196 lines
20 KiB
XML
196 lines
20 KiB
XML
<document ID-DOI="10.5281/zenodo.3374526" ID-GBIF-Dataset="2a40cf95-9ffb-4f3c-894b-cf53b8788a0a" ID-Zenodo-Dep="3374526" approvalRequired="109" approvalRequired_for_document="1" approvalRequired_for_illustrations="43" approvalRequired_for_originalDoi="1" approvalRequired_for_taxonomicNames="38" approvalRequired_for_textStreams="1" approvalRequired_for_treatments="25" checkinTime="1566477704804" checkinUser="jeremy" docAuthor="Holtz, TR jr." docDate="2004" docId="012B87EDFF8AD81B4DBE2022D98BB4FA" docLanguage="en" docName="Holtz2004Tyrannosauroidea.pdf" docOrigin="The Dinosauria, University of California Press" docStyle="DocumentStyle{}" docTitle="Tyrannosauridae Osborn 1905" docType="treatment" docVersion="9" lastPageNumber="7" masterDocId="FD12FF95FF9AD80B4D3D2775DD29B37C" masterDocTitle="Tyrannosauroidea" masterLastPageNumber="136" masterPageNumber="111" pageNumber="7" updateTime="1656386983924" updateUser="felipe">
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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>Tyrannosauroidea</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Holtz, TR jr.</mods:namePart>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:dateIssued>2004</mods:dateIssued>
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<mods:publisher>University of California Press</mods:publisher>
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<mods:name type="personal">
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<mods:roleTerm>Editor</mods:roleTerm>
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<mods:namePart>Weishampel DB</mods:namePart>
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<mods:roleTerm>Editor</mods:roleTerm>
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<mods:namePart>Dodson P</mods:namePart>
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<mods:name type="personal">
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<mods:roleTerm>Editor</mods:roleTerm>
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<mods:namePart>Osmolska H</mods:namePart>
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<mods:titleInfo>
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<mods:title>The Dinosauria</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>111</mods:start>
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<mods:classification>book chapter</mods:classification>
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<mods:identifier type="DOI">10.5281/zenodo.3374526</mods:identifier>
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<mods:identifier type="GBIF-Dataset">2a40cf95-9ffb-4f3c-894b-cf53b8788a0a</mods:identifier>
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<mods:identifier type="Zenodo-Dep">3374526</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.3483196" ID-GBIF-Taxon="163642030" ID-Zenodo-Dep="3483196" LSID="urn:lsid:plazi:treatment:012B87EDFF8AD81B4DBE2022D98BB4FA" httpUri="http://treatment.plazi.org/id/012B87EDFF8AD81B4DBE2022D98BB4FA" lastPageNumber="7" pageId="16" pageNumber="7">
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<subSubSection box="[131,560,1879,1898]" pageId="16" pageNumber="7" type="nomenclature">
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<paragraph blockId="16.[99,716,1420,1926]" box="[131,560,1879,1898]" pageId="16" pageNumber="7">
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Within Tyrannosauroidea,
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[394,560,1879,1898]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="16" pageNumber="7" type="discussion">
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<paragraph blockId="16.[99,716,1420,1926]" lastBlockId="16.[747,1364,100,1926]" pageId="16" pageNumber="7">
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has been phylogenetically defined, although this definition has been rendered problematic owing to the nature of some of the putative terminal taxa (Brochu 2002). Sereno (1998) defined this taxon as all tyrannosauroids closer to
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1007,1145,159,177]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">
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<emphasis box="[1007,1145,159,177]" italics="true" pageId="16" pageNumber="7">Tyrannosaurus</emphasis>
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</taxonomicName>
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than to
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<emphasis italics="true" pageId="16" pageNumber="7">
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<taxonomicName authorityName="Gilmore" authorityYear="1933" box="[1233,1360,158,177]" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">Alectrosaurus</taxonomicName>
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,
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[747,858,187,206]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">Aublysodon</taxonomicName>
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,
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</emphasis>
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or
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<emphasis box="[891,1030,187,205]" italics="true" pageId="16" pageNumber="7">
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<taxonomicName authorityName="Bakker, Williams & Currie" authorityYear="1988" box="[891,1025,187,205]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">Nanotyrannus</taxonomicName>
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,
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</emphasis>
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and Holtz (2001b) defined it as all descendants of the most recent common ancestor of
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<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">
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<emphasis italics="true" pageId="16" pageNumber="7">Tyrannosaurus</emphasis>
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</taxonomicName>
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and
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<emphasis box="[865,980,244,263]" italics="true" pageId="16" pageNumber="7">
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[865,976,244,263]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">Aublysodon</taxonomicName>
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.
