1624 lines
233 KiB
XML
1624 lines
233 KiB
XML
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<mods:title id="FC4243621D5721DA98D1129F6182BC68">An Older and Exceptionally Large Adult Specimen of Tyrannosaurus rex</mods:title>
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<mods:namePart id="A58DD39A332ED754160B69CBD842936F">Persons, Scott W</mods:namePart>
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<mods:namePart id="D932AFE32DC19C8E398BB93AC7DD580D">Currie, Philip J.</mods:namePart>
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<mods:namePart id="AFE93DBFAE033ACE347AE3A4C0DAE033">Erickson, Gregory M.</mods:namePart>
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<mods:title id="4371EB889C67F21918084DE9F5A08698">The Anatomical Record</mods:title>
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<mods:date id="B2C3B2A1BC1E71BF5AD219E6847AA2A1">2019</mods:date>
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<mods:number id="6CAF7D7D258FC98E1B25AB4417BFDF43">2019-03-21</mods:number>
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<mods:number id="DDF21CDEC24A95F1CCE960F4BFD0606A">302</mods:number>
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<treatment id="112887D5FFA7FF90FF56FB76FEFFFA0B" ID-DOI="http://doi.org/10.5281/zenodo.4582709" ID-GBIF-Taxon="159236947" ID-Zenodo-Dep="4582709" LSID="urn:lsid:plazi:treatment:112887D5FFA7FF90FF56FB76FEFFFA0B" httpUri="http://treatment.plazi.org/id/112887D5FFA7FF90FF56FB76FEFFFA0B" lastPageId="15" lastPageNumber="16" pageId="0" pageNumber="1">
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<paragraph id="993E36C3FFA7FF9EFF56FB76FDB8FF0F" blockId="0.[137,778,1232,1604]" lastBlockId="1.[137,778,205,682]" lastPageId="1" lastPageNumber="2" pageId="0" pageNumber="1">
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<subSubSection id="D19B6548FFA7FF9FFF56FB76FCA8FAE0" box="[164,777,1274,1296]" pageId="0" pageNumber="1" type="nomenclature">
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Henry Fairfield Osborn first introduced
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<taxonomicName id="5E814D40FFA7FF9FFDB7FB76FCA8FAE0" authorityName="Osborn" authorityYear="1905" box="[581,777,1275,1296]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
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<emphasis id="ABF5EAD1FFA7FF9FFDB7FB76FCA8FAE0" box="[581,777,1275,1296]" italics="true" pageId="0" pageNumber="1">Tyrannosaurus rex</emphasis>
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</taxonomicName>
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</subSubSection>
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<subSubSection id="D19B6548FFA7FF9EFF7BFA9AFCDEFD2E" lastPageId="1" lastPageNumber="2" pageId="0" pageNumber="1" type="discussion">
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to the scientific community and the public in 1905 (
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<bibRefCitation id="FD104B32FFA7FF9FFD45FA9BFF1FFAB8" author="Osborn HF" journalOrPublisher="New York, NY: Bulletin of the American Museum of Natural History" pageId="0" pageNumber="1" refId="ref9584" refString="Osborn HF. 1905. Tyrannosaurus and other Cretaceous carniVorous dinosaurs. New York, NY: Bulletin of the American Museum of Natural History." title="Tyrannosaurus and other Cretaceous carniVorous dinosaurs" type="book" year="1905">Osborn, 1905</bibRefCitation>
|
||
). It has reigned as the quintessential giant carnivorous dinosaur ever since (
|
||
<bibRefCitation id="FD104B32FFA7FF9FFE86FAC3FE54FA94" author="Hone D." box="[372,501,1358,1380]" journalOrPublisher="London, UK: Bloomsbury Publishing" pageId="0" pageNumber="1" refId="ref9030" refString="Hone D. 2016. The Tyrannosaur chronicles: the biology of the tyrant dinosaurs. London, UK: Bloomsbury Publishing." title="The Tyrannosaur chronicles: the biology of the tyrant dinosaurs" type="book" year="2016">Hone, 2016</bibRefCitation>
|
||
). However, as studies of
|
||
<taxonomicName id="5E814D40FFA7FF9FFF7BFAE6FF65FA70" authorityName="Osborn" authorityYear="1905" box="[137,196,1387,1408]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
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||
<emphasis id="ABF5EAD1FFA7FF9FFF7BFAE6FF65FA70" box="[137,196,1387,1408]" italics="true" pageId="0" pageNumber="1">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and the rest of the
|
||
<taxonomicName id="5E814D40FFA7FF9FFE7FFAE7FDE0FA70" authorityName="Osborn" authorityYear="1906" box="[397,577,1386,1408]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
have progressed, it has become increasingly apparent that
|
||
<taxonomicName id="5E814D40FFA7FF9FFDD6FA0AFDFEFA6C" authorityName="Osborn" authorityYear="1905" box="[548,607,1415,1436]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
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<emphasis id="ABF5EAD1FFA7FF9FFDD6FA0AFDFEFA6C" box="[548,607,1415,1436]" italics="true" pageId="0" pageNumber="1">T. rex</emphasis>
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||
</taxonomicName>
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||
is far from typical and representative of the theropod lineage as a whole. Tyrannosaurids constitute a highly derived and anatomically unusual group, possessing wide and deep skulls, stout teeth, elongate hindlimbs with arctometatarsalian pedes, and reduced forelimbs with only two functional digits (
|
||
<bibRefCitation id="FD104B32FFA7FF9FFD5CF99FFF5FF9B4" author="Brusatte SL & Norell MA & Carr TD & Erickson GM & Hutchinson JR & Balanoff AM & Bever GS & Choiniere JN & Makovicky PJ & Xu X." journalOrPublisher="Science" pageId="0" pageNumber="1" pagination="1481 - 1485" part="329" refId="ref8254" refString="Brusatte SL, Norell MA, Carr TD, Erickson GM, Hutchinson JR, Balanoff AM, Bever GS, Choiniere JN, Makovicky PJ, Xu X. 2010. Tyrannosaur paleobiology: new research on ancient exemplar organisms. Science 329: 1481 - 1485." title="Tyrannosaur paleobiology: new research on ancient exemplar organisms" type="journal article" year="2010">Brusatte et al., 2010</bibRefCitation>
|
||
). Phylogenetically, tyrannosaurids are members of the Coelurosauria, which otherwise consists of predominantly small-bodied theropods.
|
||
<taxonomicName id="5E814D40FFA7FF9FFB7BFB66FAF9FAF0" authorityName="Osborn" authorityYear="1905" box="[1161,1368,1259,1280]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA7FF9FFB7BFB66FAF9FAF0" box="[1161,1368,1259,1280]" italics="true" pageId="0" pageNumber="1">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
stands out in representing the terminal extreme for gigantism within
|
||
<taxonomicName id="5E814D40FFA7FF9FFC84FAADFB8FFAC6" authorityName="Osborn" authorityYear="1906" box="[886,1070,1312,1334]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
(
|
||
<bibRefCitation id="FD104B32FFA7FF9FFBC9FAADFAB9FAC5" author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA & Yerby SA & Brochu CA" box="[1083,1304,1312,1334]" journalOrPublisher="Nature" pageId="0" pageNumber="1" pagination="772 - 775" part="430" refId="ref8744" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA, Yerby SA, Brochu CA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 430: 772 - 775." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">Erickson et al., 2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA7FF9FFAD0FAADFA01FAC6" author="Holtz TR" box="[1314,1440,1312,1334]" editor="Weishampel DB & Dodson P & Osmolska H" journalOrPublisher="Berkeley, California: University of California Press" pageId="0" pageNumber="1" pagination="111 - 136" refId="ref8949" refString="Holtz TR. 2004. Tyrannosauroidea. In: Weishampel DB, Dodson P, Osmolska H, editors. The Dinosauria. 2 nd ed. Berkeley, California: University of California Press. p 111 - 136." title="Tyrannosauroidea" type="book chapter" volumeTitle="The Dinosauria. 2 nd ed." year="2004">Holtz, 2004</bibRefCitation>
|
||
). The iconic size of
|
||
<taxonomicName id="5E814D40FFA7FF9FFC06FAB1FB94FAA1" authorityName="Osborn" authorityYear="1905" box="[1012,1077,1340,1361]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA7FF9FFC06FAB1FB94FAA1" box="[1012,1077,1340,1361]" italics="true" pageId="0" pageNumber="1">T. rex</emphasis>
|
||
</taxonomicName>
|
||
has led to myriad questions pertaining to its development (
|
||
<bibRefCitation id="FD104B32FFA7FF9FFBA8FADBFA9EFA9B" author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA & Yerby SA & Brochu CA" box="[1114,1343,1366,1388]" journalOrPublisher="Nature" pageId="0" pageNumber="1" pagination="772 - 775" part="430" refId="ref8744" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA, Yerby SA, Brochu CA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 430: 772 - 775." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">Erickson et al., 2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA7FF9FFAB9FADBFC41FA77" author="Erickson GM & Currie P." journalOrPublisher="Science" pageId="0" pageNumber="1" pagination="213 - 217" part="313" refId="ref8687" refString="Erickson GM, Currie P. 2006. Tyrannosaur life tables: an example of nonavian dinosaur population biology. Science 313: 213 - 217." title="Tyrannosaur life tables: an example of nonavian dinosaur population biology" type="journal article" year="2006">Erickson and Currie, 2006</bibRefCitation>
|
||
), how its great size evolved (
|
||
<bibRefCitation id="FD104B32FFA7FF9FFAF9FAFCFCC1FA52" author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA & Yerby SA & Brochu CA" journalOrPublisher="Nature" pageId="0" pageNumber="1" pagination="772 - 775" part="430" refId="ref8744" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA, Yerby SA, Brochu CA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 430: 772 - 775." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">Erickson et al., 2004</bibRefCitation>
|
||
), and the ecological and physical upper bounds of terrestrial predator existence (
|
||
<bibRefCitation id="FD104B32FFA7FF9FFB9BFA2AFA82FA4D" author="Alexander R." box="[1129,1315,1447,1469]" journalOrPublisher="Zool J Linn Soc" pageId="0" pageNumber="1" pagination="1 - 25" part="83" refId="ref7954" refString="Alexander R. 1985. Mechanics of posture and gait of some large dinosaurs. Zool J Linn Soc 83: 1 - 25." title="Mechanics of posture and gait of some large dinosaurs" type="journal article" year="1985">Alexander, 1985</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA7FF9FFAC7FA2AFC4DFA27" author="Henderson DM & Snively E." journalOrPublisher="Proc Biol Sci" pageId="0" pageNumber="1" pagination="S 57 - S 60" part="271 (Suppl 3)" refId="ref8880" refString="Henderson DM, Snively E. 2004. Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs. Proc Biol Sci 271 (Suppl 3): S 57 - S 60." title="Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs." type="journal article" year="2004">Henderson and Snively, 2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA7FF9FFC05FA4FFBD7FA28" author="Holtz TR" box="[1015,1142,1474,1496]" journalOrPublisher="Bloomington, IN: Indiana University Press" pageId="0" pageNumber="1" pagination="371 - 396" refId="ref8990" refString="Holtz TR. 2008. A critical reappraisal of the obligate scavenging hypothesis for Tyrannosaurus rex and other tyrant dinosaurs. Tyrannosaurus rex the Tyrant King. Bloomington, IN: Indiana University Press. p 371 - 396." title="A critical reappraisal of the obligate scavenging hypothesis for Tyrannosaurus rex and other tyrant dinosaurs." type="book chapter" volumeTitle="Tyrannosaurus rex the Tyrant King" year="2008">Holtz, 2008</bibRefCitation>
|
||
). Nevertheless, since
|
||
<taxonomicName id="5E814D40FFA7FF9FFAAFFA4EFA3AFA28" authorityName="Osborn" authorityYear="1905" box="[1373,1435,1475,1496]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA7FF9FFAAFFA4EFA3AFA28" box="[1373,1435,1475,1496]" italics="true" pageId="0" pageNumber="1">T. rex</emphasis>
|
||
</taxonomicName>
|
||
’s discovery other enormous carnivorous dinosaurs belonging to the non-coelurosaurian tetanuran lineages Allosauroidea and
|
||
<taxonomicName id="5E814D40FFA7FF9FFCAAF99EFC53F9D9" authorityName="Stromer" authorityYear="1915" box="[856,1010,1555,1577]" class="Reptilia" family="Spinosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="family">Spinosauridae</taxonomicName>
|
||
have been found that are purported to be nearly as large as
|
||
<taxonomicName id="5E814D40FFA7FF9FFC0CF9A2FB9EF9B4" authorityName="Osborn" authorityYear="1905" box="[1022,1087,1583,1604]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA7FF9FFC0CF9A2FB9EF9B4" box="[1022,1087,1583,1604]" italics="true" pageId="0" pageNumber="1">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, if not larger (
|
||
<bibRefCitation id="FD104B32FFA7FF9FFB1AF9A3FAD4F9B4" author="Currie PJ" box="[1256,1397,1582,1604]" journalOrPublisher="Science" pageId="0" pageNumber="1" pagination="971 - 972" part="272" refId="ref8455" refString="Currie PJ. 1996. Out of Africa: meat-eating dinosaurs that challenge Tyrannosaurus rex. Science 272: 971 - 972." title="Out of Africa: meat-eating dinosaurs that challenge Tyrannosaurus rex" type="journal article" year="1996">Currie, 1996</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA7FF9EFA76F9A3FEF2FF13" author="Dal Sasso C & Maganuco S & Buffetaut E & Mendez MA" journalOrPublisher="J Vertebr Paleontol" lastPageId="1" lastPageNumber="2" pageId="0" pageNumber="1" pagination="888 - 896" part="25" refId="ref8540" refString="Dal Sasso C, Maganuco S, Buffetaut E, Mendez MA. 2005. New information on the skull of the enigmatic theropod Spinosaurus, with remarks on its size and affinities. J Vertebr Paleontol 25: 888 - 896." title="New information on the skull of the enigmatic theropod Spinosaurus, with remarks on its size and affinities" type="journal article" year="2005">Dal Sasso et al., 2005</bibRefCitation>
|
||
), suggesting
|
||
<emphasis id="ABF5EAD1FFA6FF9EFE15FF43FD96FF13" box="[487,567,205,227]" italics="true" pageId="1" pageNumber="2">T. rex’ s</emphasis>
|
||
giantism does not stand out among nonavian theropods.
|
||
<paragraph id="993E36C3FFA7FF9FFF6DF9FFFF70F87D" blockId="0.[137,778,1650,1933]" pageId="0" pageNumber="1">
|
||
<parenthesis id="D1EAFBFBFFA7FF9FFF6DF9FFFF70F87D" pageId="0" pageNumber="1">ABBREVIATIONS: AMNH = American Museum of Natural History, New York, New York; BHI = Black Hills Institute of Geological Research, Hill City, South Dakota; CM = Carnegie Museum of Natural History, Pittsburgh, Pennsylvania; FMNH = Field Museum of Natural History, Chicago, Illinois; FSAC = Faculté des Sciences Aïn Chock, Casablanca, Morocco; MOR = Museum of the Rockies, Bozeman, Montana; MSNM = Museo di Storia Naturale di Milano, Milano, Italy; MUCPv-CH = Museo de la Universidad Nacional del Comahue, El Chocón collection, Neuquén, Argentina; RSM = Royal Saskatchewan Museum, Regina, Saskatchewan, Canada; TMP = Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta, Canada</parenthesis>
|
||
</paragraph>
|
||
<footnote id="FA9A2ACDFFA7FF9FFCCDF9FFFBBFF87A" pageId="0" pageNumber="1">
|
||
<paragraph id="993E36C3FFA7FF9FFCCDF9FFFBE5F96D" blockId="0.[810,1451,1650,1930]" pageId="0" pageNumber="1">Grant sponsor: National Science Foundation; Grant numbers: EAR 0207744, DBI 0446224.</paragraph>
|
||
<paragraph id="993E36C3FFA7FF9FFCCDF92AFA08F91A" blockId="0.[810,1451,1650,1930]" pageId="0" pageNumber="1">*Correspondence to: W. Scott Persons, Department of Biological Sciences, CW 405, Biological Sciences Bldg., University of Alberta, Edmonton, Alberta T6G 2E9, Canada. Tel: 780-492-1252</paragraph>
|
||
<paragraph id="993E36C3FFA7FF9FFCCDF979FBF7F8F7" blockId="0.[810,1451,1650,1930]" box="[831,1110,1780,1799]" pageId="0" pageNumber="1">E-mail: persons@ualberta.ca</paragraph>
|
||
<paragraph id="993E36C3FFA7FF9FFCCDF89FFC44F8CD" blockId="0.[810,1451,1650,1930]" pageId="0" pageNumber="1">Received 19 May 2018; Revised 11 November 2018; Accepted 10 December 2018.</paragraph>
|
||
<paragraph id="993E36C3FFA7FF9FFCB8F8CAFB86F8AA" blockId="0.[810,1451,1650,1930]" box="[842,1063,1863,1882]" pageId="0" pageNumber="1">DOI: 10.1002/ar.24118</paragraph>
|
||
<paragraph id="993E36C3FFA7FF9FFCD8F8D2FBBFF87A" blockId="0.[810,1451,1650,1930]" pageId="0" pageNumber="1">Published online 00 Month 2019 in Wiley Online Library (wileyonlinelibrary.com).</paragraph>
|
||
</footnote>
|
||
<paragraph id="993E36C3FFA6FF9EFF56FE89FE93FD5A" blockId="1.[137,778,205,682]" pageId="1" pageNumber="2">
|
||
<materialsCitation id="29E93C9EFFA6FF9EFF56FE89FE4EFE74" ID-GBIF-Occurrence="2421817396" collectingDate="1991" collectionCode="RSM" country="Canada" location="near Eastend" pageId="1" pageNumber="2" preparations="relatively complete (roughly 65%) including skull, hip, and limbs" specimenCode="RSM P2523.8" stateProvince="Saskatchewan">
|
||
<specimenCode id="C9279EB8FFA6FF9EFF56FE89FEE3FEE9" box="[164,322,259,281]" pageId="1" pageNumber="2">RSM P2523.8</specimenCode>
|
||
(
|
||
<collectionCode id="FF90AE06FFA6FF9EFEAEFE89FE35FEE9" box="[348,404,260,281]" collectionName="RSM" pageId="1" pageNumber="2">RSM</collectionCode>
|
||
: Royal Saskatchewan Museum, Regina, SK, Canada) was discovered in an iron-stained, heavily cemented sandstone lens of the Frenchman Formation (Maastrichtian, Upper Cretaceous) near
|
||
<location id="9C5E6018FFA6FF9EFD5EFED9FCA4FE9A" LSID="urn:lsid:plazi:treatment:112887D5FFA7FF90FF56FB76FEFFFA0B:9C5E6018FFA6FF9EFD5EFED9FCA4FE9A" box="[684,773,340,362]" country="Canada" name="Eastend" pageId="1" pageNumber="2" stateProvince="Saskatchewan">Eastend</location>
|
||
,
|
||
<collectingRegion id="5B45F821FFA6FF9EFF7BFEE3FE82FE74" box="[137,291,366,388]" country="Canada" name="Saskatchewan" pageId="1" pageNumber="2">Saskatchewan</collectingRegion>
|
||
,
|
||
<collectingCountry id="E1967653FFA6FF9EFEC6FEE3FE27FE74" box="[308,390,366,388]" name="Canada" pageId="1" pageNumber="2">Canada</collectingCountry>
|
||
in
|
||
<collectingDate id="FD7BE9EBFFA6FF9EFE47FEE2FE4AFE74" box="[437,491,367,388]" pageId="1" pageNumber="2">1991</collectingDate>
|
||
.
|
||
</materialsCitation>
|
||
However, excavation did not begin in earnest until 1994. The extremely hard matrix surrounding the fossils, in combination with the size of the specimen, led to a prolonged period of preparation and delays in disseminating its discovery and importance regarding theropod giantism. Although some measurements of the limb bones have been disseminated in previous studies (
|
||
<bibRefCitation id="FD104B32FFA6FF9EFF62FDA4FE8AFDCE" author="Larson NL" box="[144,299,553,574]" journalOrPublisher="Bloomington, IN: Indiana University Press" pageId="1" pageNumber="2" pagination="56" part="Vol. 1" refId="ref9425" refString="Larson NL. 2008 a. One hundred years of Tyrannosaurus rex: the skeletons. Tyrannosaurus rex, The Tyrant King, Bloomington, IN: Indiana University Press. Vol. 1. p 56." title="One hundred years of Tyrannosaurus rex: the skeletons." type="book chapter" volumeTitle="Tyrannosaurus rex, The Tyrant King" year="2008">Larson, 2008a</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA6FF9EFEC4FDA4FDA7FDCE" author="Benson RB & Campione NE & Carrano MT & Mannion PD & Sullivan C & Upchurch P & Evans DC" box="[310,518,553,575]" journalOrPublisher="PLoS Biol" pageId="1" pageNumber="2" pagination="e 1001853" part="12" refId="ref8068" refString="Benson RB, Campione NE, Carrano MT, Mannion PD, Sullivan C, Upchurch P, Evans DC. 2014. Rates of dinosaur body mass evolution indicate 170 million years of sustained ecological innovation on the avian stem lineage. PLoS Biol 12: e 1001853." title="Rates of dinosaur body mass evolution indicate 170 million years of sustained ecological innovation on the avian stem lineage" type="journal article" year="2014">Benson et al., 2014</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA6FF9EFDE0FDA4FD5FFDCF" author="Campione NE & Evans DC & Brown CM & Carrano MT" box="[530,766,553,575]" journalOrPublisher="Methods Ecol Evol" pageId="1" pageNumber="2" pagination="913 - 923" part="5" refId="ref8337" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol Evol 5: 913 - 923." title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">Campione et al., 2014</bibRefCitation>
|
||
), including inaccurate values (owing to being made prior to full preparation of the specimen]),
|
||
<materialsCitation id="29E93C9EFFA6FF9EFDF1FDD3FD36FD84" ID-GBIF-Occurrence="2597591319" box="[515,663,606,628]" collectionCode="RSM" pageId="1" pageNumber="2" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
has never been formally described and its skeletal proportions scientifi- cally quantified.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA6FF9EFE94FD69FD8DFD0C" blockId="1.[137,778,740,944]" box="[358,556,740,764]" pageId="1" pageNumber="2">
|
||
<heading id="C27681AFFFA6FF9EFE94FD69FD8DFD0C" allCaps="true" bold="true" box="[358,556,740,764]" centered="true" fontSize="10" level="1" pageId="1" pageNumber="2" reason="0">
|
||
<emphasis id="ABF5EAD1FFA6FF9EFE94FD69FD8DFD0C" bold="true" box="[358,556,740,764]" pageId="1" pageNumber="2">DESCRIPTION</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA6FF9EFF7BFC8BFEA7FCEE" blockId="1.[137,778,740,944]" box="[137,262,774,798]" pageId="1" pageNumber="2">
|
||
<heading id="C27681AFFFA6FF9EFF7BFC8BFEA7FCEE" bold="true" box="[137,262,774,798]" fontSize="10" level="2" pageId="1" pageNumber="2" reason="0">
|
||
<emphasis id="ABF5EAD1FFA6FF9EFF7BFC8BFEA7FCEE" bold="true" box="[137,262,774,798]" pageId="1" pageNumber="2">Overview</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA6FF9EFF56FCBDFAB7FDF9" blockId="1.[137,778,740,944]" lastBlockId="1.[810,1451,205,734]" pageId="1" pageNumber="2">
|
||
<materialsCitation id="29E93C9EFFA6FF9EFF56FCBDFE9DFCB5" ID-GBIF-Occurrence="2597591311" box="[164,316,815,837]" collectionCode="RSM" pageId="1" pageNumber="2" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
is represented by an associated but disarticulated skeleton (
|
||
<figureCitation id="01BA2A46FFA6FF9EFE9AFCC7FE02FC90" box="[360,419,842,864]" captionStart="Fig. 1" captionStartId="1.[159,189,1890,1909]" captionTargetBox="[143,1443,991,1853]" captionTargetId="figure@1.[142,1447,987,1854]" captionTargetPageId="1" captionText="Fig. 1. The osteology of Tyrannosaurus rex RSM P2523.8. A, The disarticulated skull in left and right lateral views, with the braincase in posterior view. Scale bars = 10 cm. B, Composite illustration showing known skeletal elements in approximate articulation." httpUri="https://zenodo.org/record/3374668/files/figure.png" pageId="1" pageNumber="2">Fig. 1</figureCitation>
|
||
). Of special note is the abundance of incompletely coossified elements that were previously used as indices of somatic immaturity for the taxon (
|
||
<bibRefCitation id="FD104B32FFA6FF9EFD4DFC0DFEADFC40" author="Currie PJ & Hurum JH & Sabath K." journalOrPublisher="Acta Palaeontol Pol" pageId="1" pageNumber="2" pagination="227 - 234" part="48" refId="ref8512" refString="Currie PJ, Hurum JH, Sabath K. 2003. Skull structure and evolution in tyrannosaurid dinosaurs. Acta Palaeontol Pol 48: 227 - 234." title="Skull structure and evolution in tyrannosaurid dinosaurs" type="journal article" year="2003">Currie et al., 2003</bibRefCitation>
|
||
). These include quadrates that are not fused to the quadratojugals, surangulars that are not fused to the articulars, and prominently visible open sutures among the basisphenoid, exoccipital-opisthotic, and prootic elements. Some fusions within
|
||
<materialsCitation id="29E93C9EFFA6FF9EFB94FE90FB5AFEC3" ID-GBIF-Occurrence="2597591336" box="[1126,1275,285,307]" collectionCode="RSM" pageId="1" pageNumber="2" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
’s skull are also not bilaterally symmetrical. For instance, the right jugal osteoderm [the rugose cheek horn common to many tyrannosaurs (
|
||
<bibRefCitation id="FD104B32FFA6FF9EFC84FEE0FBE7FE73" author="Currie PJ & Hurum JH & Sabath K." box="[886,1094,364,387]" journalOrPublisher="Acta Palaeontol Pol" pageId="1" pageNumber="2" pagination="227 - 234" part="48" refId="ref8512" refString="Currie PJ, Hurum JH, Sabath K. 2003. Skull structure and evolution in tyrannosaurid dinosaurs. Acta Palaeontol Pol 48: 227 - 234." title="Skull structure and evolution in tyrannosaurid dinosaurs" type="journal article" year="2003">Currie et al., 2003</bibRefCitation>
|
||
)] is fused, with the suture fully obliterated. Nevertheless, its left counterpart is completely unfused (
|
||
<figureCitation id="01BA2A46FFA6FF9EFC62FE2FFC6CFE49" box="[912,973,418,441]" captionStart="Fig. 1" captionStartId="1.[159,189,1890,1909]" captionTargetBox="[143,1443,991,1853]" captionTargetId="figure@1.[142,1447,987,1854]" captionTargetPageId="1" captionText="Fig. 1. The osteology of Tyrannosaurus rex RSM P2523.8. A, The disarticulated skull in left and right lateral views, with the braincase in posterior view. Scale bars = 10 cm. B, Composite illustration showing known skeletal elements in approximate articulation." httpUri="https://zenodo.org/record/3374668/files/figure.png" pageId="1" pageNumber="2">Fig. 1</figureCitation>
|
||
). The fourth sacral centrum is not fused to either the third or the fifth sacral centra. Numerous vertebral neural arches are not fused to the centra (
|
||
<figureCitation id="01BA2A46FFA6FF9EFADFFE55FACDFE1E" box="[1325,1388,472,494]" captionStart="Fig. 2" captionStartId="2.[159,189,1647,1666]" captionTargetBox="[137,780,209,1611]" captionTargetId="figure@2.[137,780,203,1611]" captionTargetPageId="2" captionText="Fig. 2. Select vertebrae from RSM P2523.8. A, Dorsal and anterior caudal vertebrae showing variation in the fusion of the neural arch to the centrum. B, The second and third sacral vertebrae, in right lateral view. All scale bars = 10 cm." httpUri="https://zenodo.org/record/3374672/files/figure.png" pageId="1" pageNumber="2">Fig. 2</figureCitation>
|
||
), and the scapula is not fused to the coracoid (
|
||
<figureCitation id="01BA2A46FFA6FF9EFB3FFE7FFAABFDF9" box="[1229,1290,498,521]" captionStart="Fig. 3" captionStartId="2.[831,861,1035,1054]" captionTargetBox="[811,1447,205,997]" captionTargetId="figure@2.[810,1449,203,999]" captionTargetPageId="2" captionText="Fig. 3. Select appendicular elements from RSM P2523.8. A, The left scapula in lateral view. B, The right femur in anterior view. C, The right fibula in anterior view. D, Right pedal phalanx IV-1 in right lateral view." httpUri="https://zenodo.org/record/3374670/files/figure.png" pageId="1" pageNumber="2">Fig. 3</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA6FF9EFCB6FD80FCDEFD2E" blockId="1.[810,1451,205,734]" pageId="1" pageNumber="2">
|
||
As is common among tyrannosaurs (
|
||
<bibRefCitation id="FD104B32FFA6FF9EFB05FD80FCC3FDCE" author="Brusatte SL & Norell MA & Carr TD & Erickson GM & Hutchinson JR & Balanoff AM & Bever GS & Choiniere JN & Makovicky PJ & Xu X." journalOrPublisher="Science" pageId="1" pageNumber="2" pagination="1481 - 1485" part="329" refId="ref8254" refString="Brusatte SL, Norell MA, Carr TD, Erickson GM, Hutchinson JR, Balanoff AM, Bever GS, Choiniere JN, Makovicky PJ, Xu X. 2010. Tyrannosaur paleobiology: new research on ancient exemplar organisms. Science 329: 1481 - 1485." title="Tyrannosaur paleobiology: new research on ancient exemplar organisms" type="journal article" year="2010">Brusatte et al., 2010</bibRefCitation>
|
||
),
|
||
<materialsCitation id="29E93C9EFFA6FF9EFC84FDA5FBAEFDCD" ID-GBIF-Occurrence="2597591323" box="[886,1039,551,573]" collectionCode="RSM" pageId="1" pageNumber="2" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
is riddled with skeletal pathologies. These include rugosities and deformations surrounding the neural canals of caudal vertebrae 15–17, a thick ingrowth of interdental bone within the eighth alveolus of the left maxilla (
|
||
<figureCitation id="01BA2A46FFA6FF9EFC19FD1FFB8DFD58" box="[1003,1068,658,680]" captionStart="Fig. 4" captionStartId="3.[159,189,1413,1432]" captionTargetBox="[139,776,203,1378]" captionTargetId="figure@3.[139,776,203,1378]" captionTargetPageId="3" captionText="Fig. 4. Left maxilla pathology of RSM P2523.8 in lingual (A, B) and ventral (C, D) view. Numbers denote alveoli. Note the ingrowth of interdental bone within the eighth alveolus." httpUri="https://zenodo.org/record/3374676/files/figure.png" pageId="1" pageNumber="2">Fig. 4</figureCitation>
|
||
) and an incompletely healed and fractured dorsal rib head from the right side of the animal (
|
||
<figureCitation id="01BA2A46FFA6FF9EFCC0FD45FCD0FD2E" box="[818,881,712,734]" captionStart="Fig. 5" captionStartId="3.[831,861,776,795]" captionTargetBox="[814,1438,206,738]" captionTargetId="figure@3.[812,1448,203,740]" captionTargetPageId="3" captionText="Fig. 5. Pathologic dorsal rib head of RSM P2523.8. Swollen region suggests an incompletely healed injury or infection." httpUri="https://zenodo.org/record/3374674/files/figure.png" pageId="1" pageNumber="2">Fig. 5</figureCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</paragraph>
|
||
<subSubSection id="D19B6548FFA6FF9DFCD8FC8BFAE4FAEA" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2" type="diagnosis">
|
||
<paragraph id="993E36C3FFA6FF9EFCD8FC8BFC0BFCEE" blockId="1.[810,1450,774,944]" box="[810,938,774,798]" pageId="1" pageNumber="2">
|
||
<heading id="C27681AFFFA6FF9EFCD8FC8BFC0BFCEE" bold="true" box="[810,938,774,798]" fontSize="10" level="2" pageId="1" pageNumber="2" reason="0">
|
||
<emphasis id="ABF5EAD1FFA6FF9EFCD8FC8BFC0BFCEE" bold="true" box="[810,938,774,798]" pageId="1" pageNumber="2">Diagnosis</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA6FF9DFCB6FCBDFAE4FAEA" blockId="1.[810,1450,774,944]" lastBlockId="2.[810,1449,1177,1306]" lastPageId="2" lastPageNumber="3" pageId="1" pageNumber="2">
|
||
<materialsCitation id="29E93C9EFFA6FF9EFCB6FCBDFC76FCB5" ID-GBIF-Occurrence="2597591349" box="[836,983,815,837]" collectionCode="RSM" pageId="1" pageNumber="2" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
is referred to
|
||
<taxonomicName id="5E814D40FFA6FF9EFB81FCBDFB10FCB5" authorityName="Osborn" authorityYear="1905" box="[1139,1201,816,837]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA6FF9EFB81FCBDFB10FCB5" box="[1139,1201,816,837]" italics="true" pageId="1" pageNumber="2">T. rex</emphasis>
|
||
</taxonomicName>
|
||
based on the following suite of anatomical characteristics: (1) premaxillary teeth arranged in a laterally oriented tooth row and with D-shaped cross-sections; (2) a maxilla with a rounded rostral margin and a convex ventral margin; (3) “keyhole-shaped” and anteriorly directed orbits; (4) a corneal boss posterodorsal to the orbit; (5) a broad and elongate quadratojugal process and pneumatic excavation of the squamosal; (6) a maximal skull width estimated to be nearly two-thirds that of the premaxilla-occipital condyle length; (7) a long and strap-like scapula with accentuated proximal expansion; (8) a pubic shaft showing concave anterior curvature; and finally (9) an anteriorly projecting sheet of bone spanning roughly half the dorsoventral length of the ischium.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<caption id="CDFE664BFFA6FF9EFF6DF8EFFBDBF87D" httpUri="https://zenodo.org/record/3374668/files/figure.png" pageId="1" pageNumber="2" startId="1.[159,189,1890,1909]" targetBox="[143,1443,991,1853]" targetPageId="1">
|
||
<paragraph id="993E36C3FFA6FF9EFF6DF8EFFBDBF87D" blockId="1.[137,1450,1890,1933]" pageId="1" pageNumber="2">
|
||
Fig. 1. The osteology of
|
||
<taxonomicName id="5E814D40FFA6FF9EFE8BF8EEFDBDF885" authorityName="Osborn" authorityYear="1905" box="[377,540,1891,1909]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA6FF9EFE8BF8EEFDBDF885" box="[377,540,1891,1909]" italics="true" pageId="1" pageNumber="2">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
<materialsCitation id="29E93C9EFFA6FF9EFDD0F8EFFD01F885" ID-GBIF-Occurrence="2597591325" box="[546,672,1890,1909]" collectionCode="RSM" pageId="1" pageNumber="2" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
.
