[1381,1573,2609,2649]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="66" phylum="Chordata" rank="species" species="molnari">
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<emphasis box="[1381,1417,2609,2649]" italics="true" pageId="2" pageNumber="66">A</emphasis>
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.
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<emphasis box="[1436,1573,2610,2649]" italics="true" pageId="2" pageNumber="66">molnari</emphasis>
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</taxonomicName>
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is known only from skull elements and may eventually prove to be a different genus. Similar problems would result from using
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<taxonomicName authorityName="Gilmore" authorityYear="1933" box="[942,1184,208,245]" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="genus">
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<emphasis box="[942,1184,208,245]" italics="true" pageId="3" pageNumber="67">Alectrosaurus</emphasis>
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</taxonomicName>
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rather than
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[192,399,257,294]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="genus">
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<emphasis box="[192,399,257,294]" italics="true" pageId="3" pageNumber="67">Aublysodon</emphasis>
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</taxonomicName>
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as the anchor taxon for Aublysodontrnae.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="3" pageNumber="67" type="discussion">
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<paragraph blockId="3.[191,1394,157,2652]" pageId="3" pageNumber="67">
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(Note that the phylogenetic taxonomy proposed by
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<bibRefCitation author="Sereno" box="[1154,1386,312,352]" firstAuthor="Sereno" journalOrPublisher="Neues Jahrbuch fiir Geologie und Pakiontologie Abhandlungen" pageId="3" pageNumber="67" pagination="4 t- 83" part="210" refId="ref6304" refString="Sereno, P. C. 1998. A rationale for phylogenetic definitions with applica- tion to the higher-level taxonomy of Dinosauria. Neues Jahrbuch fiir Geologie und Pakiontologie Abhandlungen 210: 4 t- 83." title="A rationale for phylogenetic definitions with applica- tion to the higher-level taxonomy of Dinosauria" type="journal article" year="1998">Sereno [1998]</bibRefCitation>
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is problematic as well: his "
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<taxonomicName box="[689,988,359,396]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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" is defined as all taxa closer to
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[343,593,409,444]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="genus">
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<emphasis box="[343,593,409,444]" italics="true" pageId="3" pageNumber="67">Tyrannosaurus</emphasis>
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</taxonomicName>
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thanto
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<taxonomicName authorityName="Gilmore" authorityYear="1933" box="[729,963,409,444]" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="genus">
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<emphasis box="[729,963,409,444]" italics="true" pageId="3" pageNumber="67">Alectrosaurus</emphasis>
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</taxonomicName>
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,
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[985,1185,410,445]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="genus">
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<emphasis box="[985,1185,410,445]" italics="true" pageId="3" pageNumber="67">Aublysodon</emphasis>
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</taxonomicName>
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, and
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<taxonomicName authorityName="Bakker, Williams & Currie" authorityYear="1988" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="genus">
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<emphasis box="[1283,1385,412,445]" italics="true" pageId="3" pageNumber="67">
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<emphasis box="[193,351,458,495]" italics="true" pageId="3" pageNumber="67">Nanotyrannus</emphasis>
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</emphasis>
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</taxonomicName>
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. The latter specimen is very likely a juvenile
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<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="species" species="rex">
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<emphasis italics="true" pageId="3" pageNumber="67">Tyrannosaunts rex</emphasis>
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</taxonomicName>
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[
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<bibRefCitation author="Carr" box="[268,453,506,544]" firstAuthor="Carr" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="3" pageNumber="67" pagination="497 - 520" part="19" refId="ref4655" refString="Carr, T. D. 1999. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). Journal of Vertebrate Paleontology 19: 497 - 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">
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Carr
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<emphasis box="[365,453,509,544]" italics="true" pageId="3" pageNumber="67">1999</emphasis>
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</bibRefCitation>
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; see also below], rendering his "
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<taxonomicName box="[999,1298,510,547]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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" as a subgroup within the species
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[674,767,560,595]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="71" phylum="Chordata" rank="species" species="rex">
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<emphasis box="[674,767,560,595]" italics="true" pageId="3" pageNumber="67">T. rex</emphasis>
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</taxonomicName>
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[all specimens sharing a more recent common ancestor with the type specimen of
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[941,1030,613,646]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="71" phylum="Chordata" rank="species" species="rex">
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<emphasis box="[941,1030,613,646]" italics="true" pageId="3" pageNumber="67">T. rex</emphasis>
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</taxonomicName>
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than with the "type " of
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<taxonomicName authorityName="Bakker, Williams & Currie" authorityYear="1988" box="[240,503,657,702]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="genus">
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<emphasis box="[240,503,657,702]" italics="true" pageId="3" pageNumber="67">Nanotyrannus]</emphasis>
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</taxonomicName>
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. Furthermore,
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<bibRefCitation author="Sereno's" box="[768,1023,657,702]" firstAuthor="Sereno's" journalOrPublisher="Neues Jahrbuch fiir Geologie und Pakiontologie Abhandlungen" pageId="3" pageNumber="67" pagination="4 t- 83" part="210" refId="ref6304" refString="Sereno, P. C. 1998. A rationale for phylogenetic definitions with applica- tion to the higher-level taxonomy of Dinosauria. Neues Jahrbuch fiir Geologie und Pakiontologie Abhandlungen 210: 4 t- 83." title="A rationale for phylogenetic definitions with applica- tion to the higher-level taxonomy of Dinosauria" type="journal article" year="1998">Sereno's [1998</bibRefCitation>
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] "
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<taxonomicName authorityName="Matthew & Brown" authorityYear="1922" box="[1069,1355,660,695]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
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, " all taxa closer to
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[524,788,709,746]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="genus">
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<emphasis box="[524,788,709,746]" italics="true" pageId="3" pageNumber="67">Tyrannosaurus</emphasis>
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</taxonomicName>
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than to
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[950,1199,709,746]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="genus">
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<emphasis box="[950,1199,709,746]" italics="true" pageId="3" pageNumber="67">Albertosaurus</emphasis>
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</taxonomicName>
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,
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<taxonomicName pageId="3" pageNumber="67">
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<emphasis box="[192,306,761,796]" italics="true" pageId="3" pageNumber="67">Daspletosaurus</emphasis>
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</taxonomicName>
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, or
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<taxonomicName authorityName="Lambe" authorityYear="1914" box="[375,602,761,796]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="genus">
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<emphasis box="[375,602,761,796]" italics="true" pageId="3" pageNumber="67">Gorgosaurus</emphasis>
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</taxonomicName>
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, would be limited to the genus
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1138,1380,763,796]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="genus">
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<emphasis box="[1138,1380,763,796]" italics="true" pageId="3" pageNumber="67">Tyrannosaurus</emphasis>
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</taxonomicName>
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itself foilowing the phylogeny presented here. It is recommended that until such time as the more complete Mongolian specimens currently referred to
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<taxonomicName box="[388,739,911,946]" class="Reptilia" family="Not" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="67" phylum="Chordata" rank="species" species="olseni">
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<emphasis box="[388,739,911,946]" italics="true" pageId="3" pageNumber="67">Alectrosaurus olseni</emphasis>
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</taxonomicName>
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[see below], which seem to represent the best materiai of primitive tyrannosaur, are more adequately described, that the provisional phylogenetic taxonomy proposed here be used for tyrannosaurid systematics.)
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</paragraph>
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</subSubSection>
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</treatment>
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</document> |