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</emphasis>
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However,
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<taxonomicName authorityName="Leidy" authorityYear="1868" box="[1090,1198,244,263]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">
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<emphasis box="[1090,1198,244,263]" italics="true" pageId="16" pageNumber="7">Aublysodon</emphasis>
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</taxonomicName>
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is based on indeterminate (and lost) material, and the derived feature previously claimed for it—the lack of serrations on the teeth—may be due to postmortem damage rather than actual anatomy (Brochu 2002; Carr and Williamson, in press). Furthermore,
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<taxonomicName baseAuthorityName="Gilmore" baseAuthorityYear="1946" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="lancensis">
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<emphasis italics="true" pageId="16" pageNumber="7">Nanotyrannus lancensis</emphasis>
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</taxonomicName>
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is likely a juvenile
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1128,1302,388,406]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="rex">
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<emphasis box="[1128,1302,388,406]" italics="true" pageId="16" pageNumber="7">Tyrannosaurus rex</emphasis>
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</taxonomicName>
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(Carr 1999; Holtz 2001b; Brochu 2002; Carr and Williamson, in press); and even if it is not, it is almost certainly from a taxon more closely related to
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[913,961,474,492]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="rex">
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<emphasis box="[913,961,474,492]" italics="true" pageId="16" pageNumber="7">T. rex</emphasis>
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</taxonomicName>
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than to all other taxa previously regarded as tyrannosaurids (Holtz 2001b; Currie 2003b). Finally,
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<taxonomicName authorityName="Gilmore" authorityYear="1933" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">
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<emphasis italics="true" pageId="16" pageNumber="7">Alectrosaurus</emphasis>
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</taxonomicName>
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may be a
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<emphasis box="[930,1069,531,550]" italics="true" pageId="16" pageNumber="7">nomen dubium</emphasis>
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(Carr et al., in press), and due to its incompleteness it is phylogenetically labile (Currie 2003b).
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</paragraph>
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<paragraph blockId="16.[747,1364,100,1926]" pageId="16" pageNumber="7">
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Here,
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[833,995,588,607]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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is defined as comprising
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<emphasis italics="true" pageId="16" pageNumber="7">
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<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
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,
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<taxonomicName baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[784,960,617,636]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="bataar">Tarbosaurus bataar</taxonomicName>
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,
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<taxonomicName authorityName="Russell" authorityYear="1970" box="[966,1175,617,636]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="torosus">Daspletosaurus torosus</taxonomicName>
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,
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<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="sarcophagus">Albertosaurus sarcophagus</taxonomicName>
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,
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<taxonomicName authorityName="Lambe" authorityYear="1914" box="[825,1018,645,665]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="libratus">Gorgosaurus libratus</taxonomicName>
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,
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</emphasis>
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their most recent common ancestor, and all of its descendants. Furthermore, two subclades can be defined:
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<taxonomicName authorityName="Matthew et Brown" authorityYear="1922" box="[863,1032,702,721]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
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, a stem-based taxon composed of
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[747,918,732,750]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="rex">
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<emphasis box="[747,918,732,750]" italics="true" pageId="16" pageNumber="7">Tyrannosaurus rex</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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<emphasis box="[996,1239,760,779]" italics="true" pageId="16" pageNumber="7">
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[996,1234,760,779]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="sarcophagus">Albertosaurus sarcophagus</taxonomicName>
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;
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</emphasis>
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and
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<taxonomicName class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="subFamily" subFamily="Albertosaurinae">Albertosaurinae</taxonomicName>
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, a stem-based taxon composed of
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1157,1294,789,808]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="sarcophagus">
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<emphasis box="[1157,1294,789,808]" italics="true" pageId="16" pageNumber="7">A. sarcophagus</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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<emphasis box="[747,805,847,865]" italics="true" pageId="16" pageNumber="7">
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[747,801,847,865]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
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.
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</emphasis>
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</paragraph>
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<paragraph blockId="16.[747,1364,100,1926]" pageId="16" pageNumber="7">
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The precise position of
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1003,1171,874,893]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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, and by implication of Tyrannosauroidea, within
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<taxonomicName authorityName="Huene" authorityYear="1914" box="[1033,1171,903,922]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Avetheropoda" pageId="16" pageNumber="7" phylum="Chordata" rank="subOrder" subOrder="Coelurosauria">Coelurosauria</taxonomicName>
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is uncertain. Maniraptoriformes was defined (Holtz 1996a) as all descendants of the most recent common ancestor of
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<taxonomicName authorityName="Marsh" authorityYear="1890" box="[1104,1240,961,980]" class="Reptilia" family="Ornithomimidae" genus="Ornithomimus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">
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<emphasis box="[1104,1240,961,980]" italics="true" pageId="16" pageNumber="7">Ornithomimus</emphasis>
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</taxonomicName>
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and modern birds. Arctometatarsalia comprises those taxa closer to
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<taxonomicName authorityName="Marsh" authorityYear="1890" class="Reptilia" family="Ornithomimidae" genus="Ornithomimus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">
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<emphasis italics="true" pageId="16" pageNumber="7">Ornithomimus</emphasis>
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</taxonomicName>
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than to modern birds, while
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<taxonomicName authorityName="Marsh" authorityYear="1890" box="[1120,1256,1018,1037]" class="Reptilia" family="Ornithomimidae" genus="Ornithomimus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">
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<emphasis box="[1120,1256,1018,1037]" italics="true" pageId="16" pageNumber="7">Ornithomimus</emphasis>
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</taxonomicName>
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comprises those closer to modern birds than to Maniraptora (Holtz 2001a). The three main hypotheses of tyrannosaurid relationships within
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<taxonomicName authorityName="Huene" authorityYear="1914" box="[747,885,1104,1123]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Avetheropoda" pageId="16" pageNumber="7" phylum="Chordata" rank="subOrder" subOrder="Coelurosauria">Coelurosauria</taxonomicName>
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are (1)
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[956,1122,1104,1123]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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is a coelurosaurian clade outside Maniraptoriformes (Pérez-Moreno et al. 1994; Forster et al. 1998; Makovicky and Sues 1998; Norell et al. 2000; Rauhut 2003); (2)
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[844,1010,1190,1209]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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lies within Arctometatarsalia (Pérez-Moreno et al. 1993; Holtz 1994, 1998a, 2001a); and (3)
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<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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lies within Maniraptora (Sereno 1999a). Novas (1992b) and Wagner and Gauthier (1999) presented
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1173,1339,1276,1295]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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as one branch of an unresolved tritomy with Ornithomimosauria and Maniraptora; this would also be the strict consensus of the three hypotheses presented above.
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</paragraph>
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<paragraph blockId="16.[747,1364,100,1926]" pageId="16" pageNumber="7">
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The phylogenetic analysis presented here is the same as that presented in Holtz et al. (this vol.). This analysis comprises 76 taxa (with
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<taxonomicName baseAuthorityName="Reig" baseAuthorityYear="1963" box="[857,1013,1448,1467]" class="Reptilia" family="Herrerasauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="family">Herrerasauridae</taxonomicName>
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as an outgroup) and 638 morphological characters. DELTRAN optimization was selected for purposes of discussing the distribution of derived character states. See Holtz et al. (this vol.) for more details of the methodology and the structure of the most parsimonious trees beyond the portion examined here.
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</paragraph>
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<paragraph blockId="16.[747,1364,100,1926]" pageId="16" pageNumber="7">
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<figureCitation box="[779,889,1619,1639]" pageId="16" pageNumber="7" targetBox="[594,1360,106,542]" targetPageId="17">Figure 5.22</figureCitation>
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represents the Tyrannosauroidea portion of the 2,5 4 4 most parsimonious trees. These trees have a length of 2,444, CI = 0.354, RI = 0.715, RCI = 0.253, and HI = 0.662. In this analysis Tyrannosauroidea was found to lie outside of the Maniraptora-Ornithomimosauria clade Maniraptoriformes, but it was united with these taxa above the
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<taxonomicName authorityName="Cope" authorityYear="1871" box="[1138,1319,1763,1782]" class="Reptilia" family="Compsognathidae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="family">Compsognathidae</taxonomicName>
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in a clade that meets Sereno’s (1999a) definition of Tyrannoraptora (i.e.,
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<emphasis box="[795,937,1821,1839]" italics="true" pageId="16" pageNumber="7">
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[795,933,1821,1839]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="7" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
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,
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</emphasis>
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modern birds, their most recent common ancestor, and all of its descendants).
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</paragraph>
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<paragraph blockId="16.[747,1364,100,1926]" pageId="16" pageNumber="7">The taxa within Tyrannosauroidea are discussed here from more basal forms to those higher on the tree.</paragraph>
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</subSubSection>
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</treatment>
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</document> |