|
||
<emphasis id="ABF5EAD1FFA6FF9EFD5BF8EEFD16F885" bold="true" box="[681,695,1891,1909]" pageId="1" pageNumber="2">A</emphasis>
|
||
, The disarticulated skull in left and right lateral views, with the braincase in posterior view. Scale bars = 10 cm.
|
||
<emphasis id="ABF5EAD1FFA6FF9EFE84F8F6FE25F87D" bold="true" box="[374,388,1915,1933]" pageId="1" pageNumber="2">B</emphasis>
|
||
, Composite illustration showing known skeletal elements in approximate articulation.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="CDFE664BFFA5FF9DFCCDFB86FC3AFB96" httpUri="https://zenodo.org/record/3374670/files/figure.png" pageId="2" pageNumber="3" startId="2.[831,861,1035,1054]" targetBox="[811,1447,205,997]" targetPageId="2">
|
||
<paragraph id="993E36C3FFA5FF9DFCCDFB86FC3AFB96" blockId="2.[810,1451,1035,1126]" pageId="2" pageNumber="3">
|
||
Fig. 3. Select appendicular elements from
|
||
<materialsCitation id="29E93C9EFFA5FF9DFB2BFB86FAC1FBEE" ID-GBIF-Occurrence="2597591307" box="[1241,1376,1035,1054]" collectionCode="RSM" pageId="2" pageNumber="3" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
.
|
||
<emphasis id="ABF5EAD1FFA5FF9DFA99FB81FAD8FBEE" bold="true" box="[1387,1401,1036,1054]" pageId="2" pageNumber="3">A</emphasis>
|
||
, The left scapula in lateral view.
|
||
<emphasis id="ABF5EAD1FFA5FF9DFBC5FBA9FBE4FBC6" bold="true" box="[1079,1093,1060,1078]" pageId="2" pageNumber="3">B</emphasis>
|
||
, The right femur in anterior view.
|
||
<emphasis id="ABF5EAD1FFA5FF9DFA67FBA9FA05FBC6" bold="true" box="[1429,1444,1060,1078]" pageId="2" pageNumber="3">C</emphasis>
|
||
, The right fibula in anterior view.
|
||
<emphasis id="ABF5EAD1FFA5FF9DFBAEFBB1FBCAFBBE" bold="true" box="[1116,1131,1084,1102]" pageId="2" pageNumber="3">D</emphasis>
|
||
, Right pedal phalanx IV-1 in right lateral view.
|
||
</paragraph>
|
||
</caption>
|
||
<subSubSection id="D19B6548FFA5FF96FCD8FAE0FBBDFA0E" lastPageId="9" lastPageNumber="10" pageId="2" pageNumber="3" type="description">
|
||
<paragraph id="993E36C3FFA5FF9DFCD8FAE0FC07FA75" blockId="2.[810,1451,1389,1933]" box="[810,934,1389,1413]" pageId="2" pageNumber="3">
|
||
<heading id="C27681AFFFA5FF9DFCD8FAE0FC07FA75" bold="true" box="[810,934,1389,1413]" fontSize="10" level="2" pageId="2" pageNumber="3" reason="0">
|
||
<emphasis id="ABF5EAD1FFA5FF9DFCD8FAE0FC07FA75" bold="true" box="[810,934,1389,1413]" pageId="2" pageNumber="3">Histology</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA5FF9DFCB6FA1AFB5EF9BD" blockId="2.[810,1451,1389,1933]" pageId="2" pageNumber="3">
|
||
Transverse petrographic microscopy analysis reveals that the diaphysis of the right fibula to be almost entirely composed of dense Haversian bone (
|
||
<figureCitation id="01BA2A46FFA5FF9DFB35FA41FA94FA12" box="[1223,1333,1484,1506]" captionStart="Fig. 6" captionStartId="4.[159,189,1308,1327]" captionTargetBox="[143,1444,203,1272]" captionTargetId="figure@4.[143,1445,203,1272]" captionTargetPageId="4" captionText="Fig. 6. Osteohistology of the right fibula of RSM P2523.8. A, Outer cortical microstructure viewed using incident light microscopy. Note the plethora of secondary osteons indicating dense haversian bone remodeling (HB) and the closely packed remnants of lines of arrested growth (LAGS) nearest the periosteal surface of the element. B, Deep cortical dense haversian remodeling viewed using polarized petrographic microscopy. C, Unorganized, globular and porous, medullar-like bone tissue architecture near the lateral surface of the element. Arrows indicate the borders of secondary osteons (SO), many of which are partially or completely eroded. D, Wedl tunnels created by fungal erosion (FE) associated with the medullar-like bone tissue." httpUri="https://zenodo.org/record/3374678/files/figure.png" pageId="2" pageNumber="3">Fig. 6A,B</figureCitation>
|
||
). Between two and five tightly spaced lines of arrested growth, collectively constituting an External Fundamental System (
|
||
<bibRefCitation id="FD104B32FFA5FF9DFCC3F991FC7CF9C2" author="Cormack D." box="[817,989,1564,1586]" journalOrPublisher="Philadelphia, PA: Lippincott" pageId="2" pageNumber="3" pagination="450 - 474" refId="ref8431" refString="Cormack D. 1987. The integumentary system. Ham' s Histology. Philadelphia, PA: Lippincott. p 450 - 474." title="The integumentary system" type="book chapter" volumeTitle="Ham' s Histology" year="1987">Cormack, 1987</bibRefCitation>
|
||
), are locally present nearest the lateral periosteal surface of the element (
|
||
<figureCitation id="01BA2A46FFA5FF9DFB52F9BAFB55F9BD" box="[1184,1268,1591,1613]" captionStart="Fig. 6" captionStartId="4.[159,189,1308,1327]" captionTargetBox="[143,1444,203,1272]" captionTargetId="figure@4.[143,1445,203,1272]" captionTargetPageId="4" captionText="Fig. 6. Osteohistology of the right fibula of RSM P2523.8. A, Outer cortical microstructure viewed using incident light microscopy. Note the plethora of secondary osteons indicating dense haversian bone remodeling (HB) and the closely packed remnants of lines of arrested growth (LAGS) nearest the periosteal surface of the element. B, Deep cortical dense haversian remodeling viewed using polarized petrographic microscopy. C, Unorganized, globular and porous, medullar-like bone tissue architecture near the lateral surface of the element. Arrows indicate the borders of secondary osteons (SO), many of which are partially or completely eroded. D, Wedl tunnels created by fungal erosion (FE) associated with the medullar-like bone tissue." httpUri="https://zenodo.org/record/3374678/files/figure.png" pageId="2" pageNumber="3">Fig. 6A</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA5FF9CFCB6F9DFFD93F9D7" blockId="2.[810,1451,1389,1933]" lastBlockId="3.[137,777,1526,1575]" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3">
|
||
Our histological sampling of the fibula of
|
||
<materialsCitation id="29E93C9EFFA5FF9DFAE6F9DFFA0BF997" ID-GBIF-Occurrence="2597591352" box="[1300,1450,1617,1639]" collectionCode="RSM" pageId="2" pageNumber="3" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
reveals porous, poorly organized bone matrix composing the cortex spanning from the medial periosteal surface to approximately 20% of the centroid (
|
||
<figureCitation id="01BA2A46FFA5FF9DFB4CF92CFAB2F947" box="[1214,1299,1697,1719]" captionStart="Fig. 6" captionStartId="4.[159,189,1308,1327]" captionTargetBox="[143,1444,203,1272]" captionTargetId="figure@4.[143,1445,203,1272]" captionTargetPageId="4" captionText="Fig. 6. Osteohistology of the right fibula of RSM P2523.8. A, Outer cortical microstructure viewed using incident light microscopy. Note the plethora of secondary osteons indicating dense haversian bone remodeling (HB) and the closely packed remnants of lines of arrested growth (LAGS) nearest the periosteal surface of the element. B, Deep cortical dense haversian remodeling viewed using polarized petrographic microscopy. C, Unorganized, globular and porous, medullar-like bone tissue architecture near the lateral surface of the element. Arrows indicate the borders of secondary osteons (SO), many of which are partially or completely eroded. D, Wedl tunnels created by fungal erosion (FE) associated with the medullar-like bone tissue." httpUri="https://zenodo.org/record/3374678/files/figure.png" pageId="2" pageNumber="3">Fig. 6C</figureCitation>
|
||
). There is no definitive evidence of medullary bone associated with ovulation (
|
||
<bibRefCitation id="FD104B32FFA5FF9DFC92F95AFBF5F91D" author="Schweitzer MH & Wittmeyer JL & Horner JR" box="[864,1108,1751,1773]" journalOrPublisher="Science" pageId="2" pageNumber="3" pagination="1456 - 1460" part="308" refId="ref9672" refString="Schweitzer MH, Wittmeyer JL, Horner JR. 2005. Gender-specific reproductive tissue in ratites and Tyrannosaurus rex. Science 308: 1456 - 1460." title="Gender-specific reproductive tissue in ratites and Tyrannosaurus rex" type="journal article" year="2005">Schweitzer et al., 2005</bibRefCitation>
|
||
). Although reminiscent of previous descriptions of medullary bone in
|
||
<taxonomicName id="5E814D40FFA5FF9DFB1BF97FFA8DF8F7" authorityName="Osborn" authorityYear="1905" box="[1257,1324,1778,1799]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA5FF9DFB1BF97FFA8DF8F7" box="[1257,1324,1778,1799]" italics="true" pageId="2" pageNumber="3">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, the inner region of the element shows extensive bioerosion in the form of radiating Wedl tunnels (
|
||
<bibRefCitation id="FD104B32FFA5FF9DFB95F8AAFA02F8CD" author="Trueman C & Martill DM" box="[1127,1443,1831,1853]" journalOrPublisher="Archaeometry" pageId="2" pageNumber="3" pagination="371 - 382" part="44" refId="ref9779" refString="Trueman C, Martill DM. 2002. The long- term survival of bone: the role of bioerosion. Archaeometry 44: 371 - 382." title="The long- term survival of bone: the role of bioerosion" type="journal article" year="2002">Trueman and Martill, 2002</bibRefCitation>
|
||
) (
|
||
<figureCitation id="01BA2A46FFA5FF9DFCC3F8CCFC22F8A7" box="[817,899,1857,1879]" captionStart="Fig. 6" captionStartId="4.[159,189,1308,1327]" captionTargetBox="[143,1444,203,1272]" captionTargetId="figure@4.[143,1445,203,1272]" captionTargetPageId="4" captionText="Fig. 6. Osteohistology of the right fibula of RSM P2523.8. A, Outer cortical microstructure viewed using incident light microscopy. Note the plethora of secondary osteons indicating dense haversian bone remodeling (HB) and the closely packed remnants of lines of arrested growth (LAGS) nearest the periosteal surface of the element. B, Deep cortical dense haversian remodeling viewed using polarized petrographic microscopy. C, Unorganized, globular and porous, medullar-like bone tissue architecture near the lateral surface of the element. Arrows indicate the borders of secondary osteons (SO), many of which are partially or completely eroded. D, Wedl tunnels created by fungal erosion (FE) associated with the medullar-like bone tissue." httpUri="https://zenodo.org/record/3374678/files/figure.png" pageId="2" pageNumber="3">Fig. 6D</figureCitation>
|
||
). The diagenetic activity appears to have remodeled the former secondary osteons, whose partial borders are evident from the histology, leading to unorganized mineral globules with only superficial similarities with medullar bone structure (
|
||
<bibRefCitation id="FD104B32FFA4FF9CFEC0F99CFD86F9D7" author="Schweitzer MH & Wittmeyer JL & Horner JR" box="[306,551,1553,1575]" journalOrPublisher="Science" pageId="3" pageNumber="4" pagination="1456 - 1460" part="308" refId="ref9672" refString="Schweitzer MH, Wittmeyer JL, Horner JR. 2005. Gender-specific reproductive tissue in ratites and Tyrannosaurus rex. Science 308: 1456 - 1460." title="Gender-specific reproductive tissue in ratites and Tyrannosaurus rex" type="journal article" year="2005">Schweitzer et al., 2005</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="CDFE664BFFA5FF9DFF6DF9E2FE2BF93A" httpUri="https://zenodo.org/record/3374672/files/figure.png" pageId="2" pageNumber="3" startId="2.[159,189,1647,1666]" targetBox="[137,780,209,1611]" targetPageId="2">
|
||
<paragraph id="993E36C3FFA5FF9DFF6DF9E2FE2BF93A" blockId="2.[137,778,1647,1738]" pageId="2" pageNumber="3">
|
||
Fig. 2. Select vertebrae from
|
||
<materialsCitation id="29E93C9EFFA5FF9DFE43F9E2FD93F972" ID-GBIF-Occurrence="2597591343" box="[433,562,1647,1666]" collectionCode="RSM" pageId="2" pageNumber="3" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
.
|
||
<emphasis id="ABF5EAD1FFA5FF9DFDCCF9FDFDEDF972" bold="true" box="[574,588,1648,1666]" pageId="2" pageNumber="3">A</emphasis>
|
||
, Dorsal and anterior caudal vertebrae showing variation in the fusion of the neural arch to the centrum.
|
||
<emphasis id="ABF5EAD1FFA5FF9DFEF6F92DFEB3F942" bold="true" box="[260,274,1696,1714]" pageId="2" pageNumber="3">B</emphasis>
|
||
, The second and third sacral vertebrae, in right lateral view. All scale bars = 10 cm.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="CDFE664BFFA4FF9CFCCDFC85FB53FCC3" httpUri="https://zenodo.org/record/3374674/files/figure.png" pageId="3" pageNumber="4" startId="3.[831,861,776,795]" targetBox="[814,1438,206,738]" targetPageId="3">
|
||
<paragraph id="993E36C3FFA4FF9CFCCDFC85FB53FCC3" blockId="3.[810,1450,776,819]" pageId="3" pageNumber="4">
|
||
Fig. 5. Pathologic dorsal rib head of
|
||
<materialsCitation id="29E93C9EFFA4FF9CFB65FC85FAB8FCEB" ID-GBIF-Occurrence="2597591330" box="[1175,1305,776,795]" collectionCode="RSM" pageId="3" pageNumber="4" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
. Swollen region suggests an incompletely healed injury or infection.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="CDFE664BFFA4FF9CFF6DFA08FDA6FA39" httpUri="https://zenodo.org/record/3374676/files/figure.png" pageId="3" pageNumber="4" startId="3.[159,189,1413,1432]" targetBox="[139,776,203,1378]" targetPageId="3">
|
||
<paragraph id="993E36C3FFA4FF9CFF6DFA08FDA6FA39" blockId="3.[137,777,1413,1481]" pageId="3" pageNumber="4">
|
||
Fig. 4. Left maxilla pathology of
|
||
<materialsCitation id="29E93C9EFFA4FF9CFE3EFA08FDE8FA68" ID-GBIF-Occurrence="2597591333" box="[460,585,1413,1432]" collectionCode="RSM" pageId="3" pageNumber="4" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in lingual (
|
||
<emphasis id="ABF5EAD1FFA4FF9CFD43FA0BFD1EFA68" bold="true" box="[689,703,1414,1432]" pageId="3" pageNumber="4">A</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFA4FF9CFD3FFA0BFD7AFA68" bold="true" box="[717,731,1414,1432]" pageId="3" pageNumber="4">B</emphasis>
|
||
) and ventral (
|
||
<emphasis id="ABF5EAD1FFA4FF9CFF24FA13FF44FA40" bold="true" box="[214,229,1438,1456]" pageId="3" pageNumber="4">C</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFA4FF9CFF04FA13FEA4FA40" bold="true" box="[246,261,1438,1456]" pageId="3" pageNumber="4">D</emphasis>
|
||
) view. Numbers denote alveoli. Note the ingrowth of interdental bone within the eighth alveolus.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFA4FF9CFF7BF9D3FEEFF986" blockId="3.[137,778,1630,1934]" box="[137,334,1630,1654]" pageId="3" pageNumber="4">
|
||
<heading id="C27681AFFFA4FF9CFF7BF9D3FEEFF986" bold="true" box="[137,334,1630,1654]" fontSize="10" level="2" pageId="3" pageNumber="4" reason="0">
|
||
<emphasis id="ABF5EAD1FFA4FF9CFF7BF9D3FEEFF986" bold="true" box="[137,334,1630,1654]" pageId="3" pageNumber="4">Skull Elements</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA4FF9CFF56F905FD40F91E" blockId="3.[137,778,1630,1934]" pageId="3" pageNumber="4">
|
||
The right premaxilla is incomplete (
|
||
<figureCitation id="01BA2A46FFA4FF9CFDE5F905FDF2F96D" box="[535,595,1672,1694]" captionStart="Fig. 7" captionStartId="4.[159,189,1890,1909]" captionTargetBox="[138,775,1468,1853]" captionTargetId="figure@4.[138,777,1468,1853]" captionTargetPageId="4" captionText="Fig. 7. The right premaxilla of RSM P2523.8 in labial (A) and lingual (B) view. Four alveoli (1–4) are present." httpUri="https://zenodo.org/record/3374680/files/figure.png" pageId="3" pageNumber="4">Fig. 7</figureCitation>
|
||
), but its total size is estimated at approximately 140 mm in anteroposterior length (based on comparison with
|
||
<materialsCitation id="29E93C9EFFA4FF9CFE02F933FDF8F923" ID-GBIF-Occurrence="2421817394" box="[496,601,1725,1747]" collectionCode="BHI" pageId="3" pageNumber="4" specimenCode="BHI 3033">BHI 3033</materialsCitation>
|
||
). The premaxilla includes the symphysis and the outer walls of four alveoli.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA4FF9CFF56F97EFBCAFC43" blockId="3.[137,778,1630,1934]" lastBlockId="3.[810,1450,871,1933]" pageId="3" pageNumber="4">
|
||
In lateral view, the right maxilla of
|
||
<materialsCitation id="29E93C9EFFA4FF9CFDC3F97EFD69F8F9" ID-GBIF-Occurrence="2597591348" box="[561,712,1779,1801]" collectionCode="RSM" pageId="3" pageNumber="4" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
has a strongly convex alveolar margin (
|
||
<figureCitation id="01BA2A46FFA4FF9CFDFEF880FDF1F8D3" box="[524,592,1805,1827]" captionStart="Fig. 8" captionStartId="5.[159,189,1231,1250]" captionTargetBox="[141,1439,203,1195]" captionTargetId="figure@5.[141,1447,203,1195]" captionTargetPageId="5" captionText="Fig. 8. The right and left maxilla of RSM P2523.8 in lateral (A, C) and medial (B, D) view, respectively. (af, antorbital fenestra; mf, maxillary fenestra; 1–11, maxillary alveoli)." httpUri="https://zenodo.org/record/3374682/files/figure.png" pageId="3" pageNumber="4">Fig. 8</figureCitation>
|
||
). The maxillary fenestra spans 122 mm at its greatest anteroposterior length, while the antorbital fenestra spans 280 mm. Ten of the presumed 11 alveoli are preserved. On the ventral surface of the maxillary shelf, there are pits above dentary tooth positions 2–10. As in other tyrannosaurids, other than
|
||
<taxonomicName id="5E814D40FFA4FF9CFC9BFC0FFA9AFC68" authority="(Currie, 2003)" authorityName="Currie" authorityYear="2003" baseAuthorityName="Currie" baseAuthorityYear="2003" box="[873,1339,898,920]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="sarcophagus">
|
||
<emphasis id="ABF5EAD1FFA4FF9CFC9BFC0FFB31FC67" box="[873,1168,898,919]" italics="true" pageId="3" pageNumber="4">Albertosaurus sarcophagus</emphasis>
|
||
(
|
||
<bibRefCitation id="FD104B32FFA4FF9CFB57FC0FFA95FC68" author="Currie PJ" box="[1189,1332,898,920]" journalOrPublisher="Can J Earth Sci" pageId="3" pageNumber="4" pagination="651 - 665" part="40" refId="ref8478" refString="Currie PJ. 2003. Allometric growth in tyrannosaurids (Dinosauria: Theropoda) from the Upper Cretaceous of North America and Asia. Can J Earth Sci 40: 651 - 665." title="Allometric growth in tyrannosaurids (Dinosauria: Theropoda) from the Upper Cretaceous of North America and Asia" type="journal article" year="2003">Currie, 2003</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, only the first four pits are pronounced.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA4FF9CFCB6FC3AFBF0FB28" blockId="3.[810,1450,871,1933]" pageId="3" pageNumber="4">
|
||
The nasals are fused and are nearly complete, lacking only the slender tips of the narial processes (
|
||
<figureCitation id="01BA2A46FFA4FF9CFADAFC5FFAC8FC18" box="[1320,1385,978,1000]" captionStart="Fig. 9" captionStartId="5.[159,189,1692,1711]" captionTargetBox="[141,1446,1317,1654]" captionTargetId="figure@5.[141,1447,1317,1655]" captionTargetPageId="5" captionText="Fig. 9. The nasals of RSM P2523.8 in dorsal view (na, naris)." httpUri="https://zenodo.org/record/3374684/files/figure.png" pageId="3" pageNumber="4">Fig. 9</figureCitation>
|
||
). The nasals are 790 mm long, with the missing tips likely adding an additional 10 mm (based on comparison with BHI 3033). The external surfaces of the nasals are heavily sculptured. The contact with the maxilla is asymmetrical and is not fused. However, independent movement of either bone would have been restricted by interfingering staircase sutures, as is the case in large specimens of
|
||
<taxonomicName id="5E814D40FFA4FF9CFAF4FB00FCD7FB4D" baseAuthorityName="Russell" baseAuthorityYear="1970" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis id="ABF5EAD1FFA4FF9CFAF4FB00FCD7FB4D" italics="true" pageId="3" pageNumber="4">Daspletosaurus torosus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="5E814D40FFA4FF9CFC76FB25FBFAFB4D" baseAuthorityName="Maleev" baseAuthorityYear="1955" box="[900,1115,1192,1213]" class="Reptilia" family="Tyrannosauridae" genus="Tarbosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="bataar">
|
||
<emphasis id="ABF5EAD1FFA4FF9CFC76FB25FBFAFB4D" box="[900,1115,1192,1213]" italics="true" pageId="3" pageNumber="4">Tarbosaurus bataar</emphasis>
|
||
</taxonomicName>
|
||
, and other specimens of
|
||
<taxonomicName id="5E814D40FFA4FF9CFA9EFB25FA0BFB4D" authorityName="Osborn" authorityYear="1905" box="[1388,1450,1192,1213]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA4FF9CFA9EFB25FA0BFB4D" box="[1388,1450,1192,1213]" italics="true" pageId="3" pageNumber="4">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<bibRefCitation id="FD104B32FFA4FF9CFCC3FB4FFBE7FB28" author="Hurum JH & Sabath K." box="[817,1094,1218,1240]" journalOrPublisher="Acta Palaeontol Pol" pageId="3" pageNumber="4" pagination="2" part="48" refId="ref9151" refString="Hurum JH, Sabath K. 2003. Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared. Acta Palaeontol Pol 48: 2." title="Giant theropod dinosaurs from Asia and North America: skulls of Tarbosaurus bataar and Tyrannosaurus rex compared" type="journal article" year="2003">Hurum and Sabath, 2003</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA4FF9CFCB6FB50FA08FA12" blockId="3.[810,1450,871,1933]" pageId="3" pageNumber="4">
|
||
As in other tyrannosaurids, a shallow concavity on the medial surface of the jugal has an opening that connects with the jugal sinus (
|
||
<figureCitation id="01BA2A46FFA4FF9CFBD4FA9FFBD8FAD8" box="[1062,1145,1298,1320]" captionStart="Fig. 10" captionStartId="6.[159,189,1185,1204]" captionTargetBox="[142,1445,204,1149]" captionTargetId="figure@6.[142,1445,203,1149]" captionTargetPageId="6" captionText="Fig. 10. The right and left jugal of RSM P2523.8 in lateral (A, C) and medial (B, D) view, respectively. Rugose ossification (jugal horn) is present only on the left side (orb, orbit; pf, pneumatic fenestra; pos, postorbital suture; jh, jugal horn)." httpUri="https://zenodo.org/record/3374686/files/figure.png" pageId="3" pageNumber="4">Fig. 10</figureCitation>
|
||
). There is a large circular pneumatic fenestra in the posteroventral corner of the antorbital fossa. A large rugosity is present on the posteroventral corner of the left jugal. This rugosity, which indicates the presence of a blunt jugal “horn,” is absent on the right jugal. It is known from other tyrannosaur specimens that this rugosity is a separate ossification that typically becomes fused to the jugal during ontogeny.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA4FF9CFCB6FA6AFB24F9BE" blockId="3.[810,1450,871,1933]" pageId="3" pageNumber="4">
|
||
The left lacrimal is nearly complete (
|
||
<figureCitation id="01BA2A46FFA4FF9CFB1DFA6AFAE1FA0E" box="[1263,1344,1511,1534]" captionStart="Fig. 11" captionStartId="7.[159,189,1712,1731]" captionTargetBox="[144,1444,203,1676]" captionTargetId="figure@7.[144,1444,203,1676]" captionTargetPageId="7" captionText="Fig. 11. The right and left lacrimal of RSM P2523.8 in lateral (A, C) and medial (B, D) view, respectively (af, antorbital fenestra; orb, orbit)." httpUri="https://zenodo.org/record/3374688/files/figure.png" pageId="3" pageNumber="4">Fig. 11</figureCitation>
|
||
). It measures 360 mm in anteroposterior length and 375 mm in dorsoventral height. The two arms meet at an acute angle of slightly less than 90 degrees.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA4FF9CFCB6F9DFFA9BF96E" blockId="3.[810,1450,871,1933]" pageId="3" pageNumber="4">
|
||
Supraorbital ossifications are fused to rugosities on both postorbitals (
|
||
<figureCitation id="01BA2A46FFA4FF9CFC08F9E0FBECF973" box="[1018,1101,1645,1667]" captionStart="Fig. 12" captionStartId="8.[159,189,1498,1517]" captionTargetBox="[141,1446,203,1462]" captionTargetId="figure@8.[141,1447,203,1462]" captionTargetPageId="8" captionText="Fig. 12. The right and left postorbital of RSM P2523.8 in lateral (A, C) and medial (B, D) view, respectively (orb, orbit; pb, postorbital boss; sqp, squamosal process)." httpUri="https://zenodo.org/record/3374690/files/figure.png" pageId="3" pageNumber="4">Fig. 12</figureCitation>
|
||
). The postorbital boss is 6 cm long, 14 cm high, and 7 cm thick (lateromedial).
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA4FF9CFCB6F92FFAE0F87D" blockId="3.[810,1450,871,1933]" pageId="3" pageNumber="4">
|
||
The right squamosal is exceptionally well preserved (
|
||
<figureCitation id="01BA2A46FFA4FF9CFCC0F930FC27F923" box="[818,902,1725,1747]" captionStart="Fig. 13" captionStartId="9.[159,189,564,583]" captionTargetBox="[141,1446,203,528]" captionTargetId="figure@9.[141,1447,203,528]" captionTargetPageId="9" captionText="Fig. 13. The right squamosal of RSM P2523.8 in lateral (A), posterodorsal (B), and anterior (C) view (ltf, lateral temporal fenestra; pnc, pneumatic chamber; pos, postorbital process; qc, quadrate cotylus; qjp, quadratojugal process)." httpUri="https://zenodo.org/record/3374692/files/figure.png" pageId="3" pageNumber="4">Fig. 13</figureCitation>
|
||
). There are two large pneumatophores in the posteromedial half of the medial pneumatic chamber. The larger of these pneumatophores (measuring 78 mm in dorsoventral height and 55 mm in anteroposterior length) appear in the region dorsal to the quadrate cotyle. A smaller anteroventral pneumatophore (measuring 18 mm in dorsoventral height and 35 mm in anteroposterior length) invades the medial anteroventral portion of the squamosal.
|
||
</paragraph>
|
||
<caption id="CDFE664BFFA3FF9BFF6DFA91FA36FA7F" httpUri="https://zenodo.org/record/3374678/files/figure.png" pageId="4" pageNumber="5" startId="4.[159,189,1308,1327]" targetBox="[143,1444,203,1272]" targetPageId="4">
|
||
<paragraph id="993E36C3FFA3FF9BFF6DFA91FA36FA7F" blockId="4.[137,1452,1308,1423]" pageId="4" pageNumber="5">
|
||
Fig. 6. Osteohistology of the right fibula of
|
||
<materialsCitation id="29E93C9EFFA3FF9BFDE4FA91FD32FADF" ID-GBIF-Occurrence="2597591313" box="[534,659,1308,1327]" collectionCode="RSM" pageId="4" pageNumber="5" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
.
|
||
<emphasis id="ABF5EAD1FFA3FF9BFD69FA90FD09FADF" bold="true" box="[667,680,1309,1327]" pageId="4" pageNumber="5">A</emphasis>
|
||
, Outer cortical microstructure viewed using incident light microscopy. Note the plethora of secondary osteons indicating dense haversian bone remodeling (HB) and the closely packed remnants of lines of arrested growth (LAGS) nearest the periosteal surface of the element.
|
||
<emphasis id="ABF5EAD1FFA3FF9BFE24FAC0FE45FAAF" bold="true" box="[470,484,1357,1375]" pageId="4" pageNumber="5">B</emphasis>
|
||
, Deep cortical dense haversian remodeling viewed using polarized petrographic microscopy.
|
||
<emphasis id="ABF5EAD1FFA3FF9BFAEEFAC0FA8AFAAF" bold="true" box="[1308,1323,1357,1375]" pageId="4" pageNumber="5">C</emphasis>
|
||
, Unorganized, globular and porous, medullar-like bone tissue architecture near the lateral surface of the element. Arrows indicate the borders of secondary osteons (SO), many of which are partially or completely eroded.
|
||
<emphasis id="ABF5EAD1FFA3FF9BFD9FFAF0FDDDFA7F" bold="true" box="[621,636,1405,1423]" pageId="4" pageNumber="5">D</emphasis>
|
||
, Wedl tunnels created by fungal erosion (FE) associated with the medullar-like bone tissue.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFA3FF9BFCB6FA3AFB61F922" blockId="4.[810,1451,1463,1933]" pageId="4" pageNumber="5">
|
||
The quadrates are tall (386 mm in dorsoventral height) (
|
||
<figureCitation id="01BA2A46FFA3FF9BFCC0FA5CFC25FA18" box="[818,900,1489,1512]" captionStart="Fig. 14" captionStartId="9.[159,189,1332,1351]" captionTargetBox="[143,1444,651,1296]" captionTargetId="figure@9.[143,1444,651,1296]" captionTargetPageId="9" captionText="Fig. 14. The right quadrate of RSM P2523.8 in lateral (A) and medial (B) view (ac, articular condyles for the lower jaw; pgw, pterygoid wing; qc, quadrate cotylus; qqf, quadrate-quadratojugal fenestra." httpUri="https://zenodo.org/record/3374694/files/figure.png" pageId="4" pageNumber="5">Fig. 14</figureCitation>
|
||
), but are not fused to the quadratojugals. This is atypical for mature tyrannosaurids. Relative to the anteroposterior length of the quadrate wing of the pterygoid, the pterygoid wing of the quadrate is unusually long (22 cm) for a tyrannosaurid. Ventromedial to the pterygoid process, a 50
|
||
<emphasis id="ABF5EAD1FFA3FF9BFBF3F9D8FBB1F99A" box="[1025,1040,1621,1642]" italics="true" pageId="4" pageNumber="5">×</emphasis>
|
||
30 mm pneumatic sinus enters the anterior surface. The anterolateral–posteromedial axis of the articular condyles for the lower jaw is 18 cm wide. The quadrate–quadratojugal fenestra is large with a dorsoventral diameter of roughly 10 cm.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA3FF9BFCB6F95AFB96F8A8" blockId="4.[810,1451,1463,1933]" pageId="4" pageNumber="5">
|
||
The quadratojugal of
|
||
<materialsCitation id="29E93C9EFFA3FF9BFBB7F95AFB40F91D" ID-GBIF-Occurrence="2597591306" box="[1093,1249,1751,1773]" collectionCode="RSM" pageId="4" pageNumber="5" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
extends approximately 15 cm into the lateral temporal fenestra (
|
||
<figureCitation id="01BA2A46FFA3FF9BFABDF97FFA3CF8F8" box="[1359,1437,1778,1800]" captionStart="Fig. 15" captionStartId="10.[159,189,1554,1573]" captionTargetBox="[143,1444,203,1518]" captionTargetId="figure@10.[143,1444,203,1518]" captionTargetPageId="10" captionText="Fig. 15. The right and left quadratojugal of RSM P2523.8 in lateral (A, C) and medial (B, D) view, respectively (ltf, lateral temporal fenestra; qf, quadrate fenestra)." httpUri="https://zenodo.org/record/3374696/files/figure.png" pageId="4" pageNumber="5">Fig. 15</figureCitation>
|
||
). However, the dorsal edge of the quadratojugal shows only a shallowly overlapping contact with the long quadratojugal process of the squamosal.
|
||
</paragraph>
|
||
<caption id="CDFE664BFFA3FF9BFF6DF8EFFE43F87D" httpUri="https://zenodo.org/record/3374680/files/figure.png" pageId="4" pageNumber="5" startId="4.[159,189,1890,1909]" targetBox="[138,775,1468,1853]" targetPageId="4">
|
||
<paragraph id="993E36C3FFA3FF9BFF6DF8EFFE43F87D" blockId="4.[137,777,1890,1933]" pageId="4" pageNumber="5">
|
||
Fig. 7. The right premaxilla of
|
||
<materialsCitation id="29E93C9EFFA3FF9BFE46F8EFFD90F885" ID-GBIF-Occurrence="2597591328" box="[436,561,1890,1909]" collectionCode="RSM" pageId="4" pageNumber="5" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in labial (
|
||
<emphasis id="ABF5EAD1FFA3FF9BFD7EF8EEFD3BF885" bold="true" box="[652,666,1891,1909]" pageId="4" pageNumber="5">A</emphasis>
|
||
) and lingual (
|
||
<emphasis id="ABF5EAD1FFA3FF9BFF7CF8F6FF3DF87D" bold="true" box="[142,156,1915,1933]" pageId="4" pageNumber="5">B</emphasis>
|
||
) view. Four alveoli (1–4) are present.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFA3FF99FCB6F8D0FF72F87E" blockId="4.[810,1451,1463,1933]" lastBlockId="6.[137,779,1299,1934]" lastPageId="6" lastPageNumber="7" pageId="4" pageNumber="5">
|
||
The braincase is nearly complete and has suffered only slight taphonomic distortion (
|
||
<figureCitation id="01BA2A46FFA3FF9BFB84F8FAFB62F87D" box="[1142,1219,1911,1933]" captionStart="Fig. 16" captionStartId="11.[159,189,1070,1089]" captionTargetBox="[142,1445,203,1034]" captionTargetId="figure@11.[142,1445,203,1034]" captionTargetPageId="11" captionText="Fig. 16. The braincase of RSM P2523.8 in posterior (A) and right lateral (B) view (bt, basal tuber; btg, basiterygoid; cb, cultriform base; bo, basioccipital; bpt, basipterygoid process; bt, basal tubera; eo, exoccipital; fm foramen magnum; fo, fenestra ovalis; fr, frontal; nc, nuchal crest; oc, occipital condyle; on, orbital notch; par, parietal; ptr, posterior tympanic recess; pop, paraoccipital process; pos, postorbital suture; sc, sagittal crest; scr, subcondylar recess; soc, supraoccipital; sph, sphenethmoid; V, X-XII, cranial nerves)." httpUri="https://zenodo.org/record/3374698/files/figure.png" pageId="4" pageNumber="5">Fig. 16</figureCitation>
|
||
) with the cultriform process and most of the left paroccipital process being the only notable absences. The parietals and frontals are tightly bound to each other and to the rest of the braincase by complex interdigitating sutures. However, these are not fused. The orbitosphenoid, septosphenoid, sphenethmoid, and part of the orbital septum are fused. The frontals are partially fused, and are dorsoventrally deep between the supratemporal fossae and interorbital region. As a result, the contacts with the nasals and prefrontals slope anteroventrally. The lacrimal suture is 11 cm deep dorsoventrally and the postorbital suture is 9 cm deep dorsoventrally. The fused parietals measure 195 mm across the midline, 270 mm between the laterosphenoid contacts, and roughly 600 mm between the distal ends of the suture boundary between the paroccipital processes. The nuchal crest is incomplete on the left side, but was clearly asymmetrical, being thinner on the right side (37 mm anteroposteriorly compared with 43 mm). The parietals form a pronounced sagittal crest. The laterosphenoid extends dorsolaterally into the postorbital process roughly 21 cm from the midline and more than 5 cm beyond its contact with the frontal and parietal. The dorsal surface of the postorbital process of the laterosphenoid is smooth where it contacts a trough-like surface in the postorbital. The supraoccipital is fused indistinguishably onto the occipital surface of the parietal, as is the case in the other
|
||
<taxonomicName id="5E814D40FFA1FF99FD3AF9DEFCABF998" authorityName="Osborn" authorityYear="1905" box="[712,778,1619,1640]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA1FF99FD3AF9DEFCABF998" box="[712,778,1619,1640]" italics="true" pageId="6" pageNumber="7">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens—
|
||
<materialsCitation id="29E93C9EFFA1FF99FEFCF9E3FE04F973" ID-GBIF-Occurrence="2421817400" box="[270,421,1645,1667]" collectionCode="TMP" pageId="6" pageNumber="7" specimenCode="TMP 81.6.1">TMP 1981.6.1</materialsCitation>
|
||
,
|
||
<materialsCitation id="29E93C9EFFA1FF99FE42F9E3FDF6F973" ID-GBIF-Occurrence="2421817402" box="[432,599,1646,1667]" collectionCode="FMNH" pageId="6" pageNumber="7" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
and
|
||
<materialsCitation id="29E93C9EFFA1FF99FD7EF9E3FD50F973" ID-GBIF-Occurrence="2421817393" box="[652,753,1645,1667]" collectionCode="MOR" pageId="6" pageNumber="7" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
— but unlike other tyrannosaurids, which tend to have pronounced knob-like processes that extend conspicuously posterior to the parietal. The basipterygoid process is 10 cm below the level of the basal tuber. The basipterygoid processes are 24 cm across. The basioccipital and exoccipitals are indistinguishably fused in the occipital condyle. The prootic is fused anteriorly with the basisphenoid and laterosphenoid, and the line of contact cannot be seen. However, the exoccipital-opisthotic, prootic, and basisphenoid sutures are visible.
|
||
</paragraph>
|
||
<caption id="CDFE664BFFA2FF9AFF6DFB42FE0BFB0A" httpUri="https://zenodo.org/record/3374682/files/figure.png" pageId="5" pageNumber="6" startId="5.[159,189,1231,1250]" targetBox="[141,1439,203,1195]" targetPageId="5">
|
||
<paragraph id="993E36C3FFA2FF9AFF6DFB42FE0BFB0A" blockId="5.[137,1450,1231,1274]" pageId="5" pageNumber="6">
|
||
Fig. 8. The right and left maxilla of
|
||
<materialsCitation id="29E93C9EFFA2FF9AFE19FB42FDC8FB12" ID-GBIF-Occurrence="2597591338" box="[491,617,1231,1250]" collectionCode="RSM" pageId="5" pageNumber="6" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in lateral (
|
||
<emphasis id="ABF5EAD1FFA2FF9AFD3CFB5DFD7AFB12" bold="true" box="[718,731,1232,1250]" pageId="5" pageNumber="6">A</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFA2FF9AFD18FB5DFD58FB12" bold="true" box="[746,761,1232,1250]" pageId="5" pageNumber="6">C</emphasis>
|
||
) and medial (
|
||
<emphasis id="ABF5EAD1FFA2FF9AFC88FB5DFC29FB12" bold="true" box="[890,904,1232,1250]" pageId="5" pageNumber="6">B</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFA2FF9AFC64FB5DFC04FB12" bold="true" box="[918,933,1232,1250]" pageId="5" pageNumber="6">D</emphasis>
|
||
) view, respectively. (af, antorbital fenestra; mf, maxillary fenestra; 1–11, maxillary alveoli).
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="CDFE664BFFA2FF9AFF6DF911FD14F95F" box="[159,693,1692,1711]" httpUri="https://zenodo.org/record/3374684/files/figure.png" pageId="5" pageNumber="6" startId="5.[159,189,1692,1711]" targetBox="[141,1446,1317,1654]" targetPageId="5">
|
||
<paragraph id="993E36C3FFA2FF9AFF6DF911FD14F95F" blockId="5.[159,693,1692,1711]" box="[159,693,1692,1711]" pageId="5" pageNumber="6">
|
||
Fig. 9. The nasals of
|
||
<materialsCitation id="29E93C9EFFA2FF9AFEAAF911FE72F95F" ID-GBIF-Occurrence="2597591339" box="[344,467,1692,1711]" collectionCode="RSM" pageId="5" pageNumber="6" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in dorsal view (na, naris).
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="CDFE664BFFA1FF99FF6DFB2CFC62FB3C" httpUri="https://zenodo.org/record/3374686/files/figure.png" pageId="6" pageNumber="7" startId="6.[159,189,1185,1204]" targetBox="[142,1445,204,1149]" targetPageId="6">
|
||
<paragraph id="993E36C3FFA1FF99FF6DFB2CFC62FB3C" blockId="6.[137,1450,1185,1228]" pageId="6" pageNumber="7">
|
||
Fig. 10. The right and left jugal of
|
||
<materialsCitation id="29E93C9EFFA1FF99FE21FB2CFDEEFB44" ID-GBIF-Occurrence="2597591317" box="[467,591,1185,1204]" collectionCode="RSM" pageId="6" pageNumber="7" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in lateral (
|
||
<emphasis id="ABF5EAD1FFA1FF99FD5CFB2FFD1AFB44" bold="true" box="[686,699,1186,1204]" pageId="6" pageNumber="7">A</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFA1FF99FD3AFB2FFD76FB44" bold="true" box="[712,727,1186,1204]" pageId="6" pageNumber="7">C</emphasis>
|
||
) and medial (
|
||
<emphasis id="ABF5EAD1FFA1FF99FCA0FB2FFCC1FB44" bold="true" box="[850,864,1186,1204]" pageId="6" pageNumber="7">B</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFA1FF99FC9EFB2FFCDAFB44" bold="true" box="[876,891,1186,1204]" pageId="6" pageNumber="7">D</emphasis>
|
||
) view, respectively. Rugose ossification (jugal horn) is present only on the left side (orb, orbit; pf, pneumatic fenestra; pos, postorbital suture; jh, jugal horn).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFA1FF99FCB6FA9EFB1CFA38" blockId="6.[810,1451,1299,1934]" pageId="6" pageNumber="7">
|
||
The left ectopterygoid is better preserved than the right (
|
||
<figureCitation id="01BA2A46FFA1FF99FC82FAA0FC1FFAB3" box="[880,958,1325,1347]" captionStart="Fig. 17" captionStartId="11.[159,189,1890,1909]" captionTargetBox="[138,778,1257,1854]" captionTargetId="figure@11.[138,778,1257,1854]" captionTargetPageId="11" captionText="Fig. 17. The right and left ectopterygoid of RSM P2523.8 in lateral (A, C) and medial (B, D) view, respectively (pn, pneumatic opening)." httpUri="https://zenodo.org/record/3374700/files/figure.png" pageId="6" pageNumber="7">Fig. 17</figureCitation>
|
||
), and is inflated to a thickness of more than 7 cm. A large, 73
|
||
<emphasis id="ABF5EAD1FFA1FF99FC06FAC4FBA2FAAE" box="[1012,1027,1353,1374]" italics="true" pageId="6" pageNumber="7">×</emphasis>
|
||
31 mm, medially oriented pneumatic opening leads into a sinus that hollows out the bone and extends into at least the base of the jugal process. There also seems to be a small, 18
|
||
<emphasis id="ABF5EAD1FFA1FF99FB8BFA14FB29FA5E" box="[1145,1160,1433,1454]" italics="true" pageId="6" pageNumber="7">×</emphasis>
|
||
9 mm pneumatophore in the base of the hooked jugal process.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA1FF99FCB6FA40FC5FF973" blockId="6.[810,1451,1299,1934]" pageId="6" pageNumber="7">
|
||
The minimum length of the right dentary of
|
||
<materialsCitation id="29E93C9EFFA1FF99FA81FA40FC23FA0E" ID-GBIF-Occurrence="2597591354" collectionCode="RSM" pageId="6" pageNumber="7" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
is 810 mm, but the posterior edge is missing (
|
||
<figureCitation id="01BA2A46FFA1FF99FCC0F98EFC23F9E9" box="[818,898,1539,1561]" captionStart="Fig. 18" captionStartId="12.[159,189,813,832]" captionTargetBox="[141,1446,203,777]" captionTargetId="figure@12.[141,1446,203,777]" captionTargetPageId="12" captionText="Fig. 18. The right and left dentary of RSM P2523.8 in lateral (A, C) and medial (B, D) view, respectively (1-14, alveoli; mf, mylohyal foramen)." httpUri="https://zenodo.org/record/3374702/files/figure.png" pageId="6" pageNumber="7">Fig. 18</figureCitation>
|
||
). At the level of the sixth alveolus, the left dentary is 94.5 mm wide mediolaterally. The tooth row consists of 14 alveoli on each side and is 595 mm long. The mylohyal foramen of the splenial has a dorsoventral diameter of 5.5 cm.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA1FF99FCB6F905FBF5F87E" blockId="6.[810,1451,1299,1934]" pageId="6" pageNumber="7">
|
||
The surangulars are 645 mm long, with maximum depths of 250 mm, but in spite of their great sizes, neither is fused with the articular (
|
||
<figureCitation id="01BA2A46FFA1FF99FB5AF930FB5BF924" box="[1192,1274,1725,1748]" captionStart="Fig. 19" captionStartId="12.[159,189,1890,1909]" captionTargetBox="[140,1445,1261,1851]" captionTargetId="figure@12.[140,1447,1261,1853]" captionTargetPageId="12" captionText="Fig. 19. The right and left surangular of RSM P2523.8 in lateral (A, C) and medial (B, D) view, respectively (as, adductor shelf; l, lesion; mf, maxillary fenestra; psf, posterior surangular fenestra)." httpUri="https://zenodo.org/record/3374704/files/figure.png" pageId="6" pageNumber="7">Fig. 19</figureCitation>
|
||
). The adductor shelf protrudes 4 cm over the lateral surface. As in most other tyrannosaurids, the main vertical body of the surangular is pierced by sporadic openings. The insertion on the articular for the depressor mandibular musculature is 20 cm across and faces more posteriorly than dorsally. Posteromedial to the glenoid is a large pneumatophore that is 25 mm in diameter.
|
||
</paragraph>
|
||
<caption id="CDFE664BFFA0FF98FF6DF93DFAF8F933" box="[159,1369,1712,1731]" httpUri="https://zenodo.org/record/3374688/files/figure.png" pageId="7" pageNumber="8" startId="7.[159,189,1712,1731]" targetBox="[144,1444,203,1676]" targetPageId="7">
|
||
<paragraph id="993E36C3FFA0FF98FF6DF93DFAF8F933" blockId="7.[159,1369,1712,1731]" box="[159,1369,1712,1731]" pageId="7" pageNumber="8">
|
||
Fig. 11. The right and left lacrimal of
|
||
<materialsCitation id="29E93C9EFFA0FF98FE16F93DFDFEF933" ID-GBIF-Occurrence="2597591335" box="[484,607,1712,1731]" collectionCode="RSM" pageId="7" pageNumber="8" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in lateral (
|
||
<emphasis id="ABF5EAD1FFA0FF98FD49F93CFD69F933" bold="true" box="[699,712,1713,1731]" pageId="7" pageNumber="8">A</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFA0FF98FD21F93CFD43F933" bold="true" box="[723,738,1713,1731]" pageId="7" pageNumber="8">C</emphasis>
|
||
) and medial (
|
||
<emphasis id="ABF5EAD1FFA0FF98FCA8F93CFCC9F933" bold="true" box="[858,872,1713,1731]" pageId="7" pageNumber="8">B</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFA0FF98FC81F93CFC23F933" bold="true" box="[883,898,1713,1731]" pageId="7" pageNumber="8">D</emphasis>
|
||
) view, respectively (af, antorbital fenestra; orb, orbit).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFA0FF98FEBEF96BFDE6F90E" blockId="7.[137,777,1766,1934]" box="[332,583,1766,1790]" pageId="7" pageNumber="8">
|
||
<heading id="C27681AFFFA0FF98FEBEF96BFDE6F90E" allCaps="true" bold="true" box="[332,583,1766,1790]" centered="true" fontSize="10" level="1" pageId="7" pageNumber="8" reason="0">
|
||
<emphasis id="ABF5EAD1FFA0FF98FEBEF96BFDE6F90E" bold="true" box="[332,583,1766,1790]" pageId="7" pageNumber="8">SIZE ESTIMATION</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA0FF98FF56F89CFA31F8D3" blockId="7.[137,777,1766,1934]" lastBlockId="7.[810,1450,1773,1934]" pageId="7" pageNumber="8">
|
||
To quantify the size of
|
||
<materialsCitation id="29E93C9EFFA0FF98FE7DF89CFDBFF8D6" ID-GBIF-Occurrence="2597591363" box="[399,542,1808,1830]" collectionCode="RSM" pageId="7" pageNumber="8" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
and compare it to that of other
|
||
<taxonomicName id="5E814D40FFA0FF98FF17F8A6FE82F8B0" authorityName="Osborn" authorityYear="1905" box="[229,291,1835,1856]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA0FF98FF17F8A6FE82F8B0" box="[229,291,1835,1856]" italics="true" pageId="7" pageNumber="8">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens (
|
||
<emphasis id="ABF5EAD1FFA0FF98FE5BF8A6FE16F8B0" box="[425,439,1835,1856]" italics="true" pageId="7" pageNumber="8">n</emphasis>
|
||
= 11), a series of skeletal measurements were taken. To help ensure consistency of measurement technique, all measurements on all specimens were made by one of the authors (P.J.C.). All measurements were made by hand using standard digital calipers or measurement tape, as the size of the measured element dictated (
|
||
<tableCitation id="D4030378FFA0FF98FAC5F880FA23F8D3" box="[1335,1410,1805,1827]" captionStart="Table 1" captionText="TABLE 1. Select measurements of RSM P2523.8 and other Tyrannosaurus rex specimens, all measurements in millimeters" pageId="7" pageNumber="8" targetBox="[249,670,203,1929]" targetPageId="13">Table 1</tableCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA0FF97FCB6F8A5FD02F87E" blockId="7.[810,1450,1773,1934]" lastBlockId="8.[137,778,1619,1934]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
|
||
The standard in neontology for contrasting size across animals of disparate proportions is to use body mass (
|
||
<bibRefCitation id="FD104B32FFA0FF98FCC3F8D0FB8DF882" author="Campione NE & Evans DC & Brown CM & Carrano MT" box="[817,1068,1885,1907]" journalOrPublisher="Methods Ecol Evol" pageId="7" pageNumber="8" pagination="913 - 923" part="5" refId="ref8337" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol Evol 5: 913 - 923." title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">Campione et al., 2014</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA0FF98FBC8F8D0FB43F883" author="Erickson GM" box="[1082,1250,1885,1907]" journalOrPublisher="Annu Rev Earth Planet Sci" pageId="7" pageNumber="8" pagination="675 - 697" part="42" refId="ref8667" refString="Erickson GM. 2014. On dinosaur growth. Annu Rev Earth Planet Sci 42: 675 - 697." title="On dinosaur growth" type="journal article" year="2014">Erickson, 2014</bibRefCitation>
|
||
).
|
||
<bibRefCitation id="FD104B32FFA0FF98FB05F8D0FCCCF87E" author="Campione NE & Evans DC & Brown CM & Carrano MT" journalOrPublisher="Methods Ecol Evol" pageId="7" pageNumber="8" pagination="913 - 923" part="5" refId="ref8337" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol Evol 5: 913 - 923." title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">Campione et al. (2014)</bibRefCitation>
|
||
offered a revised equation for using minimal femoral circumference (MFC) to estimate total body mass across nonavian bipeds, dinosaurs in particular (log
|
||
<subScript id="05053486FFAFFF97FD75F9FAFD3BF975" attach="both" box="[647,666,1655,1669]" fontSize="6" pageId="8" pageNumber="9">10</subScript>
|
||
<emphasis id="ABF5EAD1FFAFFF97FD69F9E3FD10F973" box="[667,689,1646,1667]" italics="true" pageId="8" pageNumber="9">M</emphasis>
|
||
= 2.754
|
||
<emphasis id="ABF5EAD1FFAFFF97FF7BF905FF2EF96D" box="[137,143,1672,1693]" italics="true" pageId="8" pageNumber="9">·</emphasis>
|
||
log
|
||
<subScript id="05053486FFAFFF97FF5DF91CFF63F96F" attach="both" box="[175,194,1681,1695]" fontSize="6" pageId="8" pageNumber="9">10</subScript>
|
||
<emphasis id="ABF5EAD1FFAFFF97FF31F905FF75F96D" box="[195,212,1672,1693]" italics="true" pageId="8" pageNumber="9">C</emphasis>
|
||
− 0.683, where
|
||
<emphasis id="ABF5EAD1FFAFFF97FE75F904FE3CF96E" box="[391,413,1673,1694]" italics="true" pageId="8" pageNumber="9">M</emphasis>
|
||
is body mass and
|
||
<emphasis id="ABF5EAD1FFAFFF97FD9EF905FDDCF96D" box="[620,637,1672,1693]" italics="true" pageId="8" pageNumber="9">C</emphasis>
|
||
is minimum femoral circumference). One limitation of such approaches is that the hollow femoral shafts of theropods are prone to taphonomic distortion. The mid-femur shaft of
|
||
<materialsCitation id="29E93C9EFFAFFF97FD21F955FF40F8F8" ID-GBIF-Occurrence="2597591355" collectionCode="RSM" pageId="8" pageNumber="9" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
is slightly damaged, but its minimum transverse diameter (199 mm) is not deformed and indicates a femoral circumference of approximately 590 mm (based on the ELLR method of tyrannosaur femoral circumference estimation,
|
||
<bibRefCitation id="FD104B32FFAFFF97FF18F8D0FE79F883" author="Bradley GJ & Burns ME & Currie PJ" box="[234,472,1885,1907]" journalOrPublisher="Cretaceous Res" pageId="8" pageNumber="9" pagination="200 - 209" part="55" refId="ref8118" refString="Bradley GJ, Burns ME, Currie PJ. 2015. Missing data estimation in tyrannosaurid dinosaurs: Can diameter take the place of circumference?. Cretaceous Res 55: 200 - 209." title="Missing data estimation in tyrannosaurid dinosaurs: Can diameter take the place of circumference?" type="journal article" year="2015">Bradley et al., 2015</bibRefCitation>
|
||
). The body mass of
|
||
<materialsCitation id="29E93C9EFFAFFF97FD21F8D0FF40F87E" ID-GBIF-Occurrence="2597591357" collectionCode="RSM" pageId="8" pageNumber="9" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
is thus estimated to be ~8,870 kg (
|
||
<emphasis id="ABF5EAD1FFAFFF97FDA6F8F4FDC7F87E" box="[596,614,1913,1934]" italics="true" pageId="8" pageNumber="9">±</emphasis>
|
||
25%).
|
||
</paragraph>
|
||
<caption id="CDFE664BFFAFFF97FF6DFA57FEE2F9F5" httpUri="https://zenodo.org/record/3374690/files/figure.png" pageId="8" pageNumber="9" startId="8.[159,189,1498,1517]" targetBox="[141,1446,203,1462]" targetPageId="8">
|
||
<paragraph id="993E36C3FFAFFF97FF6DFA57FEE2F9F5" blockId="8.[137,1451,1498,1541]" pageId="8" pageNumber="9">
|
||
Fig. 12. The right and left postorbital of
|
||
<materialsCitation id="29E93C9EFFAFFF97FDF5FA57FD22FA1D" ID-GBIF-Occurrence="2597591340" box="[519,643,1498,1517]" collectionCode="RSM" pageId="8" pageNumber="9" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in lateral (
|
||
<emphasis id="ABF5EAD1FFAFFF97FD10FA57FCAAFA1C" bold="true" box="[738,779,1498,1516]" pageId="8" pageNumber="9">A, C</emphasis>
|
||
) and medial (
|
||
<emphasis id="ABF5EAD1FFAFFF97FC77FA57FC0FFA1C" bold="true" box="[901,942,1498,1516]" pageId="8" pageNumber="9">B, D</emphasis>
|
||
) view, respectively (orb, orbit; pb, postorbital boss; sqp, squamosal process).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFAFFF97FCB6F9DEFC24F8D2" blockId="8.[810,1450,1618,1934]" pageId="8" pageNumber="9">
|
||
Following from the 590 mm femoral circumference estimation, the relative femoral section moduli (
|
||
<bibRefCitation id="FD104B32FFAFFF97FAFAF9E0FCC1F96D" author="Erickson GM & Catanese J III & Keaveny TM" journalOrPublisher="Anat Rec" pageId="8" pageNumber="9" pagination="115 - 124" part="268" refId="ref8714" refString="Erickson GM, Catanese J III, Keaveny TM. 2002. Evolution of the biomechanical material properties of the femur. Anat Rec 268: 115 - 124." title="Evolution of the biomechanical material properties of the femur" type="journal article" year="2002">Erickson et al., 2002</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFAFFF97FC9BF905FB11F96E" author="Therrien F & Henderson DM" box="[873,1200,1672,1694]" journalOrPublisher="J Vertebr Paleontol" pageId="8" pageNumber="9" pagination="108 - 115" part="27" refId="ref9743" refString="Therrien F, Henderson DM. 2007. My theropod is bigger than yours … or not: estimating body size from skull length in theropods. J Vertebr Paleontol 27: 108 - 115." title="My theropod is bigger than yours … or not: estimating body size from skull length in theropods" type="journal article" year="2007">Therrien and Henderson, 2007</bibRefCitation>
|
||
) of
|
||
<materialsCitation id="29E93C9EFFAFFF97FB27F905FAC6F96E" ID-GBIF-Occurrence="2597591322" box="[1237,1383,1672,1694]" collectionCode="RSM" pageId="8" pageNumber="9" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
is calculated at 650,289 cm
|
||
<superScript id="6EF49B8BFFAFFF97FBE6F910FBBFF95B" attach="left" box="[1044,1054,1693,1707]" fontSize="6" pageId="8" pageNumber="9">3</superScript>
|
||
and that of
|
||
<materialsCitation id="29E93C9EFFAFFF97FB6DF92EFAE9F948" ID-GBIF-Occurrence="2597591315" box="[1183,1352,1699,1720]" collectionCode="FMNH" pageId="8" pageNumber="9" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
, the next largest
|
||
<taxonomicName id="5E814D40FFAFFF97FC73F933FC67F923" authorityName="Osborn" authorityYear="1905" box="[897,966,1726,1747]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFAFFF97FC73F933FC67F923" box="[897,966,1726,1747]" italics="true" pageId="8" pageNumber="9">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimen, is 617,781 cm
|
||
<superScript id="6EF49B8BFFAFFF97FB17F935FB4EF936" attach="left" box="[1253,1263,1720,1734]" fontSize="6" pageId="8" pageNumber="9">3</superScript>
|
||
. Section moduli therefore suggest that the
|
||
<materialsCitation id="29E93C9EFFAFFF97FBBEF955FB41F91E" ID-GBIF-Occurrence="2597591329" box="[1100,1248,1752,1774]" collectionCode="RSM" pageId="8" pageNumber="9" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
femur had a little more than 1.05 times the bending strength of
|
||
<materialsCitation id="29E93C9EFFAFFF97FAAFF97EFC20F8D2" ID-GBIF-Occurrence="2597591326" collectionCode="FMNH" pageId="8" pageNumber="9" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFAFFF96FCB6F8A5FE84F87D" blockId="8.[810,1450,1618,1934]" lastBlockId="9.[137,778,1431,1933]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
|
||
Snively (2018) suggested that comparing the surface area of the last presacral vertebrae offers another potential method of assessing relative body mass between large theropod specimens of the same taxon. Assuming a roughly horizontal body position, the presacral vertebral column of a theropod dinosaur is analogous to a cantilevered-beam loaded in bending. Gravity exerts a torque anterior to a given vertebra, equal to the distance from the vertebra to the center of mass anterior to it multiplied by the mass of this portion of the body. A vertebral centrum is subject to shear stress from this bending moment. The ability of a vertebral centrum to withstand these forces is likely proportional to the surface area of the anterior centrum face (just as the ability of a femoral shaft to withstand compressive forces is proportional to its cross-sectional area). As measured using the imaging software
|
||
<emphasis id="ABF5EAD1FFAEFF96FDCFF930FD2AF922" box="[573,651,1725,1746]" italics="true" pageId="9" pageNumber="10">imageJ</emphasis>
|
||
(
|
||
<bibRefCitation id="FD104B32FFAEFF96FD6CF930FEA2F91E" author="Schneider CA & Rasband WS & Eliceiri KW" journalOrPublisher="Nat Methods" pageId="9" pageNumber="10" pagination="671 - 675" part="9" refId="ref9642" refString="Schneider CA, Rasband WS, Eliceiri KW. 2012. NIH Image to ImageJ: 25 years of image analysis. Nat Methods 9: 671 - 675." title="NIH Image to ImageJ: 25 years of image analysis" type="journal article" year="2012">Schneider et al., 2012</bibRefCitation>
|
||
), the surface areas of the last presacral anterior centra face in
|
||
<materialsCitation id="29E93C9EFFAEFF96FECDF97FFE7AF8F8" ID-GBIF-Occurrence="2597591327" box="[319,475,1778,1800]" collectionCode="RSM" pageId="9" pageNumber="10" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
and
|
||
<materialsCitation id="29E93C9EFFAEFF96FDD7F97EFD77F8F7" ID-GBIF-Occurrence="2597591360" box="[549,726,1778,1800]" collectionCode="FMNH" pageId="9" pageNumber="10" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
are 650 and 625 cm
|
||
<superScript id="6EF49B8BFFAEFF96FECAF885FEE3F8E6" attach="left" box="[312,322,1800,1814]" fontSize="6" pageId="9" pageNumber="10">2</superScript>
|
||
, respectively. This suggests that the presacral loadings of
|
||
<materialsCitation id="29E93C9EFFAEFF96FEABF8A5FE50F8CD" ID-GBIF-Occurrence="2597591314" box="[345,497,1831,1853]" collectionCode="RSM" pageId="9" pageNumber="10" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
were 1.04 times greater than those of
|
||
<materialsCitation id="29E93C9EFFAEFF96FED8F8CEFE76F8A7" ID-GBIF-Occurrence="2597591324" box="[298,471,1858,1880]" collectionCode="FMNH" pageId="9" pageNumber="10" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
, very near the discrepancy in bending strength calculated based on femoral crosssectional area.
|
||
</paragraph>
|
||
<caption id="CDFE664BFFAEFF96FF6DFDB9FC21FDAF" httpUri="https://zenodo.org/record/3374692/files/figure.png" pageId="9" pageNumber="10" startId="9.[159,189,564,583]" targetBox="[141,1446,203,528]" targetPageId="9">
|
||
<paragraph id="993E36C3FFAEFF96FF6DFDB9FC21FDAF" blockId="9.[137,1450,564,607]" pageId="9" pageNumber="10">
|
||
Fig. 13. The right squamosal of
|
||
<materialsCitation id="29E93C9EFFAEFF96FE4FFDB9FD99FDB7" ID-GBIF-Occurrence="2597591321" box="[445,568,564,583]" collectionCode="RSM" pageId="9" pageNumber="10" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in lateral (
|
||
<emphasis id="ABF5EAD1FFAEFF96FD64FDB9FD05FDB6" bold="true" box="[662,676,564,582]" pageId="9" pageNumber="10">A</emphasis>
|
||
), posterodorsal (
|
||
<emphasis id="ABF5EAD1FFAEFF96FCC9FDB9FCE8FDB6" bold="true" box="[827,841,564,582]" pageId="9" pageNumber="10">B</emphasis>
|
||
), and anterior (
|
||
<emphasis id="ABF5EAD1FFAEFF96FC23FDB9FC41FDB6" bold="true" box="[977,992,564,582]" pageId="9" pageNumber="10">C</emphasis>
|
||
) view (ltf, lateral temporal fenestra; pnc, pneumatic chamber; pos, postorbital process; qc, quadrate cotylus; qjp, quadratojugal process).
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="CDFE664BFFAEFF96FF6DFAB9FDD6FAAF" httpUri="https://zenodo.org/record/3374694/files/figure.png" pageId="9" pageNumber="10" startId="9.[159,189,1332,1351]" targetBox="[143,1444,651,1296]" targetPageId="9">
|
||
<paragraph id="993E36C3FFAEFF96FF6DFAB9FDD6FAAF" blockId="9.[137,1451,1332,1375]" pageId="9" pageNumber="10">
|
||
Fig. 14. The right quadrate of
|
||
<materialsCitation id="29E93C9EFFAEFF96FE5CFAB9FD8BFAB7" ID-GBIF-Occurrence="2597591312" box="[430,554,1332,1351]" collectionCode="RSM" pageId="9" pageNumber="10" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in lateral (
|
||
<emphasis id="ABF5EAD1FFAEFF96FD78FAB8FD39FAB7" bold="true" box="[650,664,1333,1351]" pageId="9" pageNumber="10">A</emphasis>
|
||
) and medial (
|
||
<emphasis id="ABF5EAD1FFAEFF96FCE0FAB8FC81FAB7" bold="true" box="[786,800,1333,1351]" pageId="9" pageNumber="10">B</emphasis>
|
||
) view (ac, articular condyles for the lower jaw; pgw, pterygoid wing; qc, quadrate cotylus; qqf, quadrate-quadratojugal fenestra.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFAEFF96FCB6FA1AFBBDFA0E" blockId="9.[810,1450,1431,1534]" pageId="9" pageNumber="10">
|
||
Size comparisons of
|
||
<materialsCitation id="29E93C9EFFAEFF96FBCFFA15FB7AFA5D" ID-GBIF-Occurrence="2597591303" box="[1085,1243,1431,1453]" collectionCode="RSM" pageId="9" pageNumber="10" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
with the largest specimens of other giant theropod genera were also made using the femoral circumference method of
|
||
<bibRefCitation id="FD104B32FFAEFF96FAC9FA40FC0DFA0E" author="Campione NE & Evans DC & Brown CM & Carrano MT" journalOrPublisher="Methods Ecol Evol" pageId="9" pageNumber="10" pagination="913 - 923" part="5" refId="ref8337" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol Evol 5: 913 - 923." title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">Campione et al. (2014)</bibRefCitation>
|
||
(
|
||
<tableCitation id="D4030378FFAEFF96FC49FA6AFBAEFA0E" box="[955,1039,1511,1534]" captionStart="TABLE 2" captionStartId="14.[168,252,205,226]" captionTargetBox="[137,1450,274,647]" captionTargetPageId="14" captionText="TABLE 2. Comparison of femoral dimensions and estimated body mass of the largest theropods ±25% (mass estimation based on the formula of Campione et al. (2014))" httpUri="http://table.plazi.org/id/CDFE664BFFA9FF91FF5AFF40FBCCFF0A" pageId="9" pageNumber="10" tableUuid="CDFE664BFFA9FF91FF5AFF40FBCCFF0A">Table 2</tableCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="D19B6548FFAEFF90FCD8F9BAFEFFFA0B" lastPageId="15" lastPageNumber="16" pageId="9" pageNumber="10" type="discussion">
|
||
<paragraph id="993E36C3FFAEFF96FCD8F9BAFC65F99C" blockId="9.[810,1450,1591,1934]" pageId="9" pageNumber="10">
|
||
<heading id="C27681AFFFAEFF96FCD8F9BAFC65F99C" bold="true" fontSize="10" level="2" pageId="9" pageNumber="10" reason="0">
|
||
<emphasis id="ABF5EAD1FFAEFF96FCD8F9BAFC65F99C" bold="true" pageId="9" pageNumber="10">
|
||
The Challenge of Sizing
|
||
<taxonomicName id="5E814D40FFAEFF96FB95F9B5FC65F99C" authorityName="Stromer" authorityYear="1915" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFAEFF96FB95F9B5FC65F99C" bold="true" italics="true" pageId="9" pageNumber="10">Spinosaurus aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFAEFF96FCB6F9F3FA80F87E" blockId="9.[810,1450,1591,1934]" pageId="9" pageNumber="10">
|
||
Although using MFC to estimate bipedal dinosaur body mass has emerged as a leading technique and has been quantitatively calibrated and evaluated through comparisons with modern animals (
|
||
<bibRefCitation id="FD104B32FFAEFF96FB85F95FFA21F918" author="Campione NE & Evans DC & Brown CM & Carrano MT" box="[1143,1408,1746,1768]" journalOrPublisher="Methods Ecol Evol" pageId="9" pageNumber="10" pagination="913 - 923" part="5" refId="ref8337" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol Evol 5: 913 - 923." title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">Campione et al., 2014</bibRefCitation>
|
||
), it should be remembered that the approach makes several assumptions about the biology of the taxa it is applied to. The theropod
|
||
<taxonomicName id="5E814D40FFAEFF96FC0CF8A8FB05F8CA" authorityName="Stromer" authorityYear="1915" box="[1022,1188,1829,1850]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFAEFF96FC0CF8A8FB05F8CA" box="[1022,1188,1829,1850]" italics="true" pageId="9" pageNumber="10">S. aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
presents a particular challenge for this technique, because the locomotion and habits of these taxa are thought by many to fundamentally differ from that of most other theropods.
|
||
</paragraph>
|
||
<caption id="CDFE664BFFADFF95FF6DF99FFE93F9CD" httpUri="https://zenodo.org/record/3374696/files/figure.png" pageId="10" pageNumber="11" startId="10.[159,189,1554,1573]" targetBox="[143,1444,203,1518]" targetPageId="10">
|
||
<paragraph id="993E36C3FFADFF95FF6DF99FFE93F9CD" blockId="10.[137,1451,1554,1597]" pageId="10" pageNumber="11">
|
||
Fig. 15. The right and left quadratojugal of
|
||
<materialsCitation id="29E93C9EFFADFF95FDDAF99FFD04F9D5" ID-GBIF-Occurrence="2597591356" box="[552,677,1554,1573]" collectionCode="RSM" pageId="10" pageNumber="11" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in lateral (
|
||
<emphasis id="ABF5EAD1FFADFF95FCF4F99EFC91F9D5" bold="true" box="[774,816,1555,1573]" pageId="10" pageNumber="11">A, C</emphasis>
|
||
) and medial (
|
||
<emphasis id="ABF5EAD1FFADFF95FC5EF99EFC77F9D5" bold="true" box="[940,982,1555,1573]" pageId="10" pageNumber="11">B, D</emphasis>
|
||
) view, respectively (ltf, lateral temporal fenestra; qf, quadrate fenestra).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFADFF94FF56F9E0FC0FFA58" blockId="10.[137,778,1645,1934]" lastBlockId="11.[810,1451,1239,1661]" lastPageId="11" lastPageNumber="12" pageId="10" pageNumber="11">
|
||
The
|
||
<taxonomicName id="5E814D40FFADFF95FF2FF9E3FE22F973" authorityName="Stromer" authorityYear="1915" box="[221,387,1646,1667]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFADFF95FF2FF9E3FE22F973" box="[221,387,1646,1667]" italics="true" pageId="10" pageNumber="11">S. aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
mass estimation here presented should be interpreted with great caution, as many lines of evidence suggest it may be a substantial underestimation. This estimation was arrived at by using the reconstruction of
|
||
<taxonomicName id="5E814D40FFADFF95FF54F955FEE7F91E" authorityName="Stromer" authorityYear="1915" box="[166,326,1752,1774]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFADFF95FF54F955FEE7F91E" box="[166,326,1752,1774]" italics="true" pageId="10" pageNumber="11">S. aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
created by
|
||
<bibRefCitation id="FD104B32FFADFF95FE3FF955FD1AF91E" author="Ibrahim N & Sereno PC & Dal Sasso C & Maganuco S & Fabbri M & Martill DM & Zouhri S & Myhrvold N & Iurino DA" box="[461,699,1752,1774]" journalOrPublisher="Science" pageId="10" pageNumber="11" pagination="1613 - 1616" part="345" refId="ref9341" refString="Ibrahim N, Sereno PC, Dal Sasso C, Maganuco S, Fabbri M, Martill DM, Zouhri S, Myhrvold N, Iurino DA. 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science 345: 1613 - 1616." title="Semiaquatic adaptations in a giant predatory dinosaur" type="journal article" year="2014">Ibrahim et al. (2014)</bibRefCitation>
|
||
, which isometrically scaled know material, including the femur of specimen FSAC-KK 11888, to the size of MSNM V4047—a specimen that includes only a skull. Whether or not isometric scaling is reasonable to assume across intraspecific theropod specimens of great size variability is a debatable issue. The MFC method assumes that MFC relates directly to minimal femoral cross-sectional area, and, therefore, to the strength of the femur.
|
||
<bibRefCitation id="FD104B32FFADFF95FBBAF905FA86F96E" author="Ibrahim N & Sereno PC & Dal Sasso C & Maganuco S & Fabbri M & Martill DM & Zouhri S & Myhrvold N & Iurino DA" box="[1096,1319,1672,1694]" journalOrPublisher="Science" pageId="10" pageNumber="11" pagination="1613 - 1616" part="345" refId="ref9341" refString="Ibrahim N, Sereno PC, Dal Sasso C, Maganuco S, Fabbri M, Martill DM, Zouhri S, Myhrvold N, Iurino DA. 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science 345: 1613 - 1616." title="Semiaquatic adaptations in a giant predatory dinosaur" type="journal article" year="2014">Ibrahim et al. (2014)</bibRefCitation>
|
||
reported on the internal femoral shaft structure of
|
||
<taxonomicName id="5E814D40FFADFF95FB25F92EFAD3F948" authorityName="Stromer" authorityYear="1915" box="[1239,1394,1699,1720]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFADFF95FB25F92EFAD3F948" box="[1239,1394,1699,1720]" italics="true" pageId="10" pageNumber="11">S. aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
, and found it to be unique among theropods in lacking an open medullary cavity. This may have made the cross-sectional strength of
|
||
<taxonomicName id="5E814D40FFADFF95FC51F97EFB9AF8F8" authorityName="Stromer" authorityYear="1915" box="[931,1083,1779,1800]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFADFF95FC51F97EFB9AF8F8" box="[931,1083,1779,1800]" italics="true" pageId="10" pageNumber="11">S. aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
femora proportionally greater, relative to femoral circumference. Various researchers have also postulated that
|
||
<taxonomicName id="5E814D40FFADFF95FBE2F8A5FB10F8CE" authorityName="Stromer" authorityYear="1915" box="[1040,1201,1832,1854]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFADFF95FBE2F8A5FB10F8CE" box="[1040,1201,1832,1854]" italics="true" pageId="10" pageNumber="11">S. aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
was partially quadrupedal or even obligately quadrupedal (
|
||
<bibRefCitation id="FD104B32FFADFF95FB09F8CEFCC2F883" author="Ibrahim N & Sereno PC & Dal Sasso C & Maganuco S & Fabbri M & Martill DM & Zouhri S & Myhrvold N & Iurino DA" journalOrPublisher="Science" pageId="10" pageNumber="11" pagination="1613 - 1616" part="345" refId="ref9341" refString="Ibrahim N, Sereno PC, Dal Sasso C, Maganuco S, Fabbri M, Martill DM, Zouhri S, Myhrvold N, Iurino DA. 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science 345: 1613 - 1616." title="Semiaquatic adaptations in a giant predatory dinosaur" type="journal article" year="2014">Ibrahim et al., 2014</bibRefCitation>
|
||
), again unlike all other theropods. If so, the femur of
|
||
<taxonomicName id="5E814D40FFADFF95FCBBF8F5FC4EF87E" authorityName="Stromer" authorityYear="1915" box="[841,1007,1912,1934]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFADFF95FCBBF8F5FC4EF87E" box="[841,1007,1912,1934]" italics="true" pageId="10" pageNumber="11">S. aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
would not have been the sole weight bearer and a proportionally greater body mass could be expected relative to femoral circumference. Finally,
|
||
<bibRefCitation id="FD104B32FFACFF94FCD8FA81FBB9FAD2" author="Ibrahim N & Sereno PC & Dal Sasso C & Maganuco S & Fabbri M & Martill DM & Zouhri S & Myhrvold N & Iurino DA" box="[810,1048,1292,1314]" journalOrPublisher="Science" pageId="11" pageNumber="12" pagination="1613 - 1616" part="345" refId="ref9341" refString="Ibrahim N, Sereno PC, Dal Sasso C, Maganuco S, Fabbri M, Martill DM, Zouhri S, Myhrvold N, Iurino DA. 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science 345: 1613 - 1616." title="Semiaquatic adaptations in a giant predatory dinosaur" type="journal article" year="2014">Ibrahim et al. (2014)</bibRefCitation>
|
||
suggested that
|
||
<taxonomicName id="5E814D40FFACFF94FB20FA80FAD7FAD2" authorityName="Stromer" authorityYear="1915" box="[1234,1398,1293,1314]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFACFF94FB20FA80FAD7FAD2" box="[1234,1398,1293,1314]" italics="true" pageId="11" pageNumber="12">S. aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
was adapted for a semiaquatic lifestyle, with greatly reduced hind limb proportions. Being semiaquatic,
|
||
<taxonomicName id="5E814D40FFACFF94FAFBFACFFA0AFAA7" authorityName="Stromer" authorityYear="1915" box="[1289,1451,1346,1367]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFACFF94FAFBFACFFA0AFAA7" box="[1289,1451,1346,1367]" italics="true" pageId="11" pageNumber="12">S. aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
may have frequently relied on buoyancy in water to support its weight, and, if so, its femoral proportions may be misleading.
|
||
</paragraph>
|
||
<caption id="CDFE664BFFACFF94FF6DFBA3FC7AFB79" httpUri="https://zenodo.org/record/3374698/files/figure.png" pageId="11" pageNumber="12" startId="11.[159,189,1070,1089]" targetBox="[142,1445,203,1034]" targetPageId="11">
|
||
<paragraph id="993E36C3FFACFF94FF6DFBA3FC7AFB79" blockId="11.[137,1451,1070,1161]" pageId="11" pageNumber="12">
|
||
Fig. 16. The braincase of
|
||
<materialsCitation id="29E93C9EFFACFF94FE62FBA3FDAFFBB1" ID-GBIF-Occurrence="2597591345" box="[400,526,1070,1089]" collectionCode="RSM" pageId="11" pageNumber="12" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in posterior (
|
||
<emphasis id="ABF5EAD1FFACFF94FD7EFBA2FD3BFBB1" bold="true" box="[652,666,1071,1089]" pageId="11" pageNumber="12">A</emphasis>
|
||
) and right lateral (
|
||
<emphasis id="ABF5EAD1FFACFF94FCB7FBA2FCF2FBB1" bold="true" box="[837,851,1071,1089]" pageId="11" pageNumber="12">B</emphasis>
|
||
) view (bt, basal tuber; btg, basiterygoid; cb, cultriform base; bo, basioccipital; bpt, basipterygoid process; bt, basal tubera; eo, exoccipital; fm foramen magnum; fo, fenestra ovalis; fr, frontal; nc, nuchal crest; oc, occipital condyle; on, orbital notch; par, parietal; ptr, posterior tympanic recess; pop, paraoccipital process; pos, postorbital suture; sc, sagittal crest; scr, subcondylar recess; soc, supraoccipital; sph, sphenethmoid; V, X-XII, cranial nerves).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFACFF94FCB6FA21FACBF98C" blockId="11.[810,1451,1239,1661]" pageId="11" pageNumber="12">
|
||
Furthermore, if Ibrahim et al.’s
|
||
<bibRefCitation id="FD104B32FFACFF94FB60FA21FACBFA32" author="Ibrahim N & Sereno PC & Dal Sasso C & Maganuco S & Fabbri M & Martill DM & Zouhri S & Myhrvold N & Iurino DA" box="[1170,1386,1452,1474]" journalOrPublisher="Science" pageId="11" pageNumber="12" pagination="1613 - 1616" part="345" refId="ref9341" refString="Ibrahim N, Sereno PC, Dal Sasso C, Maganuco S, Fabbri M, Martill DM, Zouhri S, Myhrvold N, Iurino DA. 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science 345: 1613 - 1616." title="Semiaquatic adaptations in a giant predatory dinosaur" type="journal article" year="2014">Ibrahim et al. (2014)</bibRefCitation>
|
||
interpretation is correct and the hindlimbs of
|
||
<taxonomicName id="5E814D40FFACFF94FB28FA4AFACEFA2D" authorityName="Stromer" authorityYear="1915" box="[1242,1391,1479,1501]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFACFF94FB28FA4AFACEFA2D" box="[1242,1391,1479,1501]" italics="true" pageId="11" pageNumber="12">S. aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
were reduced in all dimensions relative to its body mass, mass calculations based on MFC would not be expected to yield accurate estimations. As such, it may not, at present, be possible to reasonably estimate the mass of
|
||
<emphasis id="ABF5EAD1FFACFF94FB5EF9BFFAE8F9B7" box="[1196,1353,1586,1607]" italics="true" pageId="11" pageNumber="12">
|
||
<taxonomicName id="5E814D40FFACFF94FB5EF9BFFAE4F9B7" authorityName="Stromer" authorityYear="1915" box="[1196,1349,1586,1607]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="aegyptiacus">S. aegyptiacus</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
See also
|
||
<bibRefCitation id="FD104B32FFACFF94FCD8F9C1FBD6F993" author="Therrien F & Henderson DM" box="[810,1143,1612,1635]" journalOrPublisher="J Vertebr Paleontol" pageId="11" pageNumber="12" pagination="108 - 115" part="27" refId="ref9743" refString="Therrien F, Henderson DM. 2007. My theropod is bigger than yours … or not: estimating body size from skull length in theropods. J Vertebr Paleontol 27: 108 - 115." title="My theropod is bigger than yours … or not: estimating body size from skull length in theropods" type="journal article" year="2007">Therrien and Henderson (2007))</bibRefCitation>
|
||
for a further discussion of the challenge of estimating the body mass of MSNM V4047.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFACFF94FCD8F923FAEEF936" blockId="11.[810,1450,1710,1934]" box="[810,1359,1710,1734]" pageId="11" pageNumber="12">
|
||
<heading id="C27681AFFFACFF94FCD8F923FAEEF936" bold="true" box="[810,1359,1710,1734]" fontSize="10" level="2" pageId="11" pageNumber="12" reason="0">
|
||
<emphasis id="ABF5EAD1FFACFF94FCD8F923FAEEF936" bold="true" box="[810,1359,1710,1734]" pageId="11" pageNumber="12">
|
||
Comparisons with Other
|
||
<taxonomicName id="5E814D40FFACFF94FB81F922FB1DF936" authorityName="Osborn" authorityYear="1905" box="[1139,1212,1711,1734]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFACFF94FB81F922FB1DF936" bold="true" box="[1139,1212,1711,1734]" italics="true" pageId="11" pageNumber="12">T. rex</emphasis>
|
||
</taxonomicName>
|
||
Specimens
|
||
</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFACFF93FCB6F955FAF8FBF9" blockId="11.[810,1450,1710,1934]" lastBlockId="12.[810,1450,878,1219]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">
|
||
Substantial variation in skeletal proportions has been previously recognized among adult specimens of
|
||
<taxonomicName id="5E814D40FFACFF94FA92F97EFA05F8F8" authorityName="Osborn" authorityYear="1905" box="[1376,1444,1779,1800]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFACFF94FA92F97EFA05F8F8" box="[1376,1444,1779,1800]" italics="true" pageId="11" pageNumber="12">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, and, although strong statistical support is lacking (
|
||
<bibRefCitation id="FD104B32FFACFF94FCC3F8A5FC65F8CE" author="Mallon JC" box="[817,964,1832,1854]" journalOrPublisher="Paleobiology" pageId="11" pageNumber="12" pagination="1 - 13" part="43" refId="ref9519" refString="Mallon JC. 2017. Recognizing sexual dimorphism in the fossil record: lessons from nonavian dinosaurs. Paleobiology 43: 1 - 13." title="Recognizing sexual dimorphism in the fossil record: lessons from nonavian dinosaurs" type="journal article" year="2017">Mallon, 2017</bibRefCitation>
|
||
), some researches have grouped
|
||
<taxonomicName id="5E814D40FFACFF94FADFF8A4FACDF8CE" authorityName="Osborn" authorityYear="1905" box="[1325,1388,1833,1854]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFACFF94FADFF8A4FACDF8CE" box="[1325,1388,1833,1854]" italics="true" pageId="11" pageNumber="12">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens into two morphotypes: a gracile form, with more elongate and slender skeletons, and a robust form with stockier proportions and greater estimated mass relative to body length and height (
|
||
<bibRefCitation id="FD104B32FFABFF93FE1AFCE3FCA4FC73" author="Larson P & Frey E." box="[488,773,877,899]" journalOrPublisher="J Vertebr Paleontol" pageId="12" pageNumber="13" pagination="38 A" part="12" refId="ref9492" refString="Larson P, Frey E. 1992. Sexual dimorphism in the abundant Upper Cretaceous theropod Tyrannosaurus rex. J Vertebr Paleontol 12: 38 A." title="Sexual dimorphism in the abundant Upper Cretaceous theropod Tyrannosaurus rex" type="journal article" year="1992">Larson and Frey, 1992</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFABFF93FF7BFC04FE8DFC6E" author="Larson P." box="[137,300,904,926]" journalOrPublisher="Bloomington, IN: Indiana University Press." pageId="12" pageNumber="13" pagination="103 - 130" refId="ref9463" refString="Larson P. 2008 b. Variation and sexual dimorphism in Tyrannosaurus rex. Tyrannosaurus rex: Bloomington, IN: Indiana University Press. 103 - 130." title="Variation and sexual dimorphism in Tyrannosaurus rex." type="book chapter" volumeTitle="Tyrannosaurus rex" year="2008">Larson, 2008b</bibRefCitation>
|
||
). This purported dichotomy is contentious, with arguments over whether the differences reflect sexual dimorphism or are simply the manifestation of a gradational ontogenetic continuum and/or individual variation (
|
||
<bibRefCitation id="FD104B32FFABFF93FF63FC7EFEEBFBF8" author="Carpenter K." box="[145,330,1011,1032]" journalOrPublisher="Cambridge, UK: Cambridge University Press" pageId="12" pageNumber="13" pagination="141 - 145" refId="ref8375" refString="Carpenter K. 1990. Variation in Tyrannosaurus rex. Dinosaur systematics: perspectiVes and approaches. Cambridge, UK: Cambridge University Press. p 141 - 145." title="Variation in Tyrannosaurus rex. Dinosaur systematics: perspectiVes and approaches" type="book chapter" year="1990">Carpenter, 1990</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFABFF93FEAFFC7EFDCAFBF8" author="Larson P & Frey E." box="[349,619,1011,1033]" journalOrPublisher="J Vertebr Paleontol" pageId="12" pageNumber="13" pagination="38 A" part="12" refId="ref9492" refString="Larson P, Frey E. 1992. Sexual dimorphism in the abundant Upper Cretaceous theropod Tyrannosaurus rex. J Vertebr Paleontol 12: 38 A." title="Sexual dimorphism in the abundant Upper Cretaceous theropod Tyrannosaurus rex" type="journal article" year="1992">Larson and Frey, 1992</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFABFF93FD9DFC7EFCA4FBF8" author="Larson P." box="[623,773,1011,1032]" editor="Rosenberg GD & Wolberg DL" journalOrPublisher="Knoxville: The Paleontological Society" pageId="12" pageNumber="13" pagination="139 - 155" refId="ref9386" refString="Larson P. 1994. Tyrannosaurus sex. In: Rosenberg GD, Wolberg DL, editors. Dino fest, The Paleontological Society Special Publication 7. Knoxville: The Paleontological Society. p 139 - 155." title="Tyrannosaurus sex" type="book chapter" volumeTitle="Dino fest, The Paleontological Society Special Publication 7" year="1994">Larson, 1994</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFABFF93FF7BFB83FE8BFBD3" author="Larson P." box="[137,298,1037,1059]" journalOrPublisher="Bloomington, IN: Indiana University Press." pageId="12" pageNumber="13" pagination="103 - 130" refId="ref9463" refString="Larson P. 2008 b. Variation and sexual dimorphism in Tyrannosaurus rex. Tyrannosaurus rex: Bloomington, IN: Indiana University Press. 103 - 130." title="Variation and sexual dimorphism in Tyrannosaurus rex." type="book chapter" volumeTitle="Tyrannosaurus rex" year="2008">Larson, 2008b</bibRefCitation>
|
||
). Those in the former camp speculate that
|
||
<taxonomicName id="5E814D40FFABFF93FF7BFBA4FF6EFBCE" authorityName="Osborn" authorityYear="1905" box="[137,207,1065,1086]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFABFF93FF7BFBA4FF6EFBCE" box="[137,207,1065,1086]" italics="true" pageId="12" pageNumber="13">T. rex</emphasis>
|
||
</taxonomicName>
|
||
shows reversed sexual dimorphism (
|
||
<bibRefCitation id="FD104B32FFABFF93FD74FBA5FEA1FBA8" author="Larson P & Frey E." journalOrPublisher="J Vertebr Paleontol" pageId="12" pageNumber="13" pagination="38 A" part="12" refId="ref9492" refString="Larson P, Frey E. 1992. Sexual dimorphism in the abundant Upper Cretaceous theropod Tyrannosaurus rex. J Vertebr Paleontol 12: 38 A." title="Sexual dimorphism in the abundant Upper Cretaceous theropod Tyrannosaurus rex" type="journal article" year="1992">Larson and Frey, 1992</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFABFF93FEFEFBCEFE0BFBA9" author="Larson P." box="[268,426,1091,1113]" journalOrPublisher="Bloomington, IN: Indiana University Press." pageId="12" pageNumber="13" pagination="103 - 130" refId="ref9463" refString="Larson P. 2008 b. Variation and sexual dimorphism in Tyrannosaurus rex. Tyrannosaurus rex: Bloomington, IN: Indiana University Press. 103 - 130." title="Variation and sexual dimorphism in Tyrannosaurus rex." type="book chapter" volumeTitle="Tyrannosaurus rex" year="2008">Larson, 2008b</bibRefCitation>
|
||
). As in many reptiles and birds, it is suggested that
|
||
<taxonomicName id="5E814D40FFABFF93FE90FBD3FE00FB83" authorityName="Osborn" authorityYear="1905" box="[354,417,1118,1139]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFABFF93FE90FBD3FE00FB83" box="[354,417,1118,1139]" italics="true" pageId="12" pageNumber="13">T. rex</emphasis>
|
||
</taxonomicName>
|
||
females displayed a more robust morphotype to meet the physiological rigors of oviposition (
|
||
<bibRefCitation id="FD104B32FFABFF93FF62FB1EFE84FB58" author="Larson P." box="[144,293,1171,1192]" editor="Rosenberg GD & Wolberg DL" journalOrPublisher="Knoxville: The Paleontological Society" pageId="12" pageNumber="13" pagination="139 - 155" refId="ref9386" refString="Larson P. 1994. Tyrannosaurus sex. In: Rosenberg GD, Wolberg DL, editors. Dino fest, The Paleontological Society Special Publication 7. Knoxville: The Paleontological Society. p 139 - 155." title="Tyrannosaurus sex" type="book chapter" volumeTitle="Dino fest, The Paleontological Society Special Publication 7" year="1994">Larson, 1994</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFABFF93FECBFB1EFE7DFB59" author="Larson P." box="[313,476,1171,1193]" journalOrPublisher="Bloomington, IN: Indiana University Press." pageId="12" pageNumber="13" pagination="103 - 130" refId="ref9463" refString="Larson P. 2008 b. Variation and sexual dimorphism in Tyrannosaurus rex. Tyrannosaurus rex: Bloomington, IN: Indiana University Press. 103 - 130." title="Variation and sexual dimorphism in Tyrannosaurus rex." type="book chapter" volumeTitle="Tyrannosaurus rex" year="2008">Larson, 2008b</bibRefCitation>
|
||
).
|
||
<bibRefCitation id="FD104B32FFABFF93FE04FB1EFCA8FB59" author="Schweitzer MH & Wittmeyer JL & Horner JR" box="[502,777,1171,1193]" journalOrPublisher="Science" pageId="12" pageNumber="13" pagination="1456 - 1460" part="308" refId="ref9672" refString="Schweitzer MH, Wittmeyer JL, Horner JR. 2005. Gender-specific reproductive tissue in ratites and Tyrannosaurus rex. Science 308: 1456 - 1460." title="Gender-specific reproductive tissue in ratites and Tyrannosaurus rex" type="journal article" year="2005">Schweitzer et al. (2005)</bibRefCitation>
|
||
reported medullary bone (calcium deposition stores in the cancellous and medullary cavities of birds for use during oviposition) in the femur of
|
||
<materialsCitation id="29E93C9EFFABFF93FBC8FC05FB11FC6E" ID-GBIF-Occurrence="2597591362" box="[1082,1200,904,926]" collectionCode="MOR" pageId="12" pageNumber="13" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, a specimen previously assigned to the robust morphotype (
|
||
<bibRefCitation id="FD104B32FFABFF93FB54FC2EFA9FFC49" author="Larson P." box="[1190,1342,931,953]" journalOrPublisher="Bloomington, IN: Indiana University Press." pageId="12" pageNumber="13" pagination="103 - 130" refId="ref9463" refString="Larson P. 2008 b. Variation and sexual dimorphism in Tyrannosaurus rex. Tyrannosaurus rex: Bloomington, IN: Indiana University Press. 103 - 130." title="Variation and sexual dimorphism in Tyrannosaurus rex." type="book chapter" volumeTitle="Tyrannosaurus rex" year="2008">Larson, 2008b</bibRefCitation>
|
||
). Those in the latter camp have noted that large tyrannosaurids had a highly gracile body form early in ontogeny and assumed more robust features as they matured (
|
||
<bibRefCitation id="FD104B32FFABFF93FB3AFC7EFAEFFBF9" author="Currie PJ" box="[1224,1358,1011,1033]" journalOrPublisher="Can J Earth Sci" pageId="12" pageNumber="13" pagination="651 - 665" part="40" refId="ref8478" refString="Currie PJ. 2003. Allometric growth in tyrannosaurids (Dinosauria: Theropoda) from the Upper Cretaceous of North America and Asia. Can J Earth Sci 40: 651 - 665." title="Allometric growth in tyrannosaurids (Dinosauria: Theropoda) from the Upper Cretaceous of North America and Asia" type="journal article" year="2003">Currie, 2003</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="CDFE664BFFACFF94FF6DF8EFFD62F87D" httpUri="https://zenodo.org/record/3374700/files/figure.png" pageId="11" pageNumber="12" startId="11.[159,189,1890,1909]" targetBox="[138,778,1257,1854]" targetPageId="11">
|
||
<paragraph id="993E36C3FFACFF94FF6DF8EFFD62F87D" blockId="11.[137,777,1890,1933]" pageId="11" pageNumber="12">
|
||
Fig. 17. The right and left ectopterygoid of
|
||
<materialsCitation id="29E93C9EFFACFF94FDECF8EFFD3BF885" ID-GBIF-Occurrence="2597591358" box="[542,666,1890,1909]" collectionCode="RSM" pageId="11" pageNumber="12" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in lateral (
|
||
<emphasis id="ABF5EAD1FFACFF94FD05F8EEFCA4F885" bold="true" box="[759,773,1891,1909]" pageId="11" pageNumber="12">A</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFACFF94FF7BF8F6FF39F87D" bold="true" box="[137,152,1915,1933]" pageId="11" pageNumber="12">C</emphasis>
|
||
) and medial (
|
||
<emphasis id="ABF5EAD1FFACFF94FEE2F8F6FEBFF87D" bold="true" box="[272,286,1915,1933]" pageId="11" pageNumber="12">B</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFACFF94FEDBF8F6FE99F87D" bold="true" box="[297,312,1915,1933]" pageId="11" pageNumber="12">D</emphasis>
|
||
) view, respectively (pn, pneumatic opening).
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="CDFE664BFFABFF93FF6DFCA0FAD4FCB0" box="[159,1397,813,832]" httpUri="https://zenodo.org/record/3374702/files/figure.png" pageId="12" pageNumber="13" startId="12.[159,189,813,832]" targetBox="[141,1446,203,777]" targetPageId="12">
|
||
<paragraph id="993E36C3FFABFF93FF6DFCA0FAD4FCB0" blockId="12.[159,1397,813,832]" box="[159,1397,813,832]" pageId="12" pageNumber="13">
|
||
Fig. 18. The right and left dentary of
|
||
<materialsCitation id="29E93C9EFFABFF93FE11FCA0FDFCFCB0" ID-GBIF-Occurrence="2597591347" box="[483,605,813,832]" collectionCode="RSM" pageId="12" pageNumber="13" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in lateral (
|
||
<emphasis id="ABF5EAD1FFABFF93FD4BFCA0FD67FCCF" bold="true" box="[697,710,813,831]" pageId="12" pageNumber="13">A</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFABFF93FD20FCA0FD40FCCF" bold="true" box="[722,737,813,831]" pageId="12" pageNumber="13">C</emphasis>
|
||
) and medial (
|
||
<emphasis id="ABF5EAD1FFABFF93FCAAFCA0FCC7FCCF" bold="true" box="[856,870,813,831]" pageId="12" pageNumber="13">B</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFABFF93FC83FCA0FC21FCCF" bold="true" box="[881,896,813,831]" pageId="12" pageNumber="13">D</emphasis>
|
||
) view, respectively (1-14, alveoli; mf, mylohyal foramen).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFABFF92FCB6FB83FB3AFE48" blockId="12.[810,1450,878,1219]" lastBlockId="13.[810,1450,205,920]" lastPageId="13" lastPageNumber="14" pageId="12" pageNumber="13">
|
||
In most skeletal dimensions,
|
||
<materialsCitation id="29E93C9EFFABFF93FB60FB83FA8BFBD3" ID-GBIF-Occurrence="2597591309" box="[1170,1322,1037,1059]" collectionCode="RSM" pageId="12" pageNumber="13" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
is close to
|
||
<materialsCitation id="29E93C9EFFABFF93FCD8FBA4FC75FBCD" ID-GBIF-Occurrence="2597591331" box="[810,980,1064,1086]" collectionCode="FMNH" pageId="12" pageNumber="13" specimenCode="FMNH PR2081">FMNH PR 2081</materialsCitation>
|
||
.
|
||
<materialsCitation id="29E93C9EFFABFF93FC2FFBA5FBCDFBCE" ID-GBIF-Occurrence="2597591318" box="[989,1132,1064,1086]" collectionCode="RSM" pageId="12" pageNumber="13" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
exceeds all known
|
||
<taxonomicName id="5E814D40FFABFF93FAC2FBA4FACBFBCE" authorityName="Osborn" authorityYear="1905" box="[1328,1386,1065,1086]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFABFF93FAC2FBA4FACBFBCE" box="[1328,1386,1065,1086]" italics="true" pageId="12" pageNumber="13">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens (including
|
||
<materialsCitation id="29E93C9EFFABFF93FC16FBCEFB3CFBA9" ID-GBIF-Occurrence="2597591365" box="[996,1181,1091,1113]" collectionCode="FMNH" pageId="12" pageNumber="13" specimenCode="FMNH PR2081">FMNH PR 2081</materialsCitation>
|
||
) in numerous measurements, including scapula blade width, ilium length, proximal femur width, and tibia shaft width. In additional measurement (including dentary tooth row and femur length)
|
||
<materialsCitation id="29E93C9EFFABFF93FA87FB1EFCDEFB33" ID-GBIF-Occurrence="2597591361" collectionCode="RSM" pageId="12" pageNumber="13" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
notably exceeds all
|
||
<taxonomicName id="5E814D40FFABFF93FBA6FB23FB31FB33" authorityName="Osborn" authorityYear="1905" box="[1108,1168,1198,1219]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFABFF93FBA6FB23FB31FB33" box="[1108,1168,1198,1219]" italics="true" pageId="12" pageNumber="13">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens that have been previously categorized as robust (
|
||
<bibRefCitation id="FD104B32FFAAFF92FB77FF40FABDFF13" author="Larson P." box="[1157,1308,205,227]" journalOrPublisher="Bloomington, IN: Indiana University Press." pageId="13" pageNumber="14" pagination="103 - 130" refId="ref9463" refString="Larson P. 2008 b. Variation and sexual dimorphism in Tyrannosaurus rex. Tyrannosaurus rex: Bloomington, IN: Indiana University Press. 103 - 130." title="Variation and sexual dimorphism in Tyrannosaurus rex." type="book chapter" volumeTitle="Tyrannosaurus rex" year="2008">Larson, 2008b</bibRefCitation>
|
||
). However, in many length measurements (proximodistal femoral, tibial and jaw length)
|
||
<materialsCitation id="29E93C9EFFAAFF92FC22FE8FFBFEFEE8" ID-GBIF-Occurrence="2597591302" box="[976,1119,258,280]" collectionCode="RSM" pageId="13" pageNumber="14" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
is exceeded by some individuals previously categorized as gracile (
|
||
<bibRefCitation id="FD104B32FFAAFF92FB78FE90FA83FEC3" author="Larson P." box="[1162,1314,285,307]" journalOrPublisher="Bloomington, IN: Indiana University Press." pageId="13" pageNumber="14" pagination="103 - 130" refId="ref9463" refString="Larson P. 2008 b. Variation and sexual dimorphism in Tyrannosaurus rex. Tyrannosaurus rex: Bloomington, IN: Indiana University Press. 103 - 130." title="Variation and sexual dimorphism in Tyrannosaurus rex." type="book chapter" volumeTitle="Tyrannosaurus rex" year="2008">Larson, 2008b</bibRefCitation>
|
||
). Whether or not a robust/gracile dichotomy exists among
|
||
<taxonomicName id="5E814D40FFAAFF92FB07FEB5FA91FEBD" authorityName="Osborn" authorityYear="1905" box="[1269,1328,312,333]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFAAFF92FB07FEB5FA91FEBD" box="[1269,1328,312,333]" italics="true" pageId="13" pageNumber="14">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, these comparisons indicate that
|
||
<materialsCitation id="29E93C9EFFAAFF92FBE1FEDFFB02FE98" ID-GBIF-Occurrence="2597591316" box="[1043,1187,338,360]" collectionCode="RSM" pageId="13" pageNumber="14" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
was a large and robustly proportioned individual, but likely with a shorter total hipheight and snout-vent length than other known specimens showing more elongate proportions.
|
||
</paragraph>
|
||
<caption id="CDFE664BFFABFF93FF6DF8EFFDC3F87D" httpUri="https://zenodo.org/record/3374704/files/figure.png" pageId="12" pageNumber="13" startId="12.[159,189,1890,1909]" targetBox="[140,1445,1261,1851]" targetPageId="12">
|
||
<paragraph id="993E36C3FFABFF93FF6DF8EFFDC3F87D" blockId="12.[137,1451,1890,1933]" pageId="12" pageNumber="13">
|
||
Fig. 19. The right and left surangular of
|
||
<materialsCitation id="29E93C9EFFABFF93FDE6F8EFFD33F885" ID-GBIF-Occurrence="2597591304" box="[532,658,1890,1909]" collectionCode="RSM" pageId="12" pageNumber="13" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
in lateral (
|
||
<emphasis id="ABF5EAD1FFABFF93FD04F8EEFCA2F885" bold="true" box="[758,771,1891,1909]" pageId="12" pageNumber="13">A</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFABFF93FCE3F8EEFC81F885" bold="true" box="[785,800,1891,1909]" pageId="12" pageNumber="13">C</emphasis>
|
||
) and medial (
|
||
<emphasis id="ABF5EAD1FFABFF93FC52F8EEFC0FF885" bold="true" box="[928,942,1891,1909]" pageId="12" pageNumber="13">B</emphasis>
|
||
,
|
||
<emphasis id="ABF5EAD1FFABFF93FC4EF8EEFC6AF885" bold="true" box="[956,971,1891,1909]" pageId="12" pageNumber="13">D</emphasis>
|
||
) view, respectively (as, adductor shelf; l, lesion; mf, maxillary fenestra; psf, posterior surangular fenestra).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFAAFF92FF0BFF41FEC0FEF0" box="[249,353,203,256]" pageId="13" pageNumber="14">Body mass (kg) 8,870</paragraph>
|
||
<table id="EB81C463FFAA0060FF0BFF46FD3FF879" box="[249,670,203,1929]" gridcols="13" gridrows="17" pageId="13" pageNumber="14">
|
||
<paragraph id="993E36C3FFAAFF92FE91FF46FED6FEF0" box="[355,375,203,256]" pageId="13" pageNumber="14">
|
||
<tr id="27B13481FFAA0060FF0BFF46FD3FFEF0" box="[249,670,203,256]" gridrow="0" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FE91FF46FE2FFEF0" box="[355,398,203,256]" gridcol="2" gridrow="0" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFF46FED6FEF0" box="[249,375,203,256]" colspan="2" colspanRight="1" gridcol="0" gridrow="0" pageId="13" pageNumber="14">6,740</th>
|
||
</th>
|
||
</tr>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFAAFF92FE63FF46FD3FF8CE" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FE63FF46FE73FEF0" box="[401,466,203,256]" colspan="2" colspanRight="1" gridcol="3" gridrow="0" pageId="13" pageNumber="14">
|
||
8,462 6,100
|
||
<th id="64605DFDFFAA0060FE4CFF46FE48FEF0" box="[446,489,203,256]" gridcol="5" gridrow="0" pageId="13" pageNumber="14">5,469</th>
|
||
</th>
|
||
<th id="64605DFDFFAA0060FE19FF46FDFBFEF0" box="[491,602,203,256]" colspan="4" colspanRight="3" gridcol="6" gridrow="0" pageId="13" pageNumber="14">
|
||
5,779 6,264 5,112 4,469
|
||
<th id="64605DFDFFAA0060FDB4FF46FDD0FEF0" box="[582,625,203,256]" gridcol="10" gridrow="0" pageId="13" pageNumber="14">5,025</th>
|
||
</th>
|
||
<th id="64605DFDFFAA0060FD81FF46FD3FFEF0" box="[627,670,203,256]" colspan="2" colspanRight="1" gridcol="11" gridrow="0" pageId="13" pageNumber="14">4,714 3,617</th>
|
||
<tr id="27B13481FFAA0060FF0BFE9EFD3FFEA5" box="[249,670,275,341]" gridrow="1" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFE9EFED6FEA5" box="[249,375,275,341]" colspan="2" colspanRight="1" gridcol="0" gridrow="1" pageId="13" pageNumber="14">
|
||
Pedal p. IV- 1 length 184
|
||
<td id="64605DFDFFAA0060FE91FE9EFE2FFEA5" box="[355,398,275,341]" gridcol="2" gridrow="1" pageId="13" pageNumber="14">145</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FE9EFE73FEA5" box="[401,466,275,341]" colspan="2" colspanRight="1" gridcol="3" gridrow="1" pageId="13" pageNumber="14">
|
||
154 170
|
||
<td id="64605DFDFFAA0060FE4CFE9EFE48FEA5" box="[446,489,275,341]" gridcol="5" gridrow="1" pageId="13" pageNumber="14">140</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FE9EFDFBFEA5" box="[491,602,275,341]" colspan="4" colspanRight="3" gridcol="6" gridrow="1" pageId="13" pageNumber="14">
|
||
? 176? 107
|
||
<td id="64605DFDFFAA0060FDB4FE9EFDD0FEA5" box="[582,625,275,341]" gridcol="10" gridrow="1" pageId="13" pageNumber="14">124</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FE9EFD3FFEA5" box="[627,670,275,341]" colspan="2" colspanRight="1" gridcol="11" gridrow="1" pageId="13" pageNumber="14">??</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFEE7FD3FFE25" box="[249,670,362,469]" gridrow="2" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFEE7FED6FE25" box="[249,375,362,469]" colspan="2" colspanRight="1" gridcol="0" gridrow="2" pageId="13" pageNumber="14">
|
||
Astragalus width 310
|
||
<td id="64605DFDFFAA0060FE91FEE7FE2FFE25" box="[355,398,362,469]" gridcol="2" gridrow="2" pageId="13" pageNumber="14">?</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FEE7FE73FE25" box="[401,466,362,469]" colspan="2" colspanRight="1" gridcol="3" gridrow="2" pageId="13" pageNumber="14">
|
||
? 295
|
||
<td id="64605DFDFFAA0060FE4CFEE7FE48FE25" box="[446,489,362,469]" gridcol="5" gridrow="2" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FEE7FDFBFE25" box="[491,602,362,469]" colspan="4" colspanRight="3" gridcol="6" gridrow="2" pageId="13" pageNumber="14">
|
||
280 235??
|
||
<td id="64605DFDFFAA0060FDB4FEE7FDD0FE25" box="[582,625,362,469]" gridcol="10" gridrow="2" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FEE7FD3FFE25" box="[627,670,362,469]" colspan="2" colspanRight="1" gridcol="11" gridrow="2" pageId="13" pageNumber="14">??</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFE65FD3FFDA3" box="[249,670,488,595]" gridrow="3" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFE65FED6FDA3" box="[249,375,488,595]" colspan="2" colspanRight="1" gridcol="0" gridrow="3" pageId="13" pageNumber="14">
|
||
Astragalus height 328
|
||
<td id="64605DFDFFAA0060FE91FE65FE2FFDA3" box="[355,398,488,595]" gridcol="2" gridrow="3" pageId="13" pageNumber="14">?</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FE65FE73FDA3" box="[401,466,488,595]" colspan="2" colspanRight="1" gridcol="3" gridrow="3" pageId="13" pageNumber="14">
|
||
? 280
|
||
<td id="64605DFDFFAA0060FE4CFE65FE48FDA3" box="[446,489,488,595]" gridcol="5" gridrow="3" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FE65FDFBFDA3" box="[491,602,488,595]" colspan="4" colspanRight="3" gridcol="6" gridrow="3" pageId="13" pageNumber="14">
|
||
325???
|
||
<td id="64605DFDFFAA0060FDB4FE65FDD0FDA3" box="[582,625,488,595]" gridcol="10" gridrow="3" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FE65FD3FFDA3" box="[627,670,488,595]" colspan="2" colspanRight="1" gridcol="11" gridrow="3" pageId="13" pageNumber="14">??</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFDEBFD3FFD56" box="[249,670,614,678]" gridrow="4" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFDEBFED6FD56" box="[249,375,614,678]" colspan="2" colspanRight="1" gridcol="0" gridrow="4" pageId="13" pageNumber="14">
|
||
Fibula shaft width 62
|
||
<td id="64605DFDFFAA0060FE91FDEBFE2FFD56" box="[355,398,614,678]" gridcol="2" gridrow="4" pageId="13" pageNumber="14">?</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FDEBFE73FD56" box="[401,466,614,678]" colspan="2" colspanRight="1" gridcol="3" gridrow="4" pageId="13" pageNumber="14">
|
||
59 53
|
||
<td id="64605DFDFFAA0060FE4CFDEBFE48FD56" box="[446,489,614,678]" gridcol="5" gridrow="4" pageId="13" pageNumber="14">65</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FDEBFDFBFD56" box="[491,602,614,678]" colspan="4" colspanRight="3" gridcol="6" gridrow="4" pageId="13" pageNumber="14">
|
||
55 56? 52
|
||
<td id="64605DFDFFAA0060FDB4FDEBFDD0FD56" box="[582,625,614,678]" gridcol="10" gridrow="4" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FDEBFD3FFD56" box="[627,670,614,678]" colspan="2" colspanRight="1" gridcol="11" gridrow="4" pageId="13" pageNumber="14">??</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFD37FD3FFD0A" box="[249,670,698,762]" gridrow="5" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFD37FED6FD0A" box="[249,375,698,762]" colspan="2" colspanRight="1" gridcol="0" gridrow="5" pageId="13" pageNumber="14">
|
||
Fibula length 995
|
||
<td id="64605DFDFFAA0060FE91FD37FE2FFD0A" box="[355,398,698,762]" gridcol="2" gridrow="5" pageId="13" pageNumber="14">1,025</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FD37FE73FD0A" box="[401,466,698,762]" colspan="2" colspanRight="1" gridcol="3" gridrow="5" pageId="13" pageNumber="14">
|
||
1,030 915
|
||
<td id="64605DFDFFAA0060FE4CFD37FE48FD0A" box="[446,489,698,762]" gridcol="5" gridrow="5" pageId="13" pageNumber="14">985</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FD37FDFBFD0A" box="[491,602,698,762]" colspan="4" colspanRight="3" gridcol="6" gridrow="5" pageId="13" pageNumber="14">
|
||
945 1,035??
|
||
<td id="64605DFDFFAA0060FDB4FD37FDD0FD0A" box="[582,625,698,762]" gridcol="10" gridrow="5" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FD37FD3FFD0A" box="[627,670,698,762]" colspan="2" colspanRight="1" gridcol="11" gridrow="5" pageId="13" pageNumber="14">930?</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFC83FD3FFCB6" box="[249,670,782,838]" gridrow="6" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFC83FED6FCB6" box="[249,375,782,838]" colspan="2" colspanRight="1" gridcol="0" gridrow="6" pageId="13" pageNumber="14">
|
||
Tibia shaft width 184
|
||
<td id="64605DFDFFAA0060FE91FC83FE2FFCB6" box="[355,398,782,838]" gridcol="2" gridrow="6" pageId="13" pageNumber="14">150</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FC83FE73FCB6" box="[401,466,782,838]" colspan="2" colspanRight="1" gridcol="3" gridrow="6" pageId="13" pageNumber="14">
|
||
160 150
|
||
<td id="64605DFDFFAA0060FE4CFC83FE48FCB6" box="[446,489,782,838]" gridcol="5" gridrow="6" pageId="13" pageNumber="14">155</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FC83FDFBFCB6" box="[491,602,782,838]" colspan="4" colspanRight="3" gridcol="6" gridrow="6" pageId="13" pageNumber="14">
|
||
158 170? 160
|
||
<td id="64605DFDFFAA0060FDB4FC83FDD0FCB6" box="[582,625,782,838]" gridcol="10" gridrow="6" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FC83FD3FFCB6" box="[627,670,782,838]" colspan="2" colspanRight="1" gridcol="11" gridrow="6" pageId="13" pageNumber="14">140?</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFCD7FD3FFC68" box="[249,670,858,920]" gridrow="7" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFCD7FED6FC68" box="[249,375,858,920]" colspan="2" colspanRight="1" gridcol="0" gridrow="7" pageId="13" pageNumber="14">
|
||
Tibia length 1,140
|
||
<td id="64605DFDFFAA0060FE91FCD7FE2FFC68" box="[355,398,858,920]" gridcol="2" gridrow="7" pageId="13" pageNumber="14">1,166</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FCD7FE73FC68" box="[401,466,858,920]" colspan="2" colspanRight="1" gridcol="3" gridrow="7" pageId="13" pageNumber="14">
|
||
1,140 1,060
|
||
<td id="64605DFDFFAA0060FE4CFCD7FE48FC68" box="[446,489,858,920]" gridcol="5" gridrow="7" pageId="13" pageNumber="14">1,095</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FCD7FDFBFC68" box="[491,602,858,920]" colspan="4" colspanRight="3" gridcol="6" gridrow="7" pageId="13" pageNumber="14">
|
||
1,065 1,150? 1,030
|
||
<td id="64605DFDFFAA0060FDB4FCD7FDD0FC68" box="[582,625,858,920]" gridcol="10" gridrow="7" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FCD7FD3FFC68" box="[627,670,858,920]" colspan="2" colspanRight="1" gridcol="11" gridrow="7" pageId="13" pageNumber="14">1,105 910</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFC21FD3FFBF5" box="[249,670,940,1029]" gridrow="8" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFC21FED6FBF5" box="[249,375,940,1029]" colspan="2" colspanRight="1" gridcol="0" gridrow="8" pageId="13" pageNumber="14">
|
||
Proximal femur width 426
|
||
<td id="64605DFDFFAA0060FE91FC21FE2FFBF5" box="[355,398,940,1029]" gridcol="2" gridrow="8" pageId="13" pageNumber="14">399</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FC21FE73FBF5" box="[401,466,940,1029]" colspan="2" colspanRight="1" gridcol="3" gridrow="8" pageId="13" pageNumber="14">
|
||
380 370
|
||
<td id="64605DFDFFAA0060FE4CFC21FE48FBF5" box="[446,489,940,1029]" gridcol="5" gridrow="8" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FC21FDFBFBF5" box="[491,602,940,1029]" colspan="4" colspanRight="3" gridcol="6" gridrow="8" pageId="13" pageNumber="14">
|
||
350 370? 270
|
||
<td id="64605DFDFFAA0060FDB4FC21FDD0FBF5" box="[582,625,940,1029]" gridcol="10" gridrow="8" pageId="13" pageNumber="14">330</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FC21FD3FFBF5" box="[627,670,940,1029]" colspan="2" colspanRight="1" gridcol="11" gridrow="8" pageId="13" pageNumber="14">??</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFB94FD3FFB51" box="[249,670,1049,1185]" gridrow="9" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFB94FED6FB51" box="[249,375,1049,1185]" colspan="2" colspanRight="1" gridcol="0" gridrow="9" pageId="13" pageNumber="14">
|
||
Femur circumference 590
|
||
<td id="64605DFDFFAA0060FE91FB94FE2FFB51" box="[355,398,1049,1185]" gridcol="2" gridrow="9" pageId="13" pageNumber="14">534</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FB94FE73FB51" box="[401,466,1049,1185]" colspan="2" colspanRight="1" gridcol="3" gridrow="9" pageId="13" pageNumber="14">
|
||
580 515
|
||
<td id="64605DFDFFAA0060FE4CFB94FE48FB51" box="[446,489,1049,1185]" gridcol="5" gridrow="9" pageId="13" pageNumber="14">495</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FB94FDFBFB51" box="[491,602,1049,1185]" colspan="4" colspanRight="3" gridcol="6" gridrow="9" pageId="13" pageNumber="14">
|
||
505 520 483 460
|
||
<td id="64605DFDFFAA0060FDB4FB94FDD0FB51" box="[582,625,1049,1185]" gridcol="10" gridrow="9" pageId="13" pageNumber="14">480</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FB94FD3FFB51" box="[627,670,1049,1185]" colspan="2" colspanRight="1" gridcol="11" gridrow="9" pageId="13" pageNumber="14">469 426</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFB38FD3FFB07" box="[249,670,1205,1271]" gridrow="10" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFB38FED6FB07" box="[249,375,1205,1271]" colspan="2" colspanRight="1" gridcol="0" gridrow="10" pageId="13" pageNumber="14">
|
||
Femur length 1,333
|
||
<td id="64605DFDFFAA0060FE91FB38FE2FFB07" box="[355,398,1205,1271]" gridcol="2" gridrow="10" pageId="13" pageNumber="14">1,269</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FB38FE73FB07" box="[401,466,1205,1271]" colspan="2" colspanRight="1" gridcol="3" gridrow="10" pageId="13" pageNumber="14">
|
||
1,321 1,150
|
||
<td id="64605DFDFFAA0060FE4CFB38FE48FB07" box="[446,489,1205,1271]" gridcol="5" gridrow="10" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FB38FDFBFB07" box="[491,602,1205,1271]" colspan="4" colspanRight="3" gridcol="6" gridrow="10" pageId="13" pageNumber="14">
|
||
1,350 1,280 1,232 1,210
|
||
<td id="64605DFDFFAA0060FDB4FB38FDD0FB07" box="[582,625,1205,1271]" gridcol="10" gridrow="10" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FB38FD3FFB07" box="[627,670,1205,1271]" colspan="2" colspanRight="1" gridcol="11" gridrow="10" pageId="13" pageNumber="14">1,100 1,040</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFA86FD3FFAB9" box="[249,670,1291,1353]" gridrow="11" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFA86FED6FAB9" box="[249,375,1291,1353]" colspan="2" colspanRight="1" gridcol="0" gridrow="11" pageId="13" pageNumber="14">
|
||
Ilium length 1,545
|
||
<td id="64605DFDFFAA0060FE91FA86FE2FFAB9" box="[355,398,1291,1353]" gridcol="2" gridrow="11" pageId="13" pageNumber="14">1,540</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FA86FE73FAB9" box="[401,466,1291,1353]" colspan="2" colspanRight="1" gridcol="3" gridrow="11" pageId="13" pageNumber="14">
|
||
1,525?
|
||
<td id="64605DFDFFAA0060FE4CFA86FE48FAB9" box="[446,489,1291,1353]" gridcol="5" gridrow="11" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FA86FDFBFAB9" box="[491,602,1291,1353]" colspan="4" colspanRight="3" gridcol="6" gridrow="11" pageId="13" pageNumber="14">
|
||
1,540 1,470 1,397?
|
||
<td id="64605DFDFFAA0060FDB4FA86FDD0FAB9" box="[582,625,1291,1353]" gridcol="10" gridrow="11" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FA86FD3FFAB9" box="[627,670,1291,1353]" colspan="2" colspanRight="1" gridcol="11" gridrow="11" pageId="13" pageNumber="14">1,180?</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFAD0FD3FFA59" box="[249,670,1373,1449]" gridrow="12" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFAD0FED6FA59" box="[249,375,1373,1449]" colspan="2" colspanRight="1" gridcol="0" gridrow="12" pageId="13" pageNumber="14">
|
||
Manual p. I-1 length 98.5
|
||
<td id="64605DFDFFAA0060FE91FAD0FE2FFA59" box="[355,398,1373,1449]" gridcol="2" gridrow="12" pageId="13" pageNumber="14">?</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FAD0FE73FA59" box="[401,466,1373,1449]" colspan="2" colspanRight="1" gridcol="3" gridrow="12" pageId="13" pageNumber="14">
|
||
78?
|
||
<td id="64605DFDFFAA0060FE4CFAD0FE48FA59" box="[446,489,1373,1449]" gridcol="5" gridrow="12" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FAD0FDFBFA59" box="[491,602,1373,1449]" colspan="4" colspanRight="3" gridcol="6" gridrow="12" pageId="13" pageNumber="14">
|
||
? 89??
|
||
<td id="64605DFDFFAA0060FDB4FAD0FDD0FA59" box="[582,625,1373,1449]" gridcol="10" gridrow="12" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FAD0FD3FFA59" box="[627,670,1373,1449]" colspan="2" colspanRight="1" gridcol="11" gridrow="12" pageId="13" pageNumber="14">90?</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BFA30FD3FF9FB" box="[249,670,1469,1547]" gridrow="13" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BFA30FED6F9FB" box="[249,375,1469,1547]" colspan="2" colspanRight="1" gridcol="0" gridrow="13" pageId="13" pageNumber="14">
|
||
Scapula blade width 74. 8
|
||
<td id="64605DFDFFAA0060FE91FA30FE2FF9FB" box="[355,398,1469,1547]" gridcol="2" gridrow="13" pageId="13" pageNumber="14">73</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63FA30FE73F9FB" box="[401,466,1469,1547]" colspan="2" colspanRight="1" gridcol="3" gridrow="13" pageId="13" pageNumber="14">
|
||
68 65. 7
|
||
<td id="64605DFDFFAA0060FE4CFA30FE48F9FB" box="[446,489,1469,1547]" gridcol="5" gridrow="13" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19FA30FDFBF9FB" box="[491,602,1469,1547]" colspan="4" colspanRight="3" gridcol="6" gridrow="13" pageId="13" pageNumber="14">
|
||
? 65??
|
||
<td id="64605DFDFFAA0060FDB4FA30FDD0F9FB" box="[582,625,1469,1547]" gridcol="10" gridrow="13" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81FA30FD3FF9FB" box="[627,670,1469,1547]" colspan="2" colspanRight="1" gridcol="11" gridrow="13" pageId="13" pageNumber="14">??</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BF992FD3FF98C" box="[249,670,1567,1660]" gridrow="14" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BF992FED6F98C" box="[249,375,1567,1660]" colspan="2" colspanRight="1" gridcol="0" gridrow="14" pageId="13" pageNumber="14">
|
||
Dentary tooth row length 595
|
||
<td id="64605DFDFFAA0060FE91F992FE2FF98C" box="[355,398,1567,1660]" gridcol="2" gridrow="14" pageId="13" pageNumber="14">508</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63F992FE73F98C" box="[401,466,1567,1660]" colspan="2" colspanRight="1" gridcol="3" gridrow="14" pageId="13" pageNumber="14">
|
||
585 492
|
||
<td id="64605DFDFFAA0060FE4CF992FE48F98C" box="[446,489,1567,1660]" gridcol="5" gridrow="14" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19F992FDFBF98C" box="[491,602,1567,1660]" colspan="4" colspanRight="3" gridcol="6" gridrow="14" pageId="13" pageNumber="14">
|
||
575 635 546 530
|
||
<td id="64605DFDFFAA0060FDB4F992FDD0F98C" box="[582,625,1567,1660]" gridcol="10" gridrow="14" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81F992FD3FF98C" box="[627,670,1567,1660]" colspan="2" colspanRight="1" gridcol="11" gridrow="14" pageId="13" pageNumber="14">??</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BF91DFD3FF927" box="[249,670,1680,1751]" gridrow="15" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BF91DFED6F927" box="[249,375,1680,1751]" colspan="2" colspanRight="1" gridcol="0" gridrow="15" pageId="13" pageNumber="14">
|
||
Morph Robust
|
||
<td id="64605DFDFFAA0060FE91F91DFE2FF927" box="[355,398,1680,1751]" gridcol="2" gridrow="15" pageId="13" pageNumber="14">Robust</td>
|
||
</th>
|
||
<td id="64605DFDFFAA0060FE63F91DFE73F927" box="[401,466,1680,1751]" colspan="2" colspanRight="1" gridcol="3" gridrow="15" pageId="13" pageNumber="14">
|
||
Robust Robust
|
||
<td id="64605DFDFFAA0060FE4CF91DFE48F927" box="[446,489,1680,1751]" gridcol="5" gridrow="15" pageId="13" pageNumber="14">Robust</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FE19F91DFDFBF927" box="[491,602,1680,1751]" colspan="4" colspanRight="3" gridcol="6" gridrow="15" pageId="13" pageNumber="14">
|
||
Gracile Gracile Gracile Gracile
|
||
<td id="64605DFDFFAA0060FDB4F91DFDD0F927" box="[582,625,1680,1751]" gridcol="10" gridrow="15" pageId="13" pageNumber="14">?</td>
|
||
</td>
|
||
<td id="64605DFDFFAA0060FD81F91DFD3FF927" box="[627,670,1680,1751]" colspan="2" colspanRight="1" gridcol="11" gridrow="15" pageId="13" pageNumber="14">??</td>
|
||
</tr>
|
||
<tr id="27B13481FFAA0060FF0BF97CFD3FF879" box="[249,670,1777,1929]" gridrow="16" pageId="13" pageNumber="14">
|
||
<th id="64605DFDFFAA0060FF0BF97CFD3FF879" box="[249,670,1777,1929]" colspan="13" colspanRight="12" gridcol="0" gridrow="16" pageId="13" pageNumber="14">
|
||
Specimens
|
||
<materialsCitation id="29E93C9EFFAAFF92FEBFF8D5FEC0F8B5" ID-GBIF-Occurrence="2421817401" box="[333,353,1795,1929]" collectingDate="1991" collectionCode="RSM" country="Canada" location="near Eastend" pageId="13" pageNumber="14" preparations="relatively complete (roughly 65%)" specimenCode="RSM P2523.8" stateProvince="Saskatchewan">RSM P 2523.8</materialsCitation>
|
||
<materialsCitation id="29E93C9EFFAAFF92FE91F8E8FE2FF8D0" ID-GBIF-Occurrence="2421817405" box="[355,398,1788,1928]" collectionCode="AMNH" pageId="13" pageNumber="14" specimenCode="CM 9380 (AMNH 973)">CM 9380 (AMNH 973</materialsCitation>
|
||
)
|
||
<materialsCitation id="29E93C9EFFAAFF92FE63F8CEFE04F8EF" ID-GBIF-Occurrence="2421817398" box="[401,421,1777,1928]" collectionCode="FMNH" pageId="13" pageNumber="14" specimenCode="FMNH PR2081">FMNH PR 2081</materialsCitation>
|
||
<materialsCitation id="29E93C9EFFAAFF92FE55F8D8FE1AF8BE" ID-GBIF-Occurrence="2421817397" box="[423,443,1823,1929]" collectionCode="MOR" pageId="13" pageNumber="14" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
<materialsCitation id="29E93C9EFFAAFF92FE4CF8C4FE48F8C8" ID-GBIF-Occurrence="2421817406" box="[446,489,1782,1929]" collectionCode="RTMP" pageId="13" pageNumber="14" specimenCode="RTMP 81.12.1 (NMC 9950)">RTMP 81. 12. 1, NMC 9950</materialsCitation>
|
||
<materialsCitation id="29E93C9EFFAAFF92FE19F8D2FE5EF8A9" ID-GBIF-Occurrence="2421817395" box="[491,511,1834,1929]" collectionCode="BHI" pageId="13" pageNumber="14" specimenCode="BHI 3033">BHI 3033</materialsCitation>
|
||
<materialsCitation id="29E93C9EFFAAFF92FDF0F8D8FDB7F8BE" ID-GBIF-Occurrence="2421817408" box="[514,534,1835,1929]" collectionCode="MOR" pageId="13" pageNumber="14" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
<materialsCitation id="29E93C9EFFAAFF92FDEBF8D8FD8CF8BE" ID-GBIF-Occurrence="2421817403" box="[537,557,1835,1929]" collectionCode="MOR" pageId="13" pageNumber="14" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
<materialsCitation id="29E93C9EFFAAFF92FDDDF8C4FDE2F8E1" ID-GBIF-Occurrence="2421817410" box="[559,579,1800,1929]" collectionCode="TMP" pageId="13" pageNumber="14" specimenCode="TMP 81.6.1">RTMP 81. 6. 1</materialsCitation>
|
||
<materialsCitation id="29E93C9EFFAAFF92FDB4F8E8FDD0F8EF" ID-GBIF-Occurrence="2421817404" box="[582,625,1776,1928]" collectionCode="BM" httpUri="https://data.nhm.ac.uk/dataset/collection-specimens/resource/05ff2255-c38a-40c9-b657-4ccb55ab2feb/record/5479135" pageId="13" pageNumber="14" specimenCode="BM R8040 (AMNH 5881)">BM R8040 (AMNH 5881</materialsCitation>
|
||
<materialsCitation id="29E93C9EFFAAFF92FD81F8D8FD26F8BE" ID-GBIF-Occurrence="2421817409" box="[627,647,1835,1929]" collectionCode="MOR" pageId="13" pageNumber="14" specimenCode="MOR 009">MOR 009</materialsCitation>
|
||
<materialsCitation id="29E93C9EFFAAFF92FD78F8C9FD3FF8CE" ID-GBIF-Occurrence="3396394301" box="[650,670,1807,1928]" collectionCode="USNM" httpUri="http://n2t.net/ark:/65665/37f7c8976-8ae8-49f4-931e-09f675c134fb" pageId="13" pageNumber="14" specimenCode="USNM 6183">USNM 6183</materialsCitation>
|
||
</th>
|
||
</tr>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFAAFF92FCB6FE30FC44FDA8" blockId="13.[810,1450,205,920]" pageId="13" pageNumber="14">
|
||
Medullary bone structure would only be expected in a narrow window associated with eggshell formation, and there are no histological grounds for assigning
|
||
<materialsCitation id="29E93C9EFFAAFF92FA81FE7EFC23FDD3" ID-GBIF-Occurrence="2597591350" collectionCode="RSM" pageId="13" pageNumber="14" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
to either sex. As such, no independent support for the sexual dimorphism hypothesis is possible from the present analysis.
|
||
</paragraph>
|
||
<caption id="CDFE664BFFAAFF92FF3FFE7EFF43F91D" ID-DOI="http://doi.org/10.5281/zenodo.3560559" ID-Table-UUID="CDFE664BFFAAFF92FF3FFE7EFF43F91D" ID-Zenodo-Dep="3560559" box="[205,226,358,1773]" httpUri="http://table.plazi.org/id/CDFE664BFFAAFF92FF3FFE7EFF43F91D" pageId="13" pageNumber="14" targetBox="[249,670,203,1929]" targetIsTable="true" targetPageId="13">
|
||
<paragraph id="993E36C3FFAAFF92FF3FFE7EFF43F91D" blockId="13.[205,226,358,1773]" box="[205,226,358,1773]" pageId="13" pageNumber="14">
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFE7EFF43FDFB" bold="true" box="[205,226,499,523]" pageId="13" pageNumber="14">in</emphasis>
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFEEBFF43FE1C" bold="true" box="[205,226,358,492]" pageId="13" pageNumber="14">millimeters</emphasis>
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFD9CFF43FD4B" bold="true" box="[205,226,529,699]" pageId="13" pageNumber="14">measurements</emphasis>
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFD4FFF43FD2F" bold="true" box="[205,226,706,735]" pageId="13" pageNumber="14">all</emphasis>
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFD6BFF43FD1B" bold="true" box="[205,226,742,747]" pageId="13" pageNumber="14">,</emphasis>
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFD66FF43FC95" bold="true" box="[205,226,747,869]" pageId="13" pageNumber="14">specimens</emphasis>
|
||
<taxonomicName id="5E814D40FFAAFF92FF3FFCE1FF40FC60" authority="Osborn, 1905" box="[205,225,876,912]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFCE1FF40FC60" bold="true" box="[205,225,876,912]" italics="true" pageId="13" pageNumber="14">rex</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFC1AFF40FBB8" bold="true" box="[205,225,919,1096]" italics="true" pageId="13" pageNumber="14">Tyrannosaurus</emphasis>
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFBC3FF43FB7D" bold="true" box="[205,226,1102,1165]" pageId="13" pageNumber="14">other</emphasis>
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFB19FF43FB30" bold="true" box="[205,226,1172,1216]" pageId="13" pageNumber="14">and</emphasis>
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFB4AFF43FAFF" bold="true" box="[205,226,1223,1295]" pageId="13" pageNumber="14">2523.8</emphasis>
|
||
P RSM
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFAE9FF43FA8A" bold="true" box="[205,226,1380,1402]" pageId="13" pageNumber="14">of</emphasis>
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FFA0CFF43F9DB" bold="true" box="[205,226,1409,1579]" pageId="13" pageNumber="14">measurements</emphasis>
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FF9BFFF43F989" bold="true" box="[205,226,1586,1657]" pageId="13" pageNumber="14">Select</emphasis>
|
||
. 1
|
||
<emphasis id="ABF5EAD1FFAAFF92FF3FF914FF43F91D" bold="true" box="[205,226,1689,1773]" pageId="13" pageNumber="14">TABLE</emphasis>
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFAAFF92FCB6FDD0FAEDFC68" blockId="13.[810,1450,205,920]" pageId="13" pageNumber="14">
|
||
The extent of remodeling in
|
||
<materialsCitation id="29E93C9EFFAAFF92FB6AFDD0FA95FD83" ID-GBIF-Occurrence="2597591351" box="[1176,1332,605,627]" collectionCode="RSM" pageId="13" pageNumber="14" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
precludes direct aging using a total growth line count, as has been done for other tyrannosaurid specimens (
|
||
<bibRefCitation id="FD104B32FFAAFF92FB0FFD1FFCC0FD32" author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA & Yerby SA & Brochu CA" journalOrPublisher="Nature" pageId="13" pageNumber="14" pagination="772 - 775" part="430" refId="ref8744" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA, Yerby SA, Brochu CA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 430: 772 - 775." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">Erickson et al., 2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFAAFF92FC9DFD20FBB9FD33" author="Erickson GM" box="[879,1048,685,707]" journalOrPublisher="Trends Ecol Evol" pageId="13" pageNumber="14" pagination="677 - 684" part="20" refId="ref8644" refString="Erickson GM. 2005. Assessing dinosaur growth patterns: a microscopic revolution. Trends Ecol Evol 20: 677 - 684." title="Assessing dinosaur growth patterns: a microscopic revolution" type="journal article" year="2005">Erickson, 2005</bibRefCitation>
|
||
). Nevertheless, the degree of remodeling exceeds that reported in any
|
||
<taxonomicName id="5E814D40FFAAFF92FB16FD45FA8BFD2D" authorityName="Osborn" authorityYear="1905" box="[1252,1322,712,733]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFAAFF92FB16FD45FA8BFD2D" box="[1252,1322,712,733]" italics="true" pageId="13" pageNumber="14">T. rex</emphasis>
|
||
</taxonomicName>
|
||
long bone (
|
||
<bibRefCitation id="FD104B32FFAAFF92FCC3FD6FFB87FD07" author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA & Yerby SA & Brochu CA" box="[817,1062,738,760]" journalOrPublisher="Nature" pageId="13" pageNumber="14" pagination="772 - 775" part="430" refId="ref8744" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA, Yerby SA, Brochu CA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 430: 772 - 775." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">Erickson et al., 2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFAAFF92FBC5FD6EFAC3FD08" author="Horner JR & Padian K." box="[1079,1378,738,760]" journalOrPublisher="Proc R Soc London B Biol Sci" pageId="13" pageNumber="14" pagination="1875 - 1880" part="271" refId="ref9080" refString="Horner JR, Padian K. 2004. Age and growth dynamics of Tyrannosaurus rex. Proc R Soc London B Biol Sci 271: 1875 - 1880." title="Age and growth dynamics of Tyrannosaurus rex" type="journal article" year="2004">Horner and Padian, 2004</bibRefCitation>
|
||
). This suggests the animal is of at least comparable, and likely more advanced ontogenetic status than the ~28 year old, somatically mature
|
||
<taxonomicName id="5E814D40FFAAFF92FBEAFCBEFBC1FCB8" authorityName="Osborn" authorityYear="1905" box="[1048,1120,819,840]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFAAFF92FBEAFCBEFBC1FCB8" box="[1048,1120,819,840]" italics="true" pageId="13" pageNumber="14">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimen
|
||
<materialsCitation id="29E93C9EFFAAFF92FB14FCBEFA0BFCB7" ID-GBIF-Occurrence="2597591364" box="[1254,1450,818,840]" collectionCode="FMNH" pageId="13" pageNumber="14" specimenCode="FMNH PR2081">FMNH PR 2081</materialsCitation>
|
||
(
|
||
<bibRefCitation id="FD104B32FFAAFF92FCC3FCC0FB8CFC93" author="Erickson GM & Makovicky PJ & Currie PJ & Norell MA & Yerby SA & Brochu CA" box="[817,1069,845,867]" journalOrPublisher="Nature" pageId="13" pageNumber="14" pagination="772 - 775" part="430" refId="ref8744" refString="Erickson GM, Makovicky PJ, Currie PJ, Norell MA, Yerby SA, Brochu CA. 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature 430: 772 - 775." title="Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs" type="journal article" year="2004">Erickson et al., 2004</bibRefCitation>
|
||
). Thus, the histological analysis may offer independent support for the prediction that high robustness correlates with greater maturity.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFAAFF92FCD8FC4EFAEEFC2B" blockId="13.[810,1451,963,1934]" box="[810,1359,963,987]" pageId="13" pageNumber="14">
|
||
<heading id="C27681AFFFAAFF92FCD8FC4EFAEEFC2B" bold="true" box="[810,1359,963,987]" fontSize="10" level="2" pageId="13" pageNumber="14" reason="0">
|
||
<emphasis id="ABF5EAD1FFAAFF92FCD8FC4EFAEEFC2B" bold="true" box="[810,1359,963,987]" pageId="13" pageNumber="14">Comparisons with Other Giant Theropods</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFAAFF92FCB6FC63FBAEFAFE" blockId="13.[810,1451,963,1934]" pageId="13" pageNumber="14">
|
||
Is
|
||
<taxonomicName id="5E814D40FFAAFF92FC93FC63FC01FBF3" authorityName="Osborn" authorityYear="1905" box="[865,928,1006,1027]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFAAFF92FC93FC63FC01FBF3" box="[865,928,1006,1027]" italics="true" pageId="13" pageNumber="14">T. rex</emphasis>
|
||
</taxonomicName>
|
||
the largest known terrestrial carnivore ever to have lived? Based on the size distribution for described specimens of giant theropods and our new morphometric data for
|
||
<taxonomicName id="5E814D40FFAAFF92FC62FBB3FC72FBA3" authorityName="Osborn" authorityYear="1905" box="[912,979,1086,1107]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFAAFF92FC62FBB3FC72FBA3" box="[912,979,1086,1107]" italics="true" pageId="13" pageNumber="14">T. rex</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<materialsCitation id="29E93C9EFFAAFF92FC17FBB0FB21FBA3" ID-GBIF-Occurrence="2597591334" box="[997,1152,1085,1107]" collectionCode="RSM" pageId="13" pageNumber="14" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
substantially exceeds the known size for all other giant theropods (1,650–6,260 kg) (
|
||
<tableCitation id="D4030378FFAAFF92FCC3FBFEFC25FB79" box="[817,900,1139,1161]" captionStart="TABLE 2" captionStartId="14.[168,252,205,226]" captionTargetPageId="14" captionText="TABLE 2. Comparison of femoral dimensions and estimated body mass of the largest theropods ±25% (mass estimation based on the formula of Campione et al. (2014))" httpUri="http://table.plazi.org/id/CDFE664BFFA9FF91FF5AFF40FBCCFF0A" pageId="13" pageNumber="14" tableUuid="CDFE664BFFA9FF91FF5AFF40FBCCFF0A">Table 2</tableCitation>
|
||
), including taxa previously touted as larger (
|
||
<bibRefCitation id="FD104B32FFAAFF92FA9FFBFEFC57FB53" author="Coria RA & Salgado L." journalOrPublisher="Nature" pageId="13" pageNumber="14" pagination="224 - 226" part="377" refId="ref8405" refString="Coria RA, Salgado L. 1995. A new giant carnivorous dinosaur from the Cretaceous of Patagonia. Nature 377: 224 - 226." title="A new giant carnivorous dinosaur from the Cretaceous of Patagonia" type="journal article" year="1995">Coria and Salgado, 1995</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFAAFF92FBF3FB00FB2FFB53" author="Currie PJ" box="[1025,1166,1165,1187]" journalOrPublisher="Science" pageId="13" pageNumber="14" pagination="971 - 972" part="272" refId="ref8455" refString="Currie PJ. 1996. Out of Africa: meat-eating dinosaurs that challenge Tyrannosaurus rex. Science 272: 971 - 972." title="Out of Africa: meat-eating dinosaurs that challenge Tyrannosaurus rex" type="journal article" year="1996">Currie, 1996</bibRefCitation>
|
||
). There are, however, two important caveats that should be kept in mind when interpreting these theropod mass estimations and the conclusion based on
|
||
<materialsCitation id="29E93C9EFFAAFF92FC22FB50FBCBFB03" ID-GBIF-Occurrence="2597591346" box="[976,1130,1245,1267]" collectionCode="RSM" pageId="13" pageNumber="14" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
that
|
||
<taxonomicName id="5E814D40FFAAFF92FB5FFB53FB51FB03" authorityName="Osborn" authorityYear="1905" box="[1197,1264,1246,1267]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFAAFF92FB5FFB53FB51FB03" box="[1197,1264,1246,1267]" italics="true" pageId="13" pageNumber="14">T. rex</emphasis>
|
||
</taxonomicName>
|
||
was the largest terrestrial carnivore.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFAAFF92FD4BFB4FFD6CFB35" blockId="13.[697,717,1218,1928]" box="[697,717,1218,1221]" pageId="13" pageNumber="14">.</paragraph>
|
||
<paragraph id="993E36C3FFAAFF92FD4BFB48FD6CF878" blockId="13.[697,717,1218,1928]" box="[697,717,1221,1928]" pageId="13" pageNumber="14">
|
||
<tableNote id="6467374DFFAAFF92FD4BFB48FD6CF878" box="[697,717,1221,1928]" pageId="13" pageNumber="14">) b 2008 (Larson on based morphos gracile / robust putative to Assignment</tableNote>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFAAFF92FCB6FA9EFC75F923" blockId="13.[810,1451,963,1934]" pageId="13" pageNumber="14">
|
||
First, as the largest
|
||
<taxonomicName id="5E814D40FFAAFF92FBC0FA9EFBD5FAD8" authorityName="Osborn" authorityYear="1905" box="[1074,1140,1299,1320]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFAAFF92FBC0FA9EFBD5FAD8" box="[1074,1140,1299,1320]" italics="true" pageId="13" pageNumber="14">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimen, and presumably among the oldest,
|
||
<materialsCitation id="29E93C9EFFAAFF92FC0AFAA0FB2EFAB3" ID-GBIF-Occurrence="2597591308" box="[1016,1167,1325,1347]" collectionCode="RSM" pageId="13" pageNumber="14" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
appears to represent the chance sampling of a rare somatically mature individual. Within vertebrate paleontology, a combination of taphonomic and collection biases tend to favor the recovery of larger specimens (
|
||
<bibRefCitation id="FD104B32FFAAFF92FBF6FA15FB0DFA5D" author="Damuth J." box="[1028,1196,1432,1454]" journalOrPublisher="Paleobiology" pageId="13" pageNumber="14" pagination="434 - 446" part="8" refId="ref8583" refString="Damuth J. 1982. Analysis of the preservation of community structure in assemblages of fossil mammals. Paleobiology 8: 434 - 446." title="Analysis of the preservation of community structure in assemblages of fossil mammals" type="journal article" year="1982">Damuth, 1982</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFAAFF92FB32FA15FA06FA5D" author="Brown CM & Campione NE & Giacomini HC & O' Brien LJ & Vavrek MJ & Evans DC" box="[1216,1447,1432,1454]" journalOrPublisher="Biol Lett" pageId="13" pageNumber="14" pagination="20120582" part="9" refId="ref8153" refString="Brown CM, Campione NE, Giacomini HC, O' Brien LJ, Vavrek MJ, Evans DC. 2012. Ecological modelling, size distributions and taphonomic size bias in dinosaur faunas: a comment on Codron et al. (2012). Biol Lett 9: 20120582." title="Ecological modelling, size distributions and taphonomic size bias in dinosaur faunas: a comment on Codron et al. (2012)" type="journal article" year="2012">Brown et al., 2012</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFAAFF92FCD8FA3EFBA4FA39" author="Brown CM & Evans DC & Campione NE & O' Brien LJ & Eberth DA" box="[810,1029,1459,1481]" journalOrPublisher="Palaeogeogr Palaeoclimatol Palaeoecol" pageId="13" pageNumber="14" pagination="108 - 122" part="372" refId="ref8204" refString="Brown CM, Evans DC, Campione NE, O' Brien LJ, Eberth DA. 2013. Evidence for taphonomic size bias in the Dinosaur Park Formation (Campanian, Alberta), a model Mesozoic terrestrial alluvial-paralic system. Palaeogeogr Palaeoclimatol Palaeoecol 372: 108 - 122." title="Evidence for taphonomic size bias in the Dinosaur Park Formation (Campanian, Alberta), a model Mesozoic terrestrial alluvial-paralic system" type="journal article" year="2013">Brown et al., 2013</bibRefCitation>
|
||
). However, a bias also exists in the other direction, which makes the recovery of the largest and oldest representatives of many dinosaur taxa improbable.
|
||
<bibRefCitation id="FD104B32FFAAFF92FC4EF98EFAA1F9E9" author="Erickson GM & Currie P." box="[956,1280,1539,1561]" journalOrPublisher="Science" pageId="13" pageNumber="14" pagination="213 - 217" part="313" refId="ref8687" refString="Erickson GM, Currie P. 2006. Tyrannosaur life tables: an example of nonavian dinosaur population biology. Science 313: 213 - 217." title="Tyrannosaur life tables: an example of nonavian dinosaur population biology" type="journal article" year="2006">Erickson and Currie (2006)</bibRefCitation>
|
||
, in a previous study surveying the size of all substantially complete
|
||
<taxonomicName id="5E814D40FFAAFF92FCD8F9B4FCCDF9BE" authorityName="Osborn" authorityYear="1905" box="[810,876,1593,1614]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFAAFF92FCD8F9B4FCCDF9BE" box="[810,876,1593,1614]" italics="true" pageId="13" pageNumber="14">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens, showed that the taxon had a type-B1 survivorship curve (high juvenile mortality, negligible midlife mortality, and high adulthood mortality) with very few individuals reaching full adult size. As such, the recovery of older somatically mature specimens should be rare.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFAAFF91FCB6F955FE28FBA2" blockId="13.[810,1451,963,1934]" lastBlockId="14.[137,778,791,1933]" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14">
|
||
Because a type-B1 survivorship curve and resulting sampling bias occurs throughout the
|
||
<taxonomicName id="5E814D40FFAAFF92FB76F97EFB5BF8F9" box="[1156,1274,1779,1801]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="order">Dinosauria</taxonomicName>
|
||
(
|
||
<bibRefCitation id="FD104B32FFAAFF92FAF5F97EFA07F8F9" author="Erickson GM" box="[1287,1446,1779,1801]" journalOrPublisher="Trends Ecol Evol" pageId="13" pageNumber="14" pagination="677 - 684" part="20" refId="ref8644" refString="Erickson GM. 2005. Assessing dinosaur growth patterns: a microscopic revolution. Trends Ecol Evol 20: 677 - 684." title="Assessing dinosaur growth patterns: a microscopic revolution" type="journal article" year="2005">Erickson, 2005</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFAAFF92FCD8F883FB80F8D3" author="Hone DW & Mallon JC" box="[810,1057,1805,1827]" journalOrPublisher="Palaeontology" pageId="13" pageNumber="14" pagination="535 - 545" part="60" refId="ref9053" refString="Hone DW, Mallon JC. 2017. Protracted growth impedes the detection of sexual dimorphism in non-avian dinosaurs. Palaeontology 60: 535 - 545." title="Protracted growth impedes the detection of sexual dimorphism in non-avian dinosaurs" type="journal article" year="2017">Hone and Mallon, 2017</bibRefCitation>
|
||
), and because most dinosaur taxa are known from single specimens (
|
||
<bibRefCitation id="FD104B32FFAAFF92FB7AF8A5FA3FF8CE" author="Wang SC & Dodson P." box="[1160,1438,1832,1854]" journalOrPublisher="Proceedings of the National Academy of Sciences" pageId="13" pageNumber="14" pagination="13601 - 13605" part="103" refId="ref9806" refString="Wang SC, Dodson P. 2006. Estimating the diversity of dinosaurs. Proceedings of the National Academy of Sciences 103: 13601 - 13605." title="Estimating the diversity of dinosaurs" type="journal article" year="2006">Wang and Dodson, 2006</bibRefCitation>
|
||
),
|
||
<bibRefCitation id="FD104B32FFAAFF92FCD8F8CEFC71F8A9" author="Erickson GM" box="[810,976,1859,1881]" journalOrPublisher="Trends Ecol Evol" pageId="13" pageNumber="14" pagination="677 - 684" part="20" refId="ref8644" refString="Erickson GM. 2005. Assessing dinosaur growth patterns: a microscopic revolution. Trends Ecol Evol 20: 677 - 684." title="Assessing dinosaur growth patterns: a microscopic revolution" type="journal article" year="2005">Erickson (2005)</bibRefCitation>
|
||
and
|
||
<bibRefCitation id="FD104B32FFAAFF92FBF6F8CEFB75F8A9" author="Horner JR & Goodwin MB & Myhrvold N." box="[1028,1236,1859,1881]" journalOrPublisher="PLoS One" pageId="13" pageNumber="14" pagination="e 16574" part="6" refId="ref9109" refString="Horner JR, Goodwin MB, Myhrvold N. 2011. Dinosaur census reveals abundant Tyrannosaurus and rare ontogenetic stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA. PLoS One 6: e 16574." title="Dinosaur census reveals abundant Tyrannosaurus and rare ontogenetic stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA" type="journal article" year="2011">Horner et al. (2011)</bibRefCitation>
|
||
predicted that most known species would be found to be substantially larger than currently thought. However,
|
||
<taxonomicName id="5E814D40FFAAFF92FB93F8F4FB3FF87E" authorityName="Osborn" authorityYear="1905" box="[1121,1182,1913,1934]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFAAFF92FB93F8F4FB3FF87E" box="[1121,1182,1913,1934]" italics="true" pageId="13" pageNumber="14">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is now known from over 20+ reasonably complete individuals, increasing the likelihood that one of the oldest individuals is represented.
|
||
<materialsCitation id="29E93C9EFFA9FF91FD26FCBCFF7EFC92" ID-GBIF-Occurrence="2597591310" collectionCode="RSM" pageId="14" pageNumber="15" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
and
|
||
<materialsCitation id="29E93C9EFFA9FF91FEE9FCC0FE6EFC91" ID-GBIF-Occurrence="2597591359" box="[283,463,844,866]" collectionCode="FMNH" pageId="14" pageNumber="15" specimenCode="FMNH PR2081">FMNH PR 2081</materialsCitation>
|
||
are close in estimated mass and in most skeletal dimensions, and, as both specimens are skeletally mature, they may converge upon the asymptotic mass for
|
||
<emphasis id="ABF5EAD1FFA9FF91FF03FC10FE99FC42" box="[241,312,925,946]" italics="true" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF03FC10FE92FC42" authorityName="Osborn" authorityYear="1905" box="[241,307,925,946]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
The slightly greater size of
|
||
<materialsCitation id="29E93C9EFFA9FF91FD86FC11FCABFC42" ID-GBIF-Occurrence="2597591353" box="[628,778,924,946]" collectionCode="RSM" pageId="14" pageNumber="15" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
may reflect it being relatively large for its age class. In extant vertebrates, populations within an age class typically show 20% variance in body mass, and variation is even greater in taxa showing size-based sexual dimorphism (
|
||
<bibRefCitation id="FD104B32FFA9FF91FD88FB8AFCA4FBEC" author="Hanks J." box="[634,773,1031,1053]" journalOrPublisher="Afr J Ecol" pageId="14" pageNumber="15" pagination="251 - 272" part="10" refId="ref8816" refString="Hanks J. 1972. Growth of the African elephant (Loxodonta africana). Afr J Ecol 10: 251 - 272." title="Growth of the African elephant (Loxodonta africana)" type="journal article" year="1972">Hanks, 1972</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA9FF91FF7BFBAFFECEFBC7" author="Navarro J & Vignolo P & Demaria MR & Maceira NO & Martella MB" box="[137,367,1057,1079]" journalOrPublisher="Arch Geflugelkunde" pageId="14" pageNumber="15" pagination="90 - 93" part="69" refId="ref9544" refString="Navarro J, Vignolo P, Demaria MR, Maceira NO, Martella MB. 2005. Growth curves of farmed Greater Rheas (Rhea americana albescens) from central Argentina. Arch Geflugelkunde 69: 90 - 93." title="Growth curves of farmed Greater Rheas (Rhea americana albescens) from central Argentina" type="journal article" year="2005">Navarro et al., 2005</bibRefCitation>
|
||
), which as previously explained, has been proposed for
|
||
<emphasis id="ABF5EAD1FFA9FF91FEB5FBB0FE28FBA2" box="[327,393,1085,1106]" italics="true" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FEB5FBB0FE24FBA2" authorityName="Osborn" authorityYear="1905" box="[327,389,1085,1106]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFAAFF92FDAFF967FDD0F901" box="[605,625,1770,1777]" pageId="13" pageNumber="14">)</paragraph>
|
||
</table>
|
||
<caption id="CDFE664BFFA9FF91FF5AFF40FBCCFF0A" ID-Table-UUID="CDFE664BFFA9FF91FF5AFF40FBCCFF0A" httpUri="http://table.plazi.org/id/CDFE664BFFA9FF91FF5AFF40FBCCFF0A" pageId="14" pageNumber="15" targetBox="[137,1450,274,647]" targetIsTable="true" targetPageId="14">
|
||
<paragraph id="993E36C3FFA9FF91FF5AFF40FBCCFF0A" blockId="14.[168,1419,205,250]" pageId="14" pageNumber="15">
|
||
TABLE 2. Comparison of femoral dimensions and estimated body mass of the largest theropods
|
||
<emphasis id="ABF5EAD1FFA9FF91FAF6FF43FAB4FF12" bold="true" box="[1284,1301,206,226]" italics="true" pageId="14" pageNumber="15">±</emphasis>
|
||
25% (mass estimation based on the formula of
|
||
<bibRefCitation id="FD104B32FFA9FF91FC91FF68FBCCFF0A" author="Campione NE & Evans DC & Brown CM & Carrano MT" box="[867,1133,229,250]" journalOrPublisher="Methods Ecol Evol" pageId="14" pageNumber="15" pagination="913 - 923" part="5" refId="ref8337" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol Evol 5: 913 - 923." title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">Campione et al. (2014))</bibRefCitation>
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="993E36C3FFA9FF91FF7BFEA7FA35FD77" pageId="14" pageNumber="15">
|
||
<table id="EB81C463FFA90060FF7BFE9FFA0BFD77" box="[137,1450,274,647]" gridcols="5" gridrows="14" pageId="14" pageNumber="15">
|
||
<tr id="27B13481FFA90060FF7BFE9FFA0BFECF" box="[137,1450,274,319]" gridrow="0" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFE9FFE72FECF" box="[137,467,274,319]" gridcol="0" gridrow="0" pageId="14" pageNumber="15">Species</th>
|
||
<th id="64605DFDFFA90060FDD4FE9FFD5FFECF" box="[550,766,274,319]" gridcol="1" gridrow="0" pageId="14" pageNumber="15">Specimens</th>
|
||
<th id="64605DFDFFA90060FCA3FE9FFC70FECF" box="[849,977,274,319]" gridcol="2" gridrow="0" pageId="14" pageNumber="15">Femur length (mm)</th>
|
||
<th id="64605DFDFFA90060FBD6FE9FFB52FECF" box="[1060,1267,274,319]" gridcol="3" gridrow="0" pageId="14" pageNumber="15">Femur circumference (mm)</th>
|
||
<th id="64605DFDFFA90060FAB4FE9FFA0BFECF" box="[1350,1450,274,319]" gridcol="4" gridrow="0" pageId="14" pageNumber="15">Body mass (kg)</th>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFEDFFA0BFE97" box="[137,1450,338,359]" gridrow="1" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFEDFFE72FE97" box="[137,467,338,359]" gridcol="0" gridrow="1" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFEDFFEECFE97" authorityName="Osborn" authorityYear="1905" box="[137,333,338,359]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFEDFFEECFE97" box="[137,333,338,359]" italics="true" pageId="14" pageNumber="15">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FEDFFD5FFE97" box="[550,766,338,359]" gridcol="1" gridrow="1" pageId="14" pageNumber="15">
|
||
<materialsCitation id="29E93C9EFFA9FF91FDD4FEDFFD15FE97" ID-GBIF-Occurrence="2421817399" box="[550,692,338,359]" collectingDate="1991" collectionCode="RSM" country="Canada" location="near Eastend" pageId="14" pageNumber="15" preparations="relatively complete (roughly 65%)" specimenCode="RSM P2523.8" stateProvince="Saskatchewan">RSM P2523.8</materialsCitation>
|
||
</td>
|
||
<td id="64605DFDFFA90060FCA3FEDFFC70FE97" box="[849,977,338,359]" gridcol="2" gridrow="1" pageId="14" pageNumber="15">1,330</td>
|
||
<td id="64605DFDFFA90060FBD6FEDFFB52FE97" box="[1060,1267,338,359]" gridcol="3" gridrow="1" pageId="14" pageNumber="15">590</td>
|
||
<td id="64605DFDFFA90060FAB4FEDFFA0BFE97" box="[1350,1450,338,359]" gridcol="4" gridrow="1" pageId="14" pageNumber="15">8,870</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFEE4FA0BFE8E" box="[137,1450,361,382]" gridrow="2" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFEE4FE72FE8E" box="[137,467,361,382]" gridcol="0" gridrow="2" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFEE4FE08FE8E" box="[137,425,361,382]" class="Reptilia" family="Carcharodontosauridae" genus="Acrocanthosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="atokensis">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFEE4FE08FE8E" box="[137,425,361,382]" italics="true" pageId="14" pageNumber="15">Acrocanthosaurus atokensis</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FEE4FD5FFE8E" box="[550,766,361,382]" gridcol="1" gridrow="2" pageId="14" pageNumber="15">NCSM 14345</td>
|
||
<td id="64605DFDFFA90060FCA3FEE4FC70FE8E" box="[849,977,361,382]" gridcol="2" gridrow="2" pageId="14" pageNumber="15">1,277</td>
|
||
<td id="64605DFDFFA90060FBD6FEE4FB52FE8E" box="[1060,1267,361,382]" gridcol="3" gridrow="2" pageId="14" pageNumber="15">425</td>
|
||
<td id="64605DFDFFA90060FAB4FEE4FA0BFE8E" box="[1350,1450,361,382]" gridcol="4" gridrow="2" pageId="14" pageNumber="15">3,590</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFE0CFA0BFE66" box="[137,1450,385,406]" gridrow="3" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFE0CFE72FE66" box="[137,467,385,406]" gridcol="0" gridrow="3" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFE0CFE72FE66" authorityName="Hu" authorityYear="1964" box="[137,467,385,406]" class="Reptilia" family="Neovenatoridae" genus="Chilantaisaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="tashuikouensis">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFE0CFE72FE66" box="[137,467,385,406]" italics="true" pageId="14" pageNumber="15">Chilantaisaurus tashuikouensis</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FE0CFD5FFE66" box="[550,766,385,406]" gridcol="1" gridrow="3" pageId="14" pageNumber="15">IVPP V.2884</td>
|
||
<td id="64605DFDFFA90060FCA3FE0CFC70FE66" box="[849,977,385,406]" gridcol="2" gridrow="3" pageId="14" pageNumber="15">1,190</td>
|
||
<td id="64605DFDFFA90060FBD6FE0CFB52FE66" box="[1060,1267,385,406]" gridcol="3" gridrow="3" pageId="14" pageNumber="15">430</td>
|
||
<td id="64605DFDFFA90060FAB4FE0CFA0BFE66" box="[1350,1450,385,406]" gridcol="4" gridrow="3" pageId="14" pageNumber="15">3,710</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFE17FA0BFE5F" box="[137,1450,410,431]" gridrow="4" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFE17FE72FE5F" box="[137,467,410,431]" gridcol="0" gridrow="4" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFE17FED3FE5F" box="[137,370,410,431]" class="Reptilia" family="Deinocheiridae" genus="Deinocheirus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="mirificus">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFE17FED3FE5F" box="[137,370,410,431]" italics="true" pageId="14" pageNumber="15">Deinocheirus mirificus</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FE17FD5FFE5F" box="[550,766,410,431]" gridcol="1" gridrow="4" pageId="14" pageNumber="15">MPC KID447</td>
|
||
<td id="64605DFDFFA90060FCA3FE17FC70FE5F" box="[849,977,410,431]" gridcol="2" gridrow="4" pageId="14" pageNumber="15">1,300</td>
|
||
<td id="64605DFDFFA90060FBD6FE17FB52FE5F" box="[1060,1267,410,431]" gridcol="3" gridrow="4" pageId="14" pageNumber="15">520</td>
|
||
<td id="64605DFDFFA90060FAB4FE17FA0BFE5F" box="[1350,1450,410,431]" gridcol="4" gridrow="4" pageId="14" pageNumber="15">6,260</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFE3FFA0BFE37" box="[137,1450,434,455]" gridrow="5" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFE3FFE72FE37" box="[137,467,434,455]" gridcol="0" gridrow="5" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFE3FFE2AFE37" box="[137,395,434,455]" class="Reptilia" family="Carcharodontosauridae" genus="Giganotosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="carolinii">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFE3FFE2AFE37" box="[137,395,434,455]" italics="true" pageId="14" pageNumber="15">Giganotosaurus carolinii</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FE3FFD5FFE37" box="[550,766,434,455]" gridcol="1" gridrow="5" pageId="14" pageNumber="15">MUCPv-CH 1</td>
|
||
<td id="64605DFDFFA90060FCA3FE3FFC70FE37" box="[849,977,434,455]" gridcol="2" gridrow="5" pageId="14" pageNumber="15">1,365</td>
|
||
<td id="64605DFDFFA90060FBD6FE3FFB52FE37" box="[1060,1267,434,455]" gridcol="3" gridrow="5" pageId="14" pageNumber="15">520</td>
|
||
<td id="64605DFDFFA90060FAB4FE3FFA0BFE37" box="[1350,1450,434,455]" gridcol="4" gridrow="5" pageId="14" pageNumber="15">6,260</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFE44FA0BFE2F" box="[137,1450,457,479]" gridrow="6" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFE44FE72FE2F" box="[137,467,457,479]" gridcol="0" gridrow="6" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFE47FEE2FE2F" box="[137,323,458,479]" class="Reptilia" family="Carcharodontosauridae" genus="Mapusaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rosei">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFE47FEE2FE2F" box="[137,323,458,479]" italics="true" pageId="14" pageNumber="15">Mapusaurus rosei</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FE44FD5FFE2F" box="[550,766,457,479]" gridcol="1" gridrow="6" pageId="14" pageNumber="15">PVPH.108.44</td>
|
||
<td id="64605DFDFFA90060FCA3FE44FC70FE2F" box="[849,977,457,479]" gridcol="2" gridrow="6" pageId="14" pageNumber="15">1,170</td>
|
||
<td id="64605DFDFFA90060FBD6FE44FB52FE2F" box="[1060,1267,457,479]" gridcol="3" gridrow="6" pageId="14" pageNumber="15">435</td>
|
||
<td id="64605DFDFFA90060FAB4FE44FA0BFE2F" box="[1350,1450,457,479]" gridcol="4" gridrow="6" pageId="14" pageNumber="15">3,830</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFE6CFA0BFE07" box="[137,1450,481,503]" gridrow="7" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFE6CFE72FE07" box="[137,467,481,503]" gridcol="0" gridrow="7" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFE6FFEE2FE07" box="[137,323,482,503]" class="Reptilia" family="Carcharodontosauridae" genus="Mapusaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rosei">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFE6FFEE2FE07" box="[137,323,482,503]" italics="true" pageId="14" pageNumber="15">Mapusaurus rosei</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FE6CFD5FFE07" box="[550,766,481,503]" gridcol="1" gridrow="7" pageId="14" pageNumber="15">PVPH.108.203</td>
|
||
<td id="64605DFDFFA90060FCA3FE6CFC70FE07" box="[849,977,481,503]" gridcol="2" gridrow="7" pageId="14" pageNumber="15">1,205</td>
|
||
<td id="64605DFDFFA90060FBD6FE6CFB52FE07" box="[1060,1267,481,503]" gridcol="3" gridrow="7" pageId="14" pageNumber="15">415</td>
|
||
<td id="64605DFDFFA90060FAB4FE6CFA0BFE07" box="[1350,1450,481,503]" gridcol="4" gridrow="7" pageId="14" pageNumber="15">3,370</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFE77FA0BFDFF" box="[137,1450,506,527]" gridrow="8" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFE77FE72FDFF" box="[137,467,506,527]" gridcol="0" gridrow="8" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFE77FEE2FDFF" box="[137,323,506,527]" class="Reptilia" family="Carcharodontosauridae" genus="Mapusaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rosei">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFE77FEE2FDFF" box="[137,323,506,527]" italics="true" pageId="14" pageNumber="15">Mapusaurus rosei</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FE77FD5FFDFF" box="[550,766,506,527]" gridcol="1" gridrow="8" pageId="14" pageNumber="15">PVPH.108.233</td>
|
||
<td id="64605DFDFFA90060FCA3FE77FC70FDFF" box="[849,977,506,527]" gridcol="2" gridrow="8" pageId="14" pageNumber="15">1,115</td>
|
||
<td id="64605DFDFFA90060FBD6FE77FB52FDFF" box="[1060,1267,506,527]" gridcol="3" gridrow="8" pageId="14" pageNumber="15">420</td>
|
||
<td id="64605DFDFFA90060FAB4FE77FA0BFDFF" box="[1350,1450,506,527]" gridcol="4" gridrow="8" pageId="14" pageNumber="15">3,480</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFD9FFA0BFDD7" box="[137,1450,530,551]" gridrow="9" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFD9FFE72FDD7" box="[137,467,530,551]" gridcol="0" gridrow="9" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFD9FFE34FDD7" baseAuthorityName="Chure" baseAuthorityYear="1995" box="[137,405,530,551]" class="Reptilia" family="Allosauridae" genus="Saurophaganax" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="maximus">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFD9FFE34FDD7" box="[137,405,530,551]" italics="true" pageId="14" pageNumber="15">Saurophaganax maximus</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FD9FFD5FFDD7" box="[550,766,530,551]" gridcol="1" gridrow="9" pageId="14" pageNumber="15">OMNH 01708</td>
|
||
<td id="64605DFDFFA90060FCA3FD9FFC70FDD7" box="[849,977,530,551]" gridcol="2" gridrow="9" pageId="14" pageNumber="15">1,130</td>
|
||
<td id="64605DFDFFA90060FBD6FD9FFB52FDD7" box="[1060,1267,530,551]" gridcol="3" gridrow="9" pageId="14" pageNumber="15">435</td>
|
||
<td id="64605DFDFFA90060FAB4FD9FFA0BFDD7" box="[1350,1450,530,551]" gridcol="4" gridrow="9" pageId="14" pageNumber="15">3,830</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFDA4FA0BFDCF" box="[137,1450,553,575]" gridrow="10" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFDA4FE72FDCF" box="[137,467,553,575]" gridcol="0" gridrow="10" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFDA4FE2AFDCF" authorityName="Stromer" authorityYear="1915" box="[137,395,553,575]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFDA4FE2AFDCF" box="[137,395,553,575]" italics="true" pageId="14" pageNumber="15">Spinosaurus aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FDA4FD5FFDCF" box="[550,766,553,575]" gridcol="1" gridrow="10" pageId="14" pageNumber="15">
|
||
<bibRefCitation id="FD104B32FFA9FF91FDD4FDA4FD5FFDCE" author="Ibrahim N & Sereno PC & Dal Sasso C & Maganuco S & Fabbri M & Martill DM & Zouhri S & Myhrvold N & Iurino DA" box="[550,766,553,574]" journalOrPublisher="Science" pageId="14" pageNumber="15" pagination="1613 - 1616" part="345" refId="ref9341" refString="Ibrahim N, Sereno PC, Dal Sasso C, Maganuco S, Fabbri M, Martill DM, Zouhri S, Myhrvold N, Iurino DA. 2014. Semiaquatic adaptations in a giant predatory dinosaur. Science 345: 1613 - 1616." title="Semiaquatic adaptations in a giant predatory dinosaur" type="journal article" year="2014">Ibrahim et al. (2014)</bibRefCitation>
|
||
</td>
|
||
<td id="64605DFDFFA90060FCA3FDA4FC70FDCF" box="[849,977,553,575]" gridcol="2" gridrow="10" pageId="14" pageNumber="15">830</td>
|
||
<td id="64605DFDFFA90060FBD6FDA4FB52FDCF" box="[1060,1267,553,575]" gridcol="3" gridrow="10" pageId="14" pageNumber="15">320</td>
|
||
<td id="64605DFDFFA90060FAB4FDA4FA0BFDCF" box="[1350,1450,553,575]" gridcol="4" gridrow="10" pageId="14" pageNumber="15">1,645</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFDCCFA0BFDA7" box="[137,1450,577,599]" gridrow="11" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFDCCFE72FDA7" box="[137,467,577,599]" gridcol="0" gridrow="11" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFDCCFEDCFDA7" box="[137,381,577,599]" class="Reptilia" family="Spinosauridae" genus="Suchomimus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="tenerensis">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFDCCFEDCFDA7" box="[137,381,577,599]" italics="true" pageId="14" pageNumber="15">Suchomimus tenerensis</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FDCCFD5FFDA7" box="[550,766,577,599]" gridcol="1" gridrow="11" pageId="14" pageNumber="15">MNN GDF 500</td>
|
||
<td id="64605DFDFFA90060FCA3FDCCFC70FDA7" box="[849,977,577,599]" gridcol="2" gridrow="11" pageId="14" pageNumber="15">1,080</td>
|
||
<td id="64605DFDFFA90060FBD6FDCCFB52FDA7" box="[1060,1267,577,599]" gridcol="3" gridrow="11" pageId="14" pageNumber="15">410</td>
|
||
<td id="64605DFDFFA90060FAB4FDCCFA0BFDA7" box="[1350,1450,577,599]" gridcol="4" gridrow="11" pageId="14" pageNumber="15">3,260</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFDD7FA0BFD9F" box="[137,1450,602,623]" gridrow="12" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFDD7FE72FD9F" box="[137,467,602,623]" gridcol="0" gridrow="12" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFDD7FE35FD9F" box="[137,404,602,623]" class="Reptilia" family="Carcharodontosauridae" genus="Tyrannotitan" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="chubutensis">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFDD7FE35FD9F" box="[137,404,602,623]" italics="true" pageId="14" pageNumber="15">Tyrannotitan chubutensis</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FDD7FD5FFD9F" box="[550,766,602,623]" gridcol="1" gridrow="12" pageId="14" pageNumber="15">MPEF-PV 1156</td>
|
||
<td id="64605DFDFFA90060FCA3FDD7FC70FD9F" box="[849,977,602,623]" gridcol="2" gridrow="12" pageId="14" pageNumber="15">1,265</td>
|
||
<td id="64605DFDFFA90060FBD6FDD7FB52FD9F" box="[1060,1267,602,623]" gridcol="3" gridrow="12" pageId="14" pageNumber="15">520</td>
|
||
<td id="64605DFDFFA90060FAB4FDD7FA0BFD9F" box="[1350,1450,602,623]" gridcol="4" gridrow="12" pageId="14" pageNumber="15">6,260</td>
|
||
</tr>
|
||
<tr id="27B13481FFA90060FF7BFDFFFA0BFD77" box="[137,1450,626,647]" gridrow="13" pageId="14" pageNumber="15">
|
||
<th id="64605DFDFFA90060FF7BFDFFFE72FD77" box="[137,467,626,647]" gridcol="0" gridrow="13" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BFDFFFE35FD77" box="[137,404,626,647]" class="Reptilia" family="Carcharodontosauridae" genus="Tyrannotitan" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="chubutensis">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFDFFFE35FD77" box="[137,404,626,647]" italics="true" pageId="14" pageNumber="15">Tyrannotitan chubutensis</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="64605DFDFFA90060FDD4FDFFFD5FFD77" box="[550,766,626,647]" gridcol="1" gridrow="13" pageId="14" pageNumber="15">MPEF-PV 1157</td>
|
||
<td id="64605DFDFFA90060FCA3FDFFFC70FD77" box="[849,977,626,647]" gridcol="2" gridrow="13" pageId="14" pageNumber="15">1,330</td>
|
||
<td id="64605DFDFFA90060FBD6FDFFFB52FD77" box="[1060,1267,626,647]" gridcol="3" gridrow="13" pageId="14" pageNumber="15">505</td>
|
||
<td id="64605DFDFFA90060FAB4FDFFFA0BFD77" box="[1350,1450,626,647]" gridcol="4" gridrow="13" pageId="14" pageNumber="15">5,780</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA9FF91FF7BFD2FFF58FD3E" blockId="14.[137,1450,673,718]" pageId="14" pageNumber="15">
|
||
<tableNote id="6467374DFFA9FF91FF7BFD2FFF58FD3E" pageId="14" pageNumber="15" targetBox="[137,1450,274,647]" targetPageId="14">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BFD2FFF61FD47" box="[137,192,674,695]" italics="true" pageId="14" pageNumber="15">Note.</emphasis>
|
||
There are a great number of problematic assumptions in the mass estimate of
|
||
<taxonomicName id="5E814D40FFA9FF91FBF8FD2CFAACFD47" authorityName="Stromer" authorityYear="1915" box="[1034,1293,673,695]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="aegyptiacus">
|
||
<emphasis id="ABF5EAD1FFA9FF91FBF8FD2CFAACFD47" box="[1034,1293,673,695]" italics="true" pageId="14" pageNumber="15">Spinosaurus aegyptiacus</emphasis>
|
||
</taxonomicName>
|
||
, see main text for details.
|
||
</tableNote>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA9FF91FF56FBDAFEF7FA5D" blockId="14.[137,778,791,1933]" pageId="14" pageNumber="15">
|
||
Although
|
||
<materialsCitation id="29E93C9EFFA9FF91FEE0FBDAFE0BFB9C" ID-GBIF-Occurrence="2597591342" box="[274,426,1110,1132]" collectionCode="RSM" pageId="14" pageNumber="15" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
has an estimated weight more than 40% greater than the next largest known theropod taxon, specimens of several other theropod species (including
|
||
<taxonomicName id="5E814D40FFA9FF91FF40FB2AFE1BFB4C" box="[178,442,1191,1212]" class="Reptilia" family="Carcharodontosauridae" genus="Giganotosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="carolinii">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF40FB2AFE1BFB4C" box="[178,442,1191,1212]" italics="true" pageId="14" pageNumber="15">Giganotosaurus carolinii</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="5E814D40FFA9FF91FE02FB2AFCA3FB4C" box="[496,770,1191,1212]" class="Reptilia" family="Carcharodontosauridae" genus="Tyrannotitan" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="chubutensis">
|
||
<emphasis id="ABF5EAD1FFA9FF91FE02FB2AFCA3FB4C" box="[496,770,1191,1212]" italics="true" pageId="14" pageNumber="15">Tyrannotitan chubutensis</emphasis>
|
||
</taxonomicName>
|
||
) have femoral proportions indicative of body masses greater than those of most other adult
|
||
<taxonomicName id="5E814D40FFA9FF91FE0DFB50FDE5FB02" authorityName="Osborn" authorityYear="1905" box="[511,580,1245,1266]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA9FF91FE0DFB50FDE5FB02" box="[511,580,1245,1266]" italics="true" pageId="14" pageNumber="15">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens (compare Tables 1 and 2). As such, it is likely that further sampling of these other giant theropods, all of which are represented by fewer specimens than
|
||
<taxonomicName id="5E814D40FFA9FF91FDB6FAA0FD29FAB2" authorityName="Osborn" authorityYear="1905" box="[580,648,1325,1346]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA9FF91FDB6FAA0FD29FAB2" box="[580,648,1325,1346]" italics="true" pageId="14" pageNumber="15">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, may yield larger individuals that match or surpass the size of
|
||
<materialsCitation id="29E93C9EFFA9FF91FD20FACAFF46FA87" ID-GBIF-Occurrence="2597591301" collectionCode="RSM" pageId="14" pageNumber="15" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
. In the case of
|
||
<taxonomicName id="5E814D40FFA9FF91FE69FAEFFD15FA86" box="[411,692,1377,1399]" class="Reptilia" family="Carcharodontosauridae" genus="Giganotosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="carolinii">
|
||
<emphasis id="ABF5EAD1FFA9FF91FE69FAEFFD15FA86" box="[411,692,1377,1399]" italics="true" pageId="14" pageNumber="15">Giganotosaurus carolinii</emphasis>
|
||
</taxonomicName>
|
||
, a sing dentary is known that does hint at a greater maximum size (
|
||
<bibRefCitation id="FD104B32FFA9FF91FF31FA1AFEEAFA5D" author="Calvo JO" box="[195,331,1431,1453]" journalOrPublisher="GAIA" pageId="14" pageNumber="15" pagination="117 - 122" part="15" refId="ref8303" refString="Calvo JO. 2000. New specimen of Giganotosaurus carolinii (Coria & Salgado, 1995), supports it as the largest theropod ever found. GAIA 15: 117 - 122." title="New specimen of Giganotosaurus carolinii (Coria & Salgado, 1995), supports it as the largest theropod ever found" type="journal article" year="2000">Calvo, 2000</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA9FF91FF56FA3CFC61FC8D" blockId="14.[137,778,791,1933]" lastBlockId="14.[810,1451,791,1933]" pageId="14" pageNumber="15">
|
||
Second, although
|
||
<materialsCitation id="29E93C9EFFA9FF91FE9EFA3FFDA4FA37" ID-GBIF-Occurrence="2597591332" box="[364,517,1457,1479]" collectionCode="RSM" pageId="14" pageNumber="15" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
has the greatest femoral circumference among giant theropods (
|
||
<tableCitation id="D4030378FFA9FF91FD70FA41FD7FFA12" box="[642,734,1484,1506]" captionStart="TABLE 2" captionStartId="14.[168,252,205,226]" captionTargetPageId="14" captionText="TABLE 2. Comparison of femoral dimensions and estimated body mass of the largest theropods ±25% (mass estimation based on the formula of Campione et al. (2014))" httpUri="http://table.plazi.org/id/CDFE664BFFA9FF91FF5AFF40FBCCFF0A" pageId="14" pageNumber="15" tableUuid="CDFE664BFFA9FF91FF5AFF40FBCCFF0A">Table 2</tableCitation>
|
||
), it does not show the greatest femoral length (the femora of
|
||
<taxonomicName id="5E814D40FFA9FF91FF7BF98FFE0CF9E6" box="[137,429,1537,1559]" class="Reptilia" family="Carcharodontosauridae" genus="Tyrannotitan" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="chubutensis">
|
||
<emphasis id="ABF5EAD1FFA9FF91FF7BF98FFE0CF9E6" box="[137,429,1537,1559]" italics="true" pageId="14" pageNumber="15">Tyrannotitan chubutensis</emphasis>
|
||
</taxonomicName>
|
||
MPEF-PV 1157 are equally long and those for
|
||
<taxonomicName id="5E814D40FFA9FF91FE94F991FDDBF9C1" box="[358,634,1564,1585]" class="Reptilia" family="Carcharodontosauridae" genus="Giganotosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="carolinii">
|
||
<emphasis id="ABF5EAD1FFA9FF91FE94F991FDDBF9C1" box="[358,634,1564,1585]" italics="true" pageId="14" pageNumber="15">Giganotosaurus carolinii</emphasis>
|
||
</taxonomicName>
|
||
MUCPv-CH 1 are roughly 10% longer). Though a less reliable indicator of body mass than MFC (
|
||
<bibRefCitation id="FD104B32FFA9FF91FE34F9DCFD1AF998" author="Campione NE & Evans DC & Brown CM & Carrano MT" box="[454,699,1617,1640]" journalOrPublisher="Methods Ecol Evol" pageId="14" pageNumber="15" pagination="913 - 923" part="5" refId="ref8337" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol Evol 5: 913 - 923." title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">Campione et al., 2014</bibRefCitation>
|
||
), femoral length is generally considered an isometrically stable feature across theropods. However, rather than being proportionately short-legged, the other hindlimb bones of tyrannosaurids are far more elongate than those of carcharodontosaurids (
|
||
<taxonomicName id="5E814D40FFA9FF91FE94F95AFE45F91C" box="[358,484,1751,1772]" class="Reptilia" family="Carcharodontosauridae" genus="Giganotosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="carolinii">
|
||
<emphasis id="ABF5EAD1FFA9FF91FE94F95AFE45F91C" box="[358,484,1751,1772]" italics="true" pageId="14" pageNumber="15">G. carolinii</emphasis>
|
||
</taxonomicName>
|
||
has a tibia length of only 1,060 mm, compared with 1,140 mm in
|
||
<materialsCitation id="29E93C9EFFA9FF91FDADF97FFD5EF8F7" ID-GBIF-Occurrence="2597591344" box="[607,767,1777,1799]" collectionCode="RSM" pageId="14" pageNumber="15" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
). Second, although femoral circumference has emerged as the standard for dinosaur mass estimation (and has been rigorously calibrated and tested in studies of extant animals), the method still suffers from a considerable range of error (variation from the true value of as much as 20% is generally expected) (
|
||
<bibRefCitation id="FD104B32FFA9FF91FBDFFC9AFA89FCDD" author="Campione NE & Evans DC & Brown CM & Carrano MT" box="[1069,1320,791,813]" journalOrPublisher="Methods Ecol Evol" pageId="14" pageNumber="15" pagination="913 - 923" part="5" refId="ref8337" refString="Campione NE, Evans DC, Brown CM, Carrano MT. 2014. Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions. Methods Ecol Evol 5: 913 - 923." title="Body mass estimation in non-avian bipeds using a theoretical conversion to quadruped stylopodial proportions" type="journal article" year="2014">Campione et al., 2014</bibRefCitation>
|
||
). Moreover, applying the method to comparisons between tyrannosaurs and other large theropods may constitute a particular challenge.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA9FF91FCB6FC0CFB43FA37" blockId="14.[810,1451,791,1933]" pageId="14" pageNumber="15">
|
||
There has been much debate over the absolute running potential (or lack thereof) of large tyrannosauroids, but however fast or slowly tyrannosauroids could run/walk, the comparative anatomical evidence indicates a relatively greater degree of limb exertion than in other large theropods (
|
||
<bibRefCitation id="FD104B32FFA9FF91FC5DFB8AFBF3FBEC" author="Snively E." box="[943,1106,1031,1053]" journalOrPublisher="University of Calgary" pageId="14" pageNumber="15" refId="ref9699" refString="Snively E. 2000. Functional morphology of the tyrannosaurid arctometatarsus. University of Calgary." title="Functional morphology of the tyrannosaurid arctometatarsus" type="book" year="2000">Snively, 2000</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA9FF91FB95FB8AFA07FBEC" author="Snively E & Russell AP" box="[1127,1446,1031,1053]" journalOrPublisher="J Morphol" pageId="14" pageNumber="15" pagination="215 - 227" part="255" refId="ref9715" refString="Snively E, Russell AP. 2003. Kinematic model of tyrannosaurid (Dinosauria: Theropoda) arctometatarsus function. J Morphol 255: 215 - 227." title="Kinematic model of tyrannosaurid (Dinosauria: Theropoda) arctometatarsus function" type="journal article" year="2003">Snively and Russell, 2003</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA9FF91FCD8FBAFFBEFFBC7" author="Persons WS & Currie PJ" box="[810,1102,1057,1079]" journalOrPublisher="Sci Rep" pageId="14" pageNumber="15" pagination="19828" part="6" refId="ref9610" refString="Persons WS, Currie PJ. 2016. An approach to scoring cursorial limb proportions in carnivorous dinosaurs and an attempt to account for allometry. Sci Rep 6: 19828." title="An approach to scoring cursorial limb proportions in carnivorous dinosaurs and an attempt to account for allometry" type="journal article" year="2016">Persons and Currie, 2016</bibRefCitation>
|
||
). The underlining assumptions of femoral circumference based mass estimations are that minimum femoral circumferences reflect femoral crosssectional strength, that in turn reflect the amount of stress the femur was suited to withstand, that in turn reflects how much body mass the femur needed to support. However, body mass alone is not the only factor contributing to the stress and strain applied to a limb bone. Locomotor loads from muscle, joint, and ground reaction forces are also factors. During faster and more vigorous movement, limb bones will experience greater strain than during slower and less vigorous movements. As such, when comparing the femur circumferences of two animals, it is worth considering whether the two are expected to have experienced a similar degree of limb exertion and locomotory reaction forces.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA9FF90FCB6FA41FEB7FEC3" blockId="14.[810,1451,791,1933]" lastBlockId="15.[137,778,205,1054]" lastPageId="15" lastPageNumber="16" pageId="14" pageNumber="15">
|
||
Tyrannosaurids are characterized by numerous anatomical features that suggest the group was capable of more rapid locomotion than other similarly sized theropods (including large allosauroids). These traits include an arctometatarsalian pes (
|
||
<bibRefCitation id="FD104B32FFA9FF91FB9CF9BAFB57F9BD" author="Holtz TR" box="[1134,1270,1591,1613]" journalOrPublisher="J Vertebr Paleontol" pageId="14" pageNumber="15" pagination="480 - 519" part="14" refId="ref8916" refString="Holtz TR. 1995. The arctometatarsalian pes, an unusual structure of the metatarsus of Cretaceous Theropoda (Dinosauria: Saurischia). J Vertebr Paleontol 14: 480 - 519." title="The arctometatarsalian pes, an unusual structure of the metatarsus of Cretaceous Theropoda (Dinosauria: Saurischia)" type="journal article" year="1995">Holtz, 1995</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA9FF91FAFAF9BAFA06F9BC" author="Snively E." box="[1288,1447,1591,1613]" journalOrPublisher="University of Calgary" pageId="14" pageNumber="15" refId="ref9699" refString="Snively E. 2000. Functional morphology of the tyrannosaurid arctometatarsus. University of Calgary." title="Functional morphology of the tyrannosaurid arctometatarsus" type="book" year="2000">Snively, 2000</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA9FF91FCD8F9DCFBF2F998" author="Snively E & Russell AP" box="[810,1107,1617,1640]" journalOrPublisher="J Morphol" pageId="14" pageNumber="15" pagination="215 - 227" part="255" refId="ref9715" refString="Snively E, Russell AP. 2003. Kinematic model of tyrannosaurid (Dinosauria: Theropoda) arctometatarsus function. J Morphol 255: 215 - 227." title="Kinematic model of tyrannosaurid (Dinosauria: Theropoda) arctometatarsus function" type="journal article" year="2003">Snively and Russell, 2003</bibRefCitation>
|
||
), proportionately longer distal limb bones (
|
||
<bibRefCitation id="FD104B32FFA9FF91FC4AF9E1FB14F971" author="Bakker RT & Bir G." box="[952,1205,1644,1666]" journalOrPublisher="Bloomington, IN: Indiana University Press" pageId="14" pageNumber="15" pagination="301" part="Vol. 14" refId="ref7979" refString="Bakker RT, Bir G. 2004. Dinosaur crime scene investigations: theropod behavior at Como Bluff, Wyoming, and the evolution of birdness. Feathered dragons: studies on the transition from dinosaurs to birds, Bloomington, IN: Indiana University Press. Vol. 14. p 301." title="Dinosaur crime scene investigations: theropod behavior at Como Bluff, Wyoming, and the evolution of birdness." type="book chapter" volumeTitle="Feathered dragons: studies on the transition from dinosaurs to birds" year="2004">Bakker and Bir, 2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA9FF91FB36F9E0FCC3F96D" author="Persons WS & Currie PJ" journalOrPublisher="Sci Rep" pageId="14" pageNumber="15" pagination="19828" part="6" refId="ref9610" refString="Persons WS, Currie PJ. 2016. An approach to scoring cursorial limb proportions in carnivorous dinosaurs and an attempt to account for allometry. Sci Rep 6: 19828." title="An approach to scoring cursorial limb proportions in carnivorous dinosaurs and an attempt to account for allometry" type="journal article" year="2016">Persons and Currie, 2016</bibRefCitation>
|
||
), ilia with increased muscle attachment surfaces, and more rigid posterior caudal series with potential to better aid in turning and stability (
|
||
<bibRefCitation id="FD104B32FFA9FF91FB17F931FCC2F91D" author="Bakker RT & Bir G." journalOrPublisher="Bloomington, IN: Indiana University Press" pageId="14" pageNumber="15" pagination="301" part="Vol. 14" refId="ref7979" refString="Bakker RT, Bir G. 2004. Dinosaur crime scene investigations: theropod behavior at Como Bluff, Wyoming, and the evolution of birdness. Feathered dragons: studies on the transition from dinosaurs to birds, Bloomington, IN: Indiana University Press. Vol. 14. p 301." title="Dinosaur crime scene investigations: theropod behavior at Como Bluff, Wyoming, and the evolution of birdness." type="book chapter" volumeTitle="Feathered dragons: studies on the transition from dinosaurs to birds" year="2004">Bakker and Bir, 2004</bibRefCitation>
|
||
). Greater femoral circumference with the capacity to withstand greater locomotor loads, rather than simply greater body mass, merits consideration as an explanation for the discrepancies between femoral lengths and girths in
|
||
<taxonomicName id="5E814D40FFA9FF91FC7CF8CFFC6DF8A7" authorityName="Osborn" authorityYear="1905" box="[910,972,1858,1879]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA9FF91FC7CF8CFFC6DF8A7" box="[910,972,1858,1879]" italics="true" pageId="14" pageNumber="15">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis id="ABF5EAD1FFA9FF91FBF1F8CFFB22F8A6" box="[1027,1155,1857,1879]" italics="true" pageId="14" pageNumber="15">
|
||
<taxonomicName id="5E814D40FFA9FF91FBF1F8CFFBDEF8A6" box="[1027,1151,1857,1879]" class="Reptilia" family="Carcharodontosauridae" genus="Giganotosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="carolinii">G. carolinii</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Relative locomotor activity may similarly explain the extreme discrepancies in weight estimations reported for the proportionately slim-femured carcharodontosaurid
|
||
<emphasis id="ABF5EAD1FFA8FF90FE8FFF40FD60FF12" box="[381,705,205,226]" italics="true" pageId="15" pageNumber="16">
|
||
<taxonomicName id="5E814D40FFA8FF90FE8FFF40FD1FFF12" box="[381,702,205,226]" class="Reptilia" family="Carcharodontosauridae" genus="Acrocanthosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="atokensis">Acrocanthosaurus atokensis</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
Notably the 3.5 tonnes femur derived estimation for mass vs. 5.5–7 tonnes based on volumetric estimation (
|
||
<bibRefCitation id="FD104B32FFA8FF90FD3BFE8FFEABFEC3" author="Bates KT & Manning PL & Hodgetts D & Sellers WI" journalOrPublisher="PLoS One" pageId="15" pageNumber="16" pagination="e 4532" part="4" refId="ref8033" refString="Bates KT, Manning PL, Hodgetts D, Sellers WI. 2009. Estimating mass properties of dinosaurs using laser imaging and 3 D computer modelling. PLoS One 4: e 4532." title="Estimating mass properties of dinosaurs using laser imaging and 3 D computer modelling" type="journal article" year="2009">Bates et al., 2009</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA8FF90FF56FEB5FE99FBEE" blockId="15.[137,778,205,1054]" pageId="15" pageNumber="16">
|
||
Various authors have suggested a biomechanical ceiling that caps the body mass at which the bipedal theropod body plan can function with ecologically feasible efficacy (
|
||
<bibRefCitation id="FD104B32FFA8FF90FF62FE05FE67FE6D" author="Hutchinson JR & Garcia M." box="[144,454,392,414]" journalOrPublisher="Nature" pageId="15" pageNumber="16" pagination="1018 - 1021" part="415" refId="ref9184" refString="Hutchinson JR, Garcia M. 2002. Tyrannosaurus was not a fast runner. Nature 415: 1018 - 1021." title="Tyrannosaurus was not a fast runner" type="journal article" year="2002">Hutchinson and Garcia, 2002</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA8FF90FE22FE05FCA4FE6D" author="Henderson DM & Snively E." box="[464,773,392,414]" journalOrPublisher="Proc Biol Sci" pageId="15" pageNumber="16" pagination="S 57 - S 60" part="271 (Suppl 3)" refId="ref8880" refString="Henderson DM, Snively E. 2004. Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs. Proc Biol Sci 271 (Suppl 3): S 57 - S 60." title="Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs." type="journal article" year="2004">Henderson and Snively, 2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA8FF90FF7BFE2FFE75FE48" author="Therrien F & Henderson DM" box="[137,468,418,440]" journalOrPublisher="J Vertebr Paleontol" pageId="15" pageNumber="16" pagination="108 - 115" part="27" refId="ref9743" refString="Therrien F, Henderson DM. 2007. My theropod is bigger than yours … or not: estimating body size from skull length in theropods. J Vertebr Paleontol 27: 108 - 115." title="My theropod is bigger than yours … or not: estimating body size from skull length in theropods" type="journal article" year="2007">Therrien and Henderson, 2007</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA8FF90FE12FE2FFD7AFE49" author="Hutchinson JR & Bates KT & Molnar J & Allen V & Makovicky PJ" box="[480,731,418,441]" journalOrPublisher="PLoS One" pageId="15" pageNumber="16" pagination="e 26037" part="6" refId="ref9296" refString="Hutchinson JR, Bates KT, Molnar J, Allen V, Makovicky PJ. 2011. A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth. PLoS One 6: e 26037." title="A computational analysis of limb and body dimensions in Tyrannosaurus rex with implications for locomotion, ontogeny, and growth" type="journal article" year="2011">Hutchinson et al., 2011</bibRefCitation>
|
||
). As this ceiling is approached, the typical advantages of greater body mass are thought to be increasingly opposed by sizedependent restrictions on speed and agility, including bone and joint strength, and musculoskeletal torques (
|
||
<bibRefCitation id="FD104B32FFA8FF90FD4DFD80FEA0FDCD" author="Farlow JO & Smith MB & Robinson JM" journalOrPublisher="J Vertebr Paleontol" pageId="15" pageNumber="16" pagination="713 - 725" part="15" refId="ref8780" refString="Farlow JO, Smith MB, Robinson JM. 1995. Body mass, bone " strength indicator, " and cursorial potential of Tyrannosaurus rex. J Vertebr Paleontol 15: 713 - 725." title="Body mass, bone " strength indicator, " and cursorial potential of Tyrannosaurus rex" type="journal article" year="1995">Farlow et al., 1995</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA8FF90FEF9FDA5FDE2FDCD" author="Hutchinson JR & Garcia M." box="[267,579,552,574]" journalOrPublisher="Nature" pageId="15" pageNumber="16" pagination="1018 - 1021" part="415" refId="ref9184" refString="Hutchinson JR, Garcia M. 2002. Tyrannosaurus was not a fast runner. Nature 415: 1018 - 1021." title="Tyrannosaurus was not a fast runner" type="journal article" year="2002">Hutchinson and Garcia, 2002</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA8FF90FDBCFDA5FF1FFDA8" author="Hutchinson JR & Anderson FC & Blemker SS & Delp SL" journalOrPublisher="Paleobiology" pageId="15" pageNumber="16" pagination="676 - 701" part="31" refId="ref9206" refString="Hutchinson JR, Anderson FC, Blemker SS, Delp SL. 2005. Analysis of hindlimb muscle moment arms in Tyrannosaurus rex using a three-dimensional musculoskeletal computer model: implications for stance, gait, and speed. Paleobiology 31: 676 - 701." title="Analysis of hindlimb muscle moment arms in Tyrannosaurus rex using a three-dimensional musculoskeletal computer model: implications for stance, gait, and speed" type="journal article" year="2005">Hutchinson et al., 2005</bibRefCitation>
|
||
), biomechanical safety factors (
|
||
<bibRefCitation id="FD104B32FFA8FF90FDF4FDCFFD6EFDA8" author="Farlow JO & Smith MB & Robinson JM" box="[518,719,578,600]" journalOrPublisher="J Vertebr Paleontol" pageId="15" pageNumber="16" pagination="713 - 725" part="15" refId="ref8780" refString="Farlow JO, Smith MB, Robinson JM. 1995. Body mass, bone " strength indicator, " and cursorial potential of Tyrannosaurus rex. J Vertebr Paleontol 15: 713 - 725." title="Body mass, bone " strength indicator, " and cursorial potential of Tyrannosaurus rex" type="journal article" year="1995">Farlow et al., 1995</bibRefCitation>
|
||
), and sheer inertia (
|
||
<bibRefCitation id="FD104B32FFA8FF90FEDFFDD0FDDCFD82" author="Henderson DM & Snively E." box="[301,637,605,627]" journalOrPublisher="Proc Biol Sci" pageId="15" pageNumber="16" pagination="S 57 - S 60" part="271 (Suppl 3)" refId="ref8880" refString="Henderson DM, Snively E. 2004. Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs. Proc Biol Sci 271 (Suppl 3): S 57 - S 60." title="Tyrannosaurus en pointe: allometry minimized rotational inertia of large carnivorous dinosaurs." type="journal article" year="2004">Henderson and Snively, 2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="FD104B32FFA8FF90FD7DFDD0FEBBFD7E" author="Hutchinson JR & Ng-Thow-Hing V & Anderson FC" journalOrPublisher="J Theor Biol" pageId="15" pageNumber="16" pagination="660 - 680" part="246" refId="ref9252" refString="Hutchinson JR, Ng-Thow-Hing V, Anderson FC. 2007. A 3 D interactive method for estimating body segmental parameters in animals: application to the turning and running performance of Tyrannosaurus rex. J Theor Biol 246: 660 - 680." title="A 3 D interactive method for estimating body segmental parameters in animals: application to the turning and running performance of Tyrannosaurus rex" type="journal article" year="2007">Hutchinson et al., 2007</bibRefCitation>
|
||
). Extreme giantism in Allosauroidea and
|
||
<taxonomicName id="5E814D40FFA8FF90FF7BFD1FFE81FD58" authorityName="Stromer" authorityYear="1915" box="[137,288,658,680]" class="Reptilia" family="Spinosauridae" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="16" phylum="Chordata" rank="family">Spinosauridae</taxonomicName>
|
||
may be attributable to factors that mitigate these restrictions and allowed both linages to press this ceiling, such as access to enormous, but correspondingly slow, sauropod prey in the former and the weight-supporting advantages of a semiaquatic lifestyle, in the latter. However, the evolutionary history leading to the scale of
|
||
<materialsCitation id="29E93C9EFFA8FF90FD8BFC95FCA8FCDE" ID-GBIF-Occurrence="2597591320" box="[633,777,792,814]" collectionCode="RSM" pageId="15" pageNumber="16" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
unfolded in a purely terrestrial context. Nor were large sauropods prevalent throughout the range of
|
||
<taxonomicName id="5E814D40FFA8FF90FDDDFCC3FDCAFC93" authorityName="Osborn" authorityYear="1905" box="[559,619,846,867]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA8FF90FDDDFCC3FDCAFC93" box="[559,619,846,867]" italics="true" pageId="15" pageNumber="16">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, and definitive evidence of
|
||
<taxonomicName id="5E814D40FFA8FF90FEF9FCE5FEEBFC8D" authorityName="Osborn" authorityYear="1905" box="[267,330,872,893]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA8FF90FEF9FCE5FEEBFC8D" box="[267,330,872,893]" italics="true" pageId="15" pageNumber="16">T. rex</emphasis>
|
||
</taxonomicName>
|
||
predatory attempts have been identified among the smaller, more common hadrosaurs (
|
||
<bibRefCitation id="FD104B32FFA8FF90FD55FC0FFEACFC43" author="DePalma RA & Burnham DA & Martin LD & Rothschild BM & Larson PL" journalOrPublisher="Proc Natl Acad Sci" pageId="15" pageNumber="16" pagination="12560 - 12564" part="110" refId="ref8608" refString="DePalma RA, Burnham DA, Martin LD, Rothschild BM, Larson PL. 2013. Physical evidence of predatory behavior in Tyrannosaurus rex. Proc Natl Acad Sci 110: 12560 - 12564." title="Physical evidence of predatory behavior in Tyrannosaurus rex" type="journal article" year="2013">DePalma et al., 2013</bibRefCitation>
|
||
) and purportedly ceratopsians (
|
||
<bibRefCitation id="FD104B32FFA8FF90FD89FC10FD5FFC43" author="Happ J." box="[635,766,925,947]" journalOrPublisher="Bloomington, IN: Indiana University Press" pageId="15" pageNumber="16" pagination="355 - 370" refId="ref8840" refString="Happ J. 2008. An analysis of predator- prey behavior in a head-tohead encounter between Tyrannosaurus rex and Triceratops. Tyrannosaurus rex the Tyrant King. Bloomington, IN: Indiana University Press. p 355 - 370." title="An analysis of predator- prey behavior in a head-tohead encounter between Tyrannosaurus rex and Triceratops." type="book chapter" volumeTitle="Tyrannosaurus rex the Tyrant King" year="2008">Happ, 2008</bibRefCitation>
|
||
). Tyrannosaurid giantism therefore appears to have arisen without similar mitigating factors. As such,
|
||
<taxonomicName id="5E814D40FFA8FF90FDAAFC5EFD35FC18" authorityName="Osborn" authorityYear="1905" box="[600,660,979,1000]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA8FF90FDAAFC5EFD35FC18" box="[600,660,979,1000]" italics="true" pageId="15" pageNumber="16">T. rex</emphasis>
|
||
</taxonomicName>
|
||
represents the greatest size yet known at which terrestrial predation remained viable.
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA8FF90FE99FBECFD88FB89" blockId="15.[137,778,1121,1531]" box="[363,553,1121,1145]" pageId="15" pageNumber="16">
|
||
<heading id="C27681AFFFA8FF90FE99FBECFD88FB89" allCaps="true" bold="true" box="[363,553,1121,1145]" centered="true" fontSize="10" level="1" pageId="15" pageNumber="16" reason="0">
|
||
<emphasis id="ABF5EAD1FFA8FF90FE99FBECFD88FB89" bold="true" box="[363,553,1121,1145]" pageId="15" pageNumber="16">CONCLUSION</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="993E36C3FFA8FF90FF56FB06FEFFFA0B" blockId="15.[137,778,1121,1531]" pageId="15" pageNumber="16">
|
||
<taxonomicName id="5E814D40FFA8FF90FF56FB06FEDAFB50" authorityName="Osborn" authorityYear="1905" box="[164,379,1163,1184]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA8FF90FF56FB06FEDAFB50" box="[164,379,1163,1184]" italics="true" pageId="15" pageNumber="16">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
is arguably the most extensively studied and consequently best understood nonavian dinosaur (
|
||
<bibRefCitation id="FD104B32FFA8FF90FF39FB4DFE19FB26" author="Brusatte SL & Norell MA & Carr TD & Erickson GM & Hutchinson JR & Balanoff AM & Bever GS & Choiniere JN & Makovicky PJ & Xu X." box="[203,440,1216,1238]" journalOrPublisher="Science" pageId="15" pageNumber="16" pagination="1481 - 1485" part="329" refId="ref8254" refString="Brusatte SL, Norell MA, Carr TD, Erickson GM, Hutchinson JR, Balanoff AM, Bever GS, Choiniere JN, Makovicky PJ, Xu X. 2010. Tyrannosaur paleobiology: new research on ancient exemplar organisms. Science 329: 1481 - 1485." title="Tyrannosaur paleobiology: new research on ancient exemplar organisms" type="journal article" year="2010">Brusatte et al., 2010</bibRefCitation>
|
||
). Nevertheless, questions pertaining to its ontogeny and the biomechanical challenges of its great size remain pervasive.
|
||
<materialsCitation id="29E93C9EFFA8FF90FDD2FB78FD1AFAFB" ID-GBIF-Occurrence="2597591341" box="[544,699,1269,1291]" collectionCode="RSM" pageId="15" pageNumber="16" specimenCode="RSM P2523.8">RSM P2523.8</materialsCitation>
|
||
shows the form of an elder
|
||
<taxonomicName id="5E814D40FFA8FF90FE98FA9CFE09FAD6" authorityName="Osborn" authorityYear="1905" box="[362,424,1297,1318]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA8FF90FE98FA9CFE09FAD6" box="[362,424,1297,1318]" italics="true" pageId="15" pageNumber="16">T. rex</emphasis>
|
||
</taxonomicName>
|
||
at the terminal extreme of both growth and maturation. Despite its formidable size,
|
||
<taxonomicName id="5E814D40FFA8FF90FF7BFACBFF6CFAAB" authorityName="Osborn" authorityYear="1905" box="[137,205,1350,1371]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA8FF90FF7BFACBFF6CFAAB" box="[137,205,1350,1371]" italics="true" pageId="15" pageNumber="16">T. rex</emphasis>
|
||
</taxonomicName>
|
||
was evidently not invulnerable, and the relative scarcity of other equally large and mature
|
||
<taxonomicName id="5E814D40FFA8FF90FD8FFAECFD61FA86" authorityName="Osborn" authorityYear="1905" box="[637,704,1377,1398]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF,CoL" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="ABF5EAD1FFA8FF90FD8FFAECFD61FA86" box="[637,704,1377,1398]" italics="true" pageId="15" pageNumber="16">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens suggests a high rate of mortality within the species below an 8-tonne threshold. Similarly, the various skeletal pathologies of
|
||
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record the numerous severe injuries suffered over the course of its violent and unusually long life.
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</paragraph>
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</subSubSection>
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</treatment>
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</document> |