638 lines
105 KiB
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638 lines
105 KiB
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<mods:title id="C64CA6880CA68E3A87A67EE413EA1DC4">Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs</mods:title>
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<mods:namePart id="F5A27CECB8D2A5B097D8CBEC611B9188">Gregory M. Erickson</mods:namePart>
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<mods:namePart id="5C54445999E65452B10493DE23C4916D">Peter J. Makovicky</mods:namePart>
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<mods:namePart id="30C3D356BC4BA86FF8C8B480DF4EAD03">Philip J. Currie</mods:namePart>
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<mods:namePart id="EFEABDF192DC29BFB02C98DADF1690CC">Mark A. Norell</mods:namePart>
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<mods:namePart id="B183E7A02027831D6067C739794B5915">Scott A. Yerby</mods:namePart>
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<mods:namePart id="D938AE24449FED3134AC9836C6C7B5C1">Christopher A. Brochu</mods:namePart>
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<mods:title id="A4C72D68B1DDDA2700F596B24FE65F1E">Nature</mods:title>
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<mods:date id="7F154574595A1F9ACE41B5F6AD4E574B">2004</mods:date>
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<mods:number id="172E1C6702D8626275C908DA2E665BBA">2016-03-24</mods:number>
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<mods:number id="8B3CC950DE57A2A32D176D3B15FFE2D4">430</mods:number>
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<treatment id="038987C6350EFFB6FCDED91CFE419F37" ID-DOI="http://doi.org/10.5281/zenodo.3811869" ID-GBIF-Taxon="190305014" ID-Zenodo-Dep="3811869" LSID="urn:lsid:plazi:treatment:038987C6350EFFB6FCDED91CFE419F37" httpUri="http://treatment.plazi.org/id/038987C6350EFFB6FCDED91CFE419F37" lastPageId="3" lastPageNumber="774" pageId="0" pageNumber="772">
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Stemming from more than a century of investigation, considerable understanding of tyrannosaurid osteology
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<bibRefCitation id="EFB14B21350EFFB5FB09D935FB319A35" author="Brochu, C. A." box="[1263,1271,1649,1662]" journalOrPublisher="J. Vert. Paleontol. Mem." pageId="0" pageNumber="772" pagination="1 - 138" part="22" refId="ref3757" refString="4. Brochu, C. A. Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and highresolution computed tomographic analysis of the skull. J. Vert. Paleontol. Mem. 22 (7), 1 - 138 (2003)." title="Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and highresolution computed tomographic analysis of the skull" type="journal article" year="2003">
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, myology
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<bibRefCitation id="EFB14B21350EFFB5FABED935FAA69A35" author="Carrano, M. T. & Hutchinson, J. R." box="[1368,1376,1649,1662]" journalOrPublisher="J. Morphol." pageId="0" pageNumber="772" pagination="207 - 228" part="253" refId="ref3805" refString="5. Carrano, M. T. & Hutchinson, J. R. Pelvic and hindlimb musculature of Tyrannosaurus rex (Dinosauria: Theropoda). J. Morphol. 253, 207 - 228 (2002)." title="Pelvic and hindlimb musculature of Tyrannosaurus rex (Dinosauria: Theropoda)" type="journal article" year="2002">
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<superScript id="7C559B98350EFFB5FABED935FAA69A35" attach="right" box="[1368,1376,1649,1662]" fontSize="5" pageId="0" pageNumber="772">5</superScript>
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, neurology
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<bibRefCitation id="EFB14B21350EFFB5FCA3D9C9FC8B9AD1" author="Brochu, C. A." box="[837,845,1677,1690]" journalOrPublisher="J. Vert. Paleontol." pageId="0" pageNumber="772" pagination="1 - 6" part="20" refId="ref3846" refString="6. Brochu, C. A. A digitally rendered endocast for Tyrannosaurus rex. J. Vert. Paleontol. 20, 1 - 6 (2003)." title="A digitally rendered endocast for Tyrannosaurus rex" type="journal article" year="2003">
|
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<superScript id="7C559B98350EFFB5FCA3D9C9FC8B9AD1" attach="left" box="[837,845,1677,1690]" fontSize="5" pageId="0" pageNumber="772">6</superScript>
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, behaviour
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<superScript id="7C559B98350EFFB5FC2FD9C9FC1B9AD1" attach="left" box="[969,989,1677,1690]" fontSize="5" pageId="0" pageNumber="772">
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<bibRefCitation id="EFB14B21350EFFB5FC2FD9C9FC149AD1" author="Currie, P. J." box="[969,978,1677,1690]" journalOrPublisher="GAIA" pageId="0" pageNumber="772" pagination="271 - 277" part="15" refId="ref3877" refString="7. Currie, P. J. Possible evidence of gregarious behavior in tyrannosaurids. GAIA 15, 271 - 277 (1998)." title="Possible evidence of gregarious behavior in tyrannosaurids" type="journal article" year="1998">7</bibRefCitation>
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,
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<bibRefCitation id="EFB14B21350EFFB5FC33D9C9FC1B9AD1" author="Erickson, G. M." box="[981,989,1677,1690]" journalOrPublisher="Scient. Am." pageId="0" pageNumber="772" pagination="42 - 49" part="9" refId="ref3903" refString="8. Erickson, G. M. Breathing life into Tyrannosaurus rex. Scient. Am. 9, 42 - 49 (1999)." title="Breathing life into Tyrannosaurus rex" type="journal article" year="1999">8</bibRefCitation>
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, physiology
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<superScript id="7C559B98350EFFB5FB87D9C9FBB39AD1" attach="right" box="[1121,1141,1677,1690]" fontSize="5" pageId="0" pageNumber="772">
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<bibRefCitation id="EFB14B21350EFFB5FB87D9C9FBAF9AD1" author="Erickson, G. M. & Curry-Rogers, K. & Yerby, S." box="[1121,1129,1677,1690]" journalOrPublisher="Nature" pageId="0" pageNumber="772" pagination="429 - 433" part="412" refId="ref3720" refString="3. Erickson, G. M., Curry-Rogers, K. & Yerby, S. Dinosaur growth patterns and rapid avian growth rates. Nature 412, 429 - 433 (2001)." title="Dinosaur growth patterns and rapid avian growth rates" type="journal article" year="2001">3</bibRefCitation>
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,
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<bibRefCitation id="EFB14B21350EFFB5FB8AD9C9FBB39AD1" author="Chin, K. & Tokaryk, T. T. & Erickson, G. M. & Calk, L. C." box="[1132,1141,1677,1690]" journalOrPublisher="Nature" pageId="0" pageNumber="772" pagination="680 - 682" part="393" refId="ref3930" refString="9. Chin, K., Tokaryk, T. T., Erickson, G. M. & Calk, L. C. A king-sized theropod coprolite. Nature 393, 680 - 682 (1998)." title="A king-sized theropod coprolite" type="journal article" year="1998">9</bibRefCitation>
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, physical capabilities
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<superScript id="7C559B98350EFFB5FAB8D9C9FA449AD1" attach="left" box="[1374,1410,1677,1690]" fontSize="5" pageId="0" pageNumber="772">
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<bibRefCitation id="EFB14B21350EFFB5FAB8D9C9FAB69AD1" author="Erickson, G. M." box="[1374,1392,1677,1690]" journalOrPublisher="Nature" pageId="0" pageNumber="772" pagination="706 - 708" part="382" refId="ref3972" refString="10. Erickson, G. M. et al. Bite-force estimation for Tyrannosaurus rex from tooth-marked bones. Nature 382, 706 - 708 (1996)." title="Bite-force estimation for Tyrