365 lines
55 KiB
XML
365 lines
55 KiB
XML
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<mods:title id="F16CD4A18288428856881917F9F91D03">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:name id="B281FB4F29CEBABC5FBF46CEBD0B96CA" type="personal">
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<mods:namePart id="30EDC494D7FB591EC039B18F704238CB">Christopher A. Brochu</mods:namePart>
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<mods:affiliation id="00D6ED8D2EEBDC2DA0BCA32FBB42A0C4">Department of Geology, Field Museum, 1400 S. Lake Shore Drive, Chicago, Illinois 60605</mods:affiliation>
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<mods:title id="767EAAF1D6E5BB620D4FCFEB6F10422C">Journal of Vertebrate Paleontology</mods:title>
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<mods:part id="6B49516FFBFF0C5C5283CC3C5E8C7B82">
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<mods:date id="2D606DE4A22A0E60BACC347813D39211">2003</mods:date>
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<mods:number id="5F8338CA1BA23F63EAE6075470BB01FB">2003-12-31</mods:number>
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<mods:number id="57204E26273807B70A6E6A0A62B7BEC1">22</mods:number>
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<mods:identifier id="79100C250F279AC53D1D6F4C8A3473AB" type="DOI">10.2307/3889334</mods:identifier>
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<treatment id="9A3A87D00B5C0D66FF0DAF943C42FA7B" ID-DOI="http://doi.org/10.5281/zenodo.3810809" ID-GBIF-Taxon="190303340" ID-Zenodo-Dep="3810809" LSID="urn:lsid:plazi:treatment:9A3A87D00B5C0D66FF0DAF943C42FA7B" httpUri="http://treatment.plazi.org/id/9A3A87D00B5C0D66FF0DAF943C42FA7B" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">
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<subSubSection id="5A89654D0B5C0D65FF0DAF9439B5FB71" box="[251,1164,1223,1259]" pageId="1" pageNumber="2" type="description">
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<paragraph id="122C36C60B5C0D65FF0DAF9439B5FB71" blockId="1.[251,1164,1222,1259]" box="[251,1164,1223,1259]" pageId="1" pageNumber="2">
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SYSTEMATIC REVIEW OF
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<taxonomicName id="D5934D450B5C0D65FD03AF9439B5FB71" authorityName="sensu Sereno" authorityYear="1998" box="[757,1164,1223,1259]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">TYRANNOSAURIDAE</taxonomicName>
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</paragraph>
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</subSubSection>
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<subSubSection id="5A89654D0B5C0D65FF2AAE433930F7B6" pageId="1" pageNumber="2" type="nomenclature">
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<paragraph id="122C36C60B5C0D65FF2AAE433930F7B6" blockId="1.[170,1234,1295,3150]" pageId="1" pageNumber="2">
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<taxonomicName id="D5934D450B5C0D65FF2AAE433CD4FAAE" box="[220,493,1296,1332]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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is an undoubtedly monophyletic assemblage of theropods. It is best represented in the Late Cretaceous of western North America and eastern Asia, with possible occurrences elsewhere in space and time (
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<bibRefCitation id="76024B370B5C0D65FCE0AEDE3ED8FA2B" author="Leidy, J." box="[790,993,1421,1457]" journalOrPublisher="Proceedings of the Academy of Natural Sciences of Philadelphia" pageId="1" pageNumber="2" pagination="72 - 73" part="8" refId="ref75212" refString="Leidy, J. 1856. Notice of remains of extinct Reptiles and Fishes discovered by Dr. F. V. Hayden in the Bad Lands of the Judith River, Nebraska Territory. Proceedings of the Academy of Natural Sciences of Philadelphia 8: 72 - 73." title="Notice of remains of extinct Reptiles and Fishes discovered by Dr. F. V. Hayden in the Bad Lands of the Judith River, Nebraska Territory" type="journal article" year="1856">Leidy, 1856</bibRefCitation>
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;
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<bibRefCitation id="76024B370B5C0D65FC00AEDE39F1FA2B" author="Marsh, O. C." box="[1014,1224,1421,1457]" journalOrPublisher="American Journal of Science (Series 3)" pageId="1" pageNumber="2" pagination="449 - 453" part="43" refId="ref75694" refString="Marsh, O. C. 1892. Notice of new reptiles from the Laramie Formation. American Journal of Science (Series 3) 43: 449 - 453." title="Notice of new reptiles from the Laramie Formation" type="journal article" year="1892">Marsh, 1892</bibRefCitation>
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;
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<bibRefCitation id="76024B370B5C0D65FF42AEE43C41FA41" author="Cope, E. D." box="[180,376,1463,1499]" journalOrPublisher="American Naturalist" pageId="1" pageNumber="2" pagination="756 - 758" part="26" refId="ref72823" refString="Cope, E. D. 1892. Fourth note on the Dinosauria of the Laramie. American Naturalist 26: 756 - 758." title="Fourth note on the Dinosauria of the Laramie" type="journal article" year="1892">Cope, 1892</bibRefCitation>
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; Lambe, 1904, 1914b, 1917;
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<bibRefCitation id="76024B370B5C0D65FC83AEE43965FA41" author="Maleev, E. A." box="[885,1116,1463,1499]" journalOrPublisher="C. R. Acad. Sci. USSR" pageId="1" pageNumber="2" 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>
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, 1974; Gilmore, 1933; Kurzanov, 1976b;
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<bibRefCitation id="76024B370B5C0D65FD0DAEB13ED0F99C" author="Lawson, D. A." box="[763,1001,1506,1542]" journalOrPublisher="Journal of Paleontology" pageId="1" pageNumber="2" pagination="158 - 164" part="50" refId="ref75087" refString="Lawson, D. A. 1976. Tyrannosaurus and Torvosaurus, Maestrichtian dinosaurs from trans-Pecos Texas. Journal of Paleontology 50: 158 - 164." title="Tyrannosaurus and Torvosaurus, Maestrichtian dinosaurs from trans-Pecos Texas" type="journal article" year="1976">Lawson, 1976</bibRefCitation>
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;
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<bibRefCitation id="76024B370B5C0D65FBF7AEB139FEF99C" author="Dong, Z. - M." box="[1025,1223,1506,1542]" journalOrPublisher="Vertebrata PalAsiatica" pageId="1" pageNumber="2" pagination="59 - 66" part="15" refId="ref73312" refString="Dong, Z. - M. 1977. On the dinosaurian remains from Turpan, Zinjiang. Vertebrata PalAsiatica 15: 59 - 66." title="On the dinosaurian remains from Turpan, Zinjiang" type="journal article" year="1977">Dong, 1977</bibRefCitation>
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, 1979; Carpenter, 1982;
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<bibRefCitation id="76024B370B5C0D65FDBCAD5F3EDCF9AA" author="Schwimmer, D. R. & G. D. Williams & J. L. Dobie & W. G. Siesser" box="[586,997,1548,1584]" journalOrPublisher="Journal of Paleontology" pageId="1" pageNumber="2" pagination="288 - 296" part="67" refId="ref77732" refString="Schwimmer, D. R., G. D. Williams, J. L. Dobie, and W. G. Siesser. 1993. Late Cretaceous dinosaurs from the Blufftown Formation in western Georgia and eastern Alabama. Journal of Paleontology 67: 288 - 296." title="Late Cretaceous dinosaurs from the Blufftown Formation in western Georgia and eastern Alabama" type="journal article" year="1993">Schwimmer et al, 1993</bibRefCitation>
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;
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<bibRefCitation id="76024B370B5C0D65FC09AD5F3C87F9C1" author="Lehman, T. M. & K. Carpenter" journalOrPublisher="Journal of Paleontology" pageId="1" pageNumber="2" pagination="1026 - 1032" part="64" refId="ref75173" refString="Lehman, T. M., and K. Carpenter. 1990. A partial skeleton of the tyrannosaurid dinosaur Aublysodon from the Upper Cretaceous of New Mexico. Journal of Paleontology 64: 1026 - 1032." title="A partial skeleton of the tyrannosaurid dinosaur Aublysodon from the Upper Cretaceous of New Mexico" type="journal article" year="1990">Lehman and Carpenter, 1990</bibRefCitation>
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;
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<bibRefCitation id="76024B370B5C0D65FE22AD643E33F9C1" author="Currie, P. J. & J. K. Rigby & R. E. Sloan" box="[468,778,1591,1627]" editor="K. Carpenter & P. J. Currie" journalOrPublisher="Cambridge University Press, Cambridge" pageId="1" pageNumber="2" pagination="107 - 125" refId="ref72848" refString="Currie, P. J., J. K. Rigby, and R. E. Sloan. 1990. Theropod teeth from the Judith River Formation; pp. 107 - 125 in K. Carpenter and P. J. Currie (eds.), Dinosaur Systematics, Cambridge University Press, Cambridge." title="Theropod teeth from the Judith River Formation" type="book chapter" volumeTitle="Dinosaur Systematics" year="1990">Currie et al., 1990</bibRefCitation>
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;
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<bibRefCitation id="76024B370B5C0D65FCD6AD643913F9C1" author="Carpenter, K." box="[800,1066,1591,1627]" editor="K. Carpenter & P. J. Currie" journalOrPublisher="Cambridge University Press, New York" pageId="1" pageNumber="2" pagination="141 - 145" refId="ref71853" refString="Carpenter, K. 1990. Variation in Tyrannosaurus rex; pp. 141 - 145 in K. Carpenter and P. J. Currie (eds.), Dinosaur Systematics. Cambridge University Press, New York." title="Variation in Tyrannosaurus rex" type="book chapter" volumeTitle="Dinosaur Systematics" year="1990">Carpenter, 1990</bibRefCitation>
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; Carr and Williamson, 2000;
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<bibRefCitation id="76024B370B5C0D65FE03AD313E37F91C" author="Hutt, S. & D. Naish & D. M. Martill & M. J. Barker & P. Newbery" box="[501,782,1634,1670]" journalOrPublisher="Cretaceous Research" pageId="1" pageNumber="2" pagination="227 - 242" part="22" refId="ref74512" refString="Hutt, S., D. Naish, D. M. Martill, M. J. Barker, and P. Newbery. 2001. A preliminary account of a new tyrannosauroid theropod from the Wessex Formation (Early Cretaceous) of southern England. Cretaceous Research 22: 227 - 242." title="A preliminary account of a new tyrannosauroid theropod from the Wessex Formation (Early Cretaceous) of southern England" type="journal article" year="2001">Hutt et al., 2001</bibRefCitation>
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; Schwimmer and Kiernan, 2001; Carr et al., in press). Members of the group share several derived character states, including D-shaped incisiform premaxillary teeth; a squamosal recess; a quadratojugal-squamosal flange bisecting the infratemporal fenestra; an enlarged surangular foramen; a reduced retroarticular process; contact between the lacrymal and postorbital over the orbit; a functionally didactyl hand; an enlarged lesser trochanter on the femur; and an arctometatarsalian pes (see Holtz, 2001a for review). These are among the most popular dinosaurs in the public and were surely among the dominant predators of their ecosystems.
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</paragraph>
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</subSubSection>
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<subSubSection id="5A89654D0B5C0D65FF2CA3603BE7FCF9" pageId="1" pageNumber="2" type="description">
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<paragraph id="122C36C60B5C0D65FF2CA3603977F5C9" blockId="1.[170,1234,1295,3150]" pageId="1" pageNumber="2">
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Given the popularity of tyrannosaurids in both academic and lay circles, one would expect
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<taxonomicName id="D5934D450B5C0D65FD50A30D3E8FF718" authorityName="sensu Sereno" authorityYear="1998" box="[678,950,2142,2178]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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to be one of the best-characterized dinosaurian groups. In fact, there is considerable debate concerning virtually any aspect of tyrannosaurid systematics, from group membership and diagnosis to the position of
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<taxonomicName id="D5934D450B5C0D65FEA1A25B3F5EF6B6" authorityName="sensu Sereno" authorityYear="1998" box="[343,615,2312,2348]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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within Theropoda. Small tyrannosaurs might be dwarf taxa or young examples of other largebodied species (Carpenter, 1992;
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<bibRefCitation id="76024B370B5C0D65FD17A20E3EABF61B" author="Carr, T. D." box="[737,914,2397,2433]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="1" pageNumber="2" 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>
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; Currie and Dong, 2001a). Many names are based on fragmentary remains, and whether all are properly diagnosable is an open question. The group seems to bounce around in various phylogenetic analyses relative to other tetanuran groups. Indeed, even the meaning of
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<taxonomicName id="D5934D450B5C0D65FF5BA17C3C84F5C9" authorityName="sensu Sereno" authorityYear="1998" box="[173,445,2607,2643]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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itself is controversial and problematic.
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</paragraph>
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<paragraph id="122C36C60B5C0D65FF21A1093F80F43E" blockId="1.[170,1234,1295,3150]" pageId="1" pageNumber="2">
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Unambiguous skeletal remains of tyrannosaurids are restricted to the latest Cretaceous.
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<taxonomicName id="D5934D450B5C0D65FD8EA1D73EF7F532" authority="Gilmore, 1933" box="[632,974,2692,2728]" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="olseni">
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<emphasis id="20E7EAD40B5C0D65FD8EA1D73EF7F532" box="[632,974,2692,2728]" italics="true" pageId="1" pageNumber="2">Alectrosaurus olseni</emphasis>
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</taxonomicName>
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Gilmore, 1933 was formerly thought to be of Turonian or Cenomanian age, but more recent work indicates a Campanian age for the Iren Dabasu beds where
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<emphasis id="20E7EAD40B5C0D65FDFAA0573F9CF4B2" box="[524,677,2820,2856]" italics="true" pageId="1" pageNumber="2">A. olseni</emphasis>
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was found (Eberth and Currie, 1993). The better-known Asian and North American taxa also derive from Campanian or Maastrichtian units (
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<bibRefCitation id="76024B370B5C0D65FC2FA00B39F9F4E6" author="Russell, D. A." box="[985,1216,2904,2940]" journalOrPublisher="National Museum of Natural Sciences Publications in Palaeontology" pageId="1" pageNumber="2" pagination="1 - 34" part="1" refId="ref77430" refString="Russell, D. A. 1970. Tyrannosaurs from the Late Cretaceous of western Canada. National Museum of Natural Sciences Publications in Palaeontology 1: 1 - 34." title="Tyrannosaurs from the Late Cretaceous of western Canada" type="journal article" year="1970">Russell, 1970</bibRefCitation>
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; Carpenter, 1992; Holtz, 2001a).
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</paragraph>
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<paragraph id="122C36C60B5C0D65FF23A0FF3A60FE0A" blockId="1.[170,1234,1295,3150]" lastBlockId="1.[1282,2335,278,1545]" pageId="1" pageNumber="2">
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A partial skeleton from the Barremian of the Isle of Wight—
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<emphasis id="20E7EAD40B5C0D65FF5CA0843CCBF461" box="[170,498,3031,3067]" italics="true" pageId="1" pageNumber="2">Eotyrannus lengi—</emphasis>
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was described as a possible tyrannosauroid by
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<bibRefCitation id="76024B370B5C0D65FF17A7533F3AF3BE" author="Hutt, S. & D. Naish & D. M. Martill & M. J. Barker & P. Newbery" box="[225,515,3072,3108]" journalOrPublisher="Cretaceous Research" pageId="1" pageNumber="2" pagination="227 - 242" part="22" refId="ref74512" refString="Hutt, S., D. Naish, D. M. Martill, M. J. Barker, and P. Newbery. 2001. A preliminary account of a new tyrannosauroid theropod from the Wessex Formation (Early Cretaceous) of southern England. Cretaceous Research 22: 227 - 242." title="A preliminary account of a new tyrannosauroid theropod from the Wessex Formation (Early Cretaceous) of southern England" type="journal article" year="2001">Hutt et al. (2001)</bibRefCitation>
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. It shares several features, including fused nasals and very tyrannosaurid-like premaxillary dentition, with other tyrannosaurids. If alliance with tyrannosaurs is upheld, it would extend the known range of the lineage to the Barremian and its geographic range to Europe.
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</paragraph>
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<paragraph id="122C36C60B5C0D65FADBAAC93BE7FCF9" blockId="1.[1282,2335,278,1545]" pageId="1" pageNumber="2">
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Isolated teeth have been used to extend the stratigraphic range of
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<taxonomicName id="D5934D450B5C0D65FA55AA963B8AFE73" authorityName="sensu Sereno" authorityYear="1998" box="[1443,1715,453,489]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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into pre-Campanian units (
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<bibRefCitation id="76024B370B5C0D65F779AA9638AFFD89" author="Eaton, J. G. & J. I. Kirkland & J. H. Hutchison & R. Denton & R. C. O'Neill & J. M. Parrish" journalOrPublisher="Geological Society of America Bulletin" pageId="1" pageNumber="2" pagination="560 - 567" part="109" refId="ref73408" refString="Eaton, J. G., J. I. Kirkland, J. H. Hutchison, R. Denton, R. C. O'Neill, and J. M. Parrish. 1997. Nonmarine extinction across the Cenomanian-Turanian boundary, southwestern Utah, with a comparison to the Cretaceous-Tertiary extinction event. Geological Society of America Bulletin 109: 560 - 567." title="Nonmarine extinction across the Cenomanian-Turanian boundary, southwestern Utah, with a comparison to the Cretaceous-Tertiary extinction event" type="journal article" year="1997">Eaton et al., 1997</bibRefCitation>
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;
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<bibRefCitation id="76024B370B5C0D65FA5CAABC3BE0FD89" author="Cifelli, R. L. & R. L. Nydham & J. D. Gardner & A. Weil & J. G. Eaton & J. I. Kirkland & S. K. Madsen" box="[1450,1753,495,531]" journalOrPublisher="Utah Geological Survey Miscellaneous Publications" pageId="1" pageNumber="2" pagination="219 - 242" part="99" refId="ref72463" refString="Cifelli, R. L., R. L. Nydham, J. D. Gardner, A. Weil, J. G. Eaton, J. I. Kirkland, and S. K. Madsen. 1999. Medial Cretaceous vertebrates from the Cedar Mountain Formation, Elliot County, Utah: the Mussentuchit Local Fauna. Utah Geological Survey Miscellaneous Publications 99: 219 - 242." title="Medial Cretaceous vertebrates from the Cedar Mountain Formation, Elliot County, Utah: the Mussentuchit Local Fauna" type="journal article" year="1999">Cifelli et al., 1999</bibRefCitation>
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;
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<bibRefCitation id="76024B370B5C0D65F91BAABC3AE2FD89" author="Manabe, M." box="[1773,2011,495,531]" journalOrPublisher="Journal of Paleontology" pageId="1" pageNumber="2" pagination="1176 - 1178" part="73" refId="ref75671" refString="Manabe, M. 1999. The early evolution of Tyrannosauridae in Asia. Journal of Paleontology 73: 1176 - 1178." title="The early evolution of Tyrannosauridae in Asia" type="journal article" year="1999">Manabe, 1999</bibRefCitation>
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;
|
||
<bibRefCitation id="76024B370B5C0D65F819AABC342CFD89" author="Lucas, S. G. & A. B. Heckert & R. M. Sullivan" box="[2031,2325,495,531]" journalOrPublisher="New Mexico Museum of Natural History and Science" pageId="1" pageNumber="2" pagination="83 - 90" part="17" refId="ref75303" refString="Lucas, S. G., A. B. Heckert, and R. M. Sullivan. 2000. Cretaceous dinosaurs in New Mexico. New Mexico Museum of Natural History and Science 17: 83 - 90." title="Cretaceous dinosaurs in New Mexico" type="journal article" year="2000">Lucas et al., 2000</bibRefCitation>
|
||
; Carr and Williamson, 2000). Indeed, the first tyrannosaurid remains described in the scientific literature were isolated teeth (
|
||
<bibRefCitation id="76024B370B5C0D65FAE6A93E38DCFD0B" author="Leidy, J." box="[1296,1509,621,657]" journalOrPublisher="Proceedings of the Academy of Natural Sciences of Philadelphia" pageId="1" pageNumber="2" pagination="72 - 73" part="8" refId="ref75212" refString="Leidy, J. 1856. Notice of remains of extinct Reptiles and Fishes discovered by Dr. F. V. Hayden in the Bad Lands of the Judith River, Nebraska Territory. Proceedings of the Academy of Natural Sciences of Philadelphia 8: 72 - 73." title="Notice of remains of extinct Reptiles and Fishes discovered by Dr. F. V. Hayden in the Bad Lands of the Judith River, Nebraska Territory" type="journal article" year="1856">Leidy, 1856</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="76024B370B5C0D65F9F5A93E3BD8FD0B" author="Marsh, O. C." box="[1539,1761,621,657]" journalOrPublisher="American Journal of Science (Series 3)" pageId="1" pageNumber="2" pagination="449 - 453" part="43" refId="ref75694" refString="Marsh, O. C. 1892. Notice of new reptiles from the Laramie Formation. American Journal of Science (Series 3) 43: 449 - 453." title="Notice of new reptiles from the Laramie Formation" type="journal article" year="1892">Marsh, 1892</bibRefCitation>
|
||
). But although isolated tyrannosaurid teeth may be biogeographically and biostratigraphically informative, it is unclear how useful they are in diagnosing particular tyrannosaurid taxa, especially in light of suspected ontogenetic changes in tyrannosaurid dentition (
|
||
<bibRefCitation id="76024B370B5C0D65F7AFA845342CFCA0" author="Carr, T. D." box="[2137,2325,790,826]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="1" pageNumber="2" 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>
|
||
; Carr and Williamson, 2000).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="5A89654D0B5C0D65FADBA8383AEAFB79" pageId="1" pageNumber="2" type="materials_examined">
|
||
<paragraph id="122C36C60B5C0D65FADBA8383AEAFB79" blockId="1.[1282,2335,278,1545]" pageId="1" pageNumber="2">
|
||
Other pre-Campanian nondental remains possibly related to
|
||
<taxonomicName id="D5934D450B5C0D65FAF5A8C63B2AFC23" authorityName="sensu Sereno" authorityYear="1998" box="[1283,1555,917,953]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
include the Jurassic
|
||
<emphasis id="20E7EAD40B5C0D65F899A8C63561FC23" box="[1903,2136,917,953]" italics="true" pageId="1" pageNumber="2">Stokesosaurus</emphasis>
|
||
from North America (
|
||
<bibRefCitation id="76024B370B5C0D65FA5AA8933BA7FC7E" author="Madsen, J. H." box="[1452,1694,960,996]" journalOrPublisher="Journal of Paleontology" pageId="1" pageNumber="2" pagination="27 - 31" part="48" refId="ref75386" refString="Madsen, J. H. 1974. A new theropod dinosaur from the Upper Jurassic of Utah. Journal of Paleontology 48: 27 - 31." title="A new theropod dinosaur from the Upper Jurassic of Utah" type="journal article" year="1974">Madsen, 1974</bibRefCitation>
|
||
; Chure and Madsen, 1998);
|
||
<emphasis id="20E7EAD40B5C0D65F77DA8933422FC7E" box="[2187,2331,960,996]" italics="true" pageId="1" pageNumber="2">
|
||
<taxonomicName id="D5934D450B5C0D65F77DA893342CFC7E" baseAuthorityName="Kurzanov" baseAuthorityYear="1976" box="[2187,2325,960,996]" class="Reptilia" family="Dromaeosauridae" genus="Itemirus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">Itemirus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
known from an isolated braincase from the Cretaceous of Asia (Kurzanov, 1976b); and
|
||
<emphasis id="20E7EAD40B5C0D65F946AF453A94FBA0" box="[1712,1965,1046,1082]" italics="true" pageId="1" pageNumber="2">
|
||
<taxonomicName id="D5934D450B5C0D65F946AF453A9EFBA0" box="[1712,1959,1046,1082]" class="Reptilia" family="Metriacanthosauridae" genus="Siamotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">Siamotyrannus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
an Early Cretaceous form from Thailand based on hip and hindlimb elements (Buffetaut et al., 1997). Available material is incomplete enough to render phylogenetic analysis problematic (Holtz, 2001a) and comparisons with
|
||
<materialsCitation id="A2FB3C9B0B5C0D65F9C0AFEC3A01FB79" ID-GBIF-Occurrence="3396382519" box="[1590,1848,1215,1251]" collectionCode="FMNH" pageId="1" pageNumber="2" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
difficult.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="5A89654D0B5C0D66FADAAFB93C42FA7B" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2" type="description">
|
||
<paragraph id="122C36C60B5C0D65FADAAFB93890F993" blockId="1.[1282,2335,278,1545]" pageId="1" pageNumber="2">
|
||
Although the focus of this study is the description of a single specimen, a review of tyrannosaurid taxonomy is relevant because other tyrannosaurs form the core of the comparative basis used here. This section focuses on alpha-level taxonomy, reviewing taxa currently thought to pertain to
|
||
<taxonomicName id="D5934D450B5C0D65F817AEC035C9FA2D" authorityName="sensu Sereno" authorityYear="1998" box="[2017,2288,1427,1463]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
in the literature, with particular emphasis to those examined for this study.
|
||
</paragraph>
|
||
<paragraph id="122C36C60B5C0D65FAF4AD1B3A20F9F1" blockId="1.[1282,1817,1606,1643]" box="[1282,1817,1608,1643]" pageId="1" pageNumber="2">
|
||
<emphasis id="20E7EAD40B5C0D65FAF4AD1B3894F9F1" bold="true" box="[1282,1453,1608,1643]" pageId="1" pageNumber="2">Definition</emphasis>
|
||
of
|
||
<taxonomicName id="D5934D450B5C0D65FA1AAD1B3A20F9F1" authorityName="sensu Sereno" authorityYear="1998" box="[1516,1817,1608,1643]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">
|
||
<emphasis id="20E7EAD40B5C0D65FA1AAD1B3A20F9F1" bold="true" box="[1516,1817,1608,1643]" pageId="1" pageNumber="2">Tyrannosauridae</emphasis>
|
||
</taxonomicName>
|
||
</paragraph>
|
||
<paragraph id="122C36C60B5C0D65FADAADC03ACEF844" blockId="1.[1274,2332,1681,3159]" pageId="1" pageNumber="2">
|
||
It seems odd that this would be a problem—after all, everyone knows what a tyrannosaur is. But as we move from a typologically-oriented nomenclatural system to one centered on phylogeny, we seek a meaning of “
|
||
<taxonomicName id="D5934D450B5C0D65F896AC41354AF8AC" authorityName="sensu Sereno" authorityYear="1998" box="[1888,2163,1810,1846]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
” that ties the name to an evolutionary phenomenon rather than a collection of characters that imply some sort of “ideal” tyrant dinosaur. Unfortunately, the assemblage of animals to fall under the aegis of
|
||
<taxonomicName id="D5934D450B5C0D65FA24ACE93BD8F844" authorityName="sensu Sereno" authorityYear="1998" box="[1490,1761,1978,2014]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
is controversial.
|
||
</paragraph>
|
||
<paragraph id="122C36C60B5C0D65FADFACB53B9BF5C3" blockId="1.[1274,2332,1681,3159]" pageId="1" pageNumber="2">
|
||
<taxonomicName id="D5934D450B5C0D65FADFACB53AA3F790" authority="sensu Sereno (1998)" authorityName="sensu Sereno" authorityYear="1998" box="[1321,1946,2022,2058]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae sensu Sereno (1998)</taxonomicName>
|
||
is a stem-based group name including tyrannosauroids closer to
|
||
<taxonomicName id="D5934D450B5C0D65F838A35C35FBF7A9" box="[1998,2242,2063,2099]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5C0D65F838A35C35FBF7A9" box="[1998,2242,2063,2099]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
than to
|
||
<emphasis id="20E7EAD40B5C0D65FAC5A3693A3BF7C4" box="[1331,1794,2106,2142]" italics="true" pageId="1" pageNumber="2">
|
||
Alectrosaurus,
|
||
<taxonomicName id="D5934D450B5C0D65F9C1A3693BC5F7C4" authorityName="Leidy" authorityYear="1868" box="[1591,1788,2106,2142]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">Aublysodon</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
or
|
||
<emphasis id="20E7EAD40B5C0D65F8B9A3693507F7C4" box="[1871,2110,2106,2142]" italics="true" pageId="1" pageNumber="2">
|
||
<taxonomicName id="D5934D450B5C0D65F8B9A3693500F7C4" box="[1871,2105,2106,2142]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">Nanotyrannus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
This would mean that
|
||
<emphasis id="20E7EAD40B5C0D65FA42A3363BA2F713" box="[1460,1691,2149,2185]" italics="true" pageId="1" pageNumber="2">Alectrosaurus</emphasis>
|
||
and
|
||
<taxonomicName id="D5934D450B5C0D65F900A3363AE4F713" box="[1782,2013,2149,2185]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5C0D65F900A3363AE4F713" box="[1782,2013,2149,2185]" italics="true" pageId="1" pageNumber="2">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
would not be tyrannosaurids, as they would be in most previous taxonomic treatments of the group (e.g.,
|
||
<bibRefCitation id="76024B370B5C0D65F913A3E83AFCF745" author="Russell, D. A." box="[1765,1989,2235,2271]" journalOrPublisher="National Museum of Natural Sciences Publications in Palaeontology" pageId="1" pageNumber="2" pagination="1 - 34" part="1" refId="ref77430" refString="Russell, D. A. 1970. Tyrannosaurs from the Late Cretaceous of western Canada. National Museum of Natural Sciences Publications in Palaeontology 1: 1 - 34." title="Tyrannosaurs from the Late Cretaceous of western Canada" type="journal article" year="1970">Russell, 1970</bibRefCitation>
|
||
; Bakker et al., 1988; Molnar et
|
||
<emphasis id="20E7EAD40B5C0D65FA42A3B638EFF693" box="[1460,1494,2277,2313]" italics="true" pageId="1" pageNumber="2">al</emphasis>
|
||
., 1990; Carpenter, 1992;
|
||
<bibRefCitation id="76024B370B5C0D65F872A3B63574F693" author="Holtz, T. R." box="[1924,2125,2277,2313]" journalOrPublisher="Journal of Paleontology" pageId="1" pageNumber="2" pagination="1100 - 1117" part="65" refId="ref74056" refString="Holtz, T. R. 1994. The phylogenetic position of the Tyrannosauridae: implications for theropod systematics. Journal of Paleontology 65: 1100 - 1117." title="The phylogenetic position of the Tyrannosauridae: implications for theropod systematics" type="journal article" year="1994">Holtz, 1994</bibRefCitation>
|
||
), but would instead be part of the more-inclusive Tyrannosauroidea.
|
||
<taxonomicName id="D5934D450B5C0D65F722A24338E1F6C4" authorityName="Matthew & Brown" authorityYear="1922" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
|
||
, in this system, is also a stem-based group name, this time including tyrannosaurids closer to
|
||
<taxonomicName id="D5934D450B5C0D65F824A23735FFF612" box="[2002,2246,2404,2440]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5C0D65F824A23735FFF612" box="[2002,2246,2404,2440]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
than to
|
||
<emphasis id="20E7EAD40B5C0D65FADDA2DC3A01F629" box="[1323,1848,2447,2483]" italics="true" pageId="1" pageNumber="2">
|
||
<taxonomicName id="D5934D450B5C0D65FADDA2DC3B21F629" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" baseAuthorityName="Witmer" baseAuthorityYear="1997" box="[1323,1560,2447,2483]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">Albertosaurus</taxonomicName>
|
||
,
|
||
<taxonomicName id="D5934D450B5C0D65F9D8A2DC3A08F629" baseAuthorityName="Russell" baseAuthorityYear="1970" box="[1582,1841,2447,2483]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
or
|
||
<emphasis id="20E7EAD40B5C0D65F88AA2DC3560F629" box="[1916,2137,2447,2483]" italics="true" pageId="1" pageNumber="2">
|
||
<taxonomicName id="D5934D450B5C0D65F88AA2DC356DF629" box="[1916,2132,2447,2483]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
No phylogenetic analysis accompanied these definitions, but the definitions imply a set of relationships congruent with that of Bakker et al. (1988), in which
|
||
<taxonomicName id="D5934D450B5C0D65F97BA15E3A4DF5AB" box="[1677,1908,2573,2609]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5C0D65F97BA15E3A4DF5AB" box="[1677,1908,2573,2609]" italics="true" pageId="1" pageNumber="2">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
is a very basal member of the tyrannosaur group.
|
||
</paragraph>
|
||
<paragraph id="122C36C60B5C0D66FAD1A1313C30FE5A" blockId="1.[1274,2332,1681,3159]" lastBlockId="2.[172,1230,282,1505]" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">
|
||
In this case,
|
||
<taxonomicName id="D5934D450B5C0D65FA0BA1313A35F51C" authorityName="sensu Sereno" authorityYear="1998" box="[1533,1804,2658,2694]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
and
|
||
<taxonomicName id="D5934D450B5C0D65F892A131354AF51C" authorityName="Matthew & Brown" authorityYear="1922" box="[1892,2163,2658,2694]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
|
||
both rely on
|
||
<taxonomicName id="D5934D450B5C0D65FAC3A1DF3B25F52A" box="[1333,1564,2700,2736]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5C0D65FAC3A1DF3B25F52A" box="[1333,1564,2700,2736]" italics="true" pageId="1" pageNumber="2">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="D5934D450B5C0D65F980A1DF3A01F52A" authorityName="Leidy" authorityYear="1868" box="[1654,1848,2700,2736]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5C0D65F980A1DF3A01F52A" box="[1654,1848,2700,2736]" italics="true" pageId="1" pageNumber="2">Aublysodon</emphasis>
|
||
</taxonomicName>
|
||
as specifiers.
|
||
<taxonomicName id="D5934D450B5C0D65F7D8A1DF342CF52A" box="[2094,2325,2700,2736]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5C0D65F7D8A1DF342CF52A" box="[2094,2325,2700,2736]" italics="true" pageId="1" pageNumber="2">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
is very likely based on an immature specimen of
|
||
<taxonomicName id="D5934D450B5C0D65F7D7A1E53809F49F" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="20E7EAD40B5C0D65F7D7A1E53809F49F" italics="true" pageId="1" pageNumber="2">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="76024B370B5C0D65FAB1A1B23BEAF49F" author="Rhozhdestvensky, A. K." box="[1351,1747,2785,2821]" journalOrPublisher="Paleontologeskii Zhournal" pageId="1" pageNumber="2" pagination="95 - 109" part="2" refId="ref76907" refString="Rhozhdestvensky, A. K. 1965. [Growth changes and some problems of systematics of Asian dinosaurs.] Paleontologeskii Zhournal 2: 95 - 109. [Russian]" title="Growth changes and some problems of systematics of Asian dinosaurs." type="journal article" year="1965">Rhozhdestvensky, 1965</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="76024B370B5C0D65F91EA1B23AA1F49F" author="Carr, T. D." box="[1768,1944,2785,2821]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="1" pageNumber="2" 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>
|
||
). I agree with Carr and Williamson (2000, 2001) that the “dental tyrannosaur”
|
||
<taxonomicName id="D5934D450B5C0D65F751A05F3862F4C0" authorityName="Leidy" authorityYear="1868" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5C0D65F751A05F3862F4C0" italics="true" pageId="1" pageNumber="2">Aublysodon</emphasis>
|
||
</taxonomicName>
|
||
is a
|
||
<emphasis id="20E7EAD40B5C0D65FA43A0653B8BF4C0" box="[1461,1714,2870,2906]" italics="true" pageId="1" pageNumber="2">nomen dubium.</emphasis>
|
||
As currently defined,
|
||
<taxonomicName id="D5934D450B5C0D65F7C6A0653BB4F41E" authority="sensu Sereno, 1998" authorityName="sensu Sereno" authorityYear="1998" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae sensu Sereno, 1998</taxonomicName>
|
||
is redundant with
|
||
<taxonomicName id="D5934D450B5C0D65F82AA03338DCF435" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="20E7EAD40B5C0D65F82AA033342CF41E" box="[2012,2325,2912,2948]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus rex</emphasis>
|
||
<bibRefCitation id="76024B370B5C0D65FB0AA0D838DCF435" author="Osborn, H. F." box="[1276,1509,2955,2991]" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="1" pageNumber="2" pagination="259 - 265" part="21" refId="ref76275" refString="Osborn, H. F. 1905. Tyrannosaurus and other Cretaceous carnivorous dinosaurs. Bulletin of the American Museum of Natural History 21: 259 - 265." title="Tyrannosaurus and other Cretaceous carnivorous dinosaurs" type="journal article" year="1905">Osborn, 1905</bibRefCitation>
|
||
</taxonomicName>
|
||
. Indeed, if we could work out the reticulation within
|
||
<emphasis id="20E7EAD40B5C0D65FA84A0E638E3F443" box="[1394,1498,2997,3033]" italics="true" pageId="1" pageNumber="2">
|
||
<taxonomicName id="D5934D450B5C0D65FA84A0E638ECF443" authorityName="Osborn" authorityYear="1905" box="[1394,1493,2997,3033]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
<taxonomicName id="D5934D450B5C0D65FA1CA0E63BC0F443" authorityName="sensu Sereno" authorityYear="1998" box="[1514,1785,2997,3033]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
could even conceivably refer to a subset of
|
||
<emphasis id="20E7EAD40B5C0D65FA37A08C3B13F399" box="[1473,1578,3039,3075]" italics="true" pageId="1" pageNumber="2">
|
||
<taxonomicName id="D5934D450B5C0D65FA37A08C3B1DF399" authorityName="Osborn" authorityYear="1905" box="[1473,1572,3039,3075]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
because the types of
|
||
<taxonomicName id="D5934D450B5C0D65F869A08C35BFF399" box="[1951,2182,3039,3075]" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5C0D65F869A08C35BFF399" box="[1951,2182,3039,3075]" italics="true" pageId="1" pageNumber="2">Nanotyrannus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="D5934D450B5C0D65F717A08C38F1F3B7" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5C0D65F717A08C38F1F3B7" italics="true" pageId="1" pageNumber="2">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
presumably shared a common ancestor after the lineage as a whole arose. This would also render Tyrannosauridae and
|
||
<taxonomicName id="D5934D450B5F0D66FEBFAA4F3F62FEDA" authorityName="Matthew & Brown" authorityYear="1922" box="[329,603,284,320]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
|
||
synonymous, since
|
||
<taxonomicName id="D5934D450B5F0D66FC4FAA4939F1FEA4" authorityName="Matthew & Brown" authorityYear="1922" box="[953,1224,282,318]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
|
||
is based on
|
||
<taxonomicName id="D5934D450B5F0D66FE81AA143FB1FEF1" authorityName="sensu Sereno" authorityYear="1998" box="[375,648,327,363]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
and the only tyrannosaurid would be
|
||
<emphasis id="20E7EAD40B5F0D66FF1EAA223F14FE0E" box="[232,557,368,405]" italics="true" pageId="2" pageNumber="3">
|
||
<taxonomicName id="D5934D450B5F0D66FF1EAA223F11FE0E" authorityName="Osborn" authorityYear="1905" box="[232,552,368,405]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Both names would refer to the same clade.
|
||
</paragraph>
|
||
<paragraph id="122C36C60B5F0D66FF20AA953C63FC21" blockId="2.[172,1230,282,1505]" pageId="2" pageNumber="3">
|
||
Holtz (2001a) defined
|
||
<taxonomicName id="D5934D450B5F0D66FD8CAA973EB3FE72" authorityName="sensu Sereno" authorityYear="1998" box="[634,906,452,488]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
as a node-based group comprising the last common ancestor of
|
||
<taxonomicName id="D5934D450B5F0D66FC23AABE39F0FD8B" box="[981,1225,493,529]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5F0D66FC23AABE39F0FD8B" box="[981,1225,493,529]" italics="true" pageId="2" pageNumber="3">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="D5934D450B5F0D66FF02A9493C8FFDA4" authorityName="Leidy" authorityYear="1868" box="[244,438,538,574]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5F0D66FF02A9493C8FFDA4" box="[244,438,538,574]" italics="true" pageId="2" pageNumber="3">Aublysodon</emphasis>
|
||
</taxonomicName>
|
||
and all of its descendents. Node-based groups allow more precise consolidation of a group’s diagnosis, and this particular definition avoids problems associated with
|
||
<emphasis id="20E7EAD40B5F0D66FB8CA93F3C63FD24" italics="true" pageId="2" pageNumber="3">
|
||
<taxonomicName id="D5934D450B5F0D66FB8CA93F3C6DFD24" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Nanotyrannus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
but it still uses
|
||
<taxonomicName id="D5934D450B5F0D66FD88A9CB3E79FD26" authorityName="Leidy" authorityYear="1868" box="[638,832,664,700]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5F0D66FD88A9CB3E79FD26" box="[638,832,664,700]" italics="true" pageId="2" pageNumber="3">Aublysodon</emphasis>
|
||
</taxonomicName>
|
||
as a specifier for
|
||
<taxonomicName id="D5934D450B5F0D66FB70A9C53CB0FD73" authorityName="sensu Sereno" authorityYear="1998" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
. If
|
||
<taxonomicName id="D5934D450B5F0D66FE35A9973FBCFD72" authorityName="Leidy" authorityYear="1868" box="[451,645,708,744]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5F0D66FE35A9973FBCFD72" box="[451,645,708,744]" italics="true" pageId="2" pageNumber="3">Aublysodon</emphasis>
|
||
</taxonomicName>
|
||
is undiagnosable, the definition of
|
||
<taxonomicName id="D5934D450B5F0D66FF58A9BC3C86FC89" authorityName="sensu Sereno" authorityYear="1998" box="[174,447,751,787]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
is rendered ambiguous.
|
||
<taxonomicName id="D5934D450B5F0D66FCA1A9BF395EFC8A" authorityName="Matthew & Brown" authorityYear="1922" box="[855,1127,748,784]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
|
||
is the stem-based group name including
|
||
<taxonomicName id="D5934D450B5F0D66FD03A8443ED3FCA1" box="[757,1002,791,827]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5F0D66FD03A8443ED3FCA1" box="[757,1002,791,827]" italics="true" pageId="2" pageNumber="3">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
and tyrannosaurids closer to it than to
|
||
<emphasis id="20E7EAD40B5F0D66FD7AA8123E60FCFF" box="[652,857,833,869]" italics="true" pageId="2" pageNumber="3">
|
||
<taxonomicName id="D5934D450B5F0D66FD7AA8123E6AFCFF" authorityName="Leidy" authorityYear="1868" box="[652,851,833,869]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Aublysodon</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
but if
|
||
<taxonomicName id="D5934D450B5F0D66FC2CA81339A5FCFE" authorityName="Leidy" authorityYear="1868" box="[986,1180,832,868]" class="Reptilia" family="Tyrannosauridae" genus="Aublysodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis id="20E7EAD40B5F0D66FC2CA81339A5FCFE" box="[986,1180,832,868]" italics="true" pageId="2" pageNumber="3">Aublysodon</emphasis>
|
||
</taxonomicName>
|
||
is invalid,
|
||
<taxonomicName id="D5934D450B5F0D66FECCA83F3F72FC0A" authorityName="Matthew & Brown" authorityYear="1922" box="[314,587,876,912]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
|
||
and
|
||
<taxonomicName id="D5934D450B5F0D66FD50A8383E8EFC15" authorityName="sensu Sereno" authorityYear="1998" box="[678,951,875,911]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
would again be redundant.
|
||
</paragraph>
|
||
<paragraph id="122C36C60B5F0D66FF2FA8913C42FA7B" blockId="2.[172,1230,282,1505]" pageId="2" pageNumber="3">
|
||
Stem-based group names are most appropriate when the membership of a group is unclear (Sereno, 1999). Some potential tyrannosaurs are controversial, but there is a core group of taxa that are universally understood to belong to this group, and a node-based group name is most appropriate. Using more than two names in the definition can circumvent the problem of
|
||
<emphasis id="20E7EAD40B5F0D66FB79AFC33C65FB79" italics="true" pageId="2" pageNumber="3">Aublysodon's</emphasis>
|
||
potential invalidity; status changes in one specifier will not change the meaning of the group. I thus amend Holtz’s definition as follows:
|
||
<taxonomicName id="D5934D450B5F0D66FDD2AE403E0CFAAD" authorityName="sensu Sereno" authorityYear="1998" box="[548,821,1299,1335]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
is the last common ancestor of
|
||
<emphasis id="20E7EAD40B5F0D66FEA5AE6D3CCFFA22" italics="true" pageId="2" pageNumber="3">
|
||
<taxonomicName id="D5934D450B5F0D66FEA5AE6D3FB6FAF8" authorityName="Osborn" authorityYear="1905" box="[339,655,1342,1378]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||
,
|
||
<taxonomicName id="D5934D450B5F0D66FD51AE6E3ECFFAFA" authorityName="Maleev" authorityYear="1955" box="[679,1014,1340,1377]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="bataar">Tarbosaurus bataar</taxonomicName>
|
||
,
|
||
<taxonomicName id="D5934D450B5F0D66FBFCAE6F3CF0FA17" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="sarcophagus">Albertosaurus sarcophagus</taxonomicName>
|
||
,
|
||
<taxonomicName id="D5934D450B5F0D66FE13AE3B3F83FA16" box="[485,698,1384,1420]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="genus">Gorgosaurus</taxonomicName>
|
||
libratus,
|
||
<taxonomicName id="D5934D450B5F0D66FC91AE3539FDFA13" authority="Gilmore, 1933" box="[871,1220,1381,1418]" class="Reptilia" family="Tyrannosauridae" genus="Alectrosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="olseni">Alectrosaurus olseni</taxonomicName>
|
||
,
|
||
<taxonomicName id="D5934D450B5F0D66FF58AEC73CC9FA22" authority="Kurzanov 1976" box="[174,496,1428,1464]" class="Reptilia" family="Tyrannosauridae" genus="Alioramus" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="remotus">Alioramus remotus</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<taxonomicName id="D5934D450B5F0D66FDA2AEC13EE7FA2F" authorityName="Russell" authorityYear="1970" box="[596,990,1425,1462]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis id="20E7EAD40B5F0D66FDA2AEC13EE7FA2F" box="[596,990,1425,1462]" italics="true" pageId="2" pageNumber="3">Daspletosaurus torosus</emphasis>
|
||
</taxonomicName>
|
||
and all of its descendents.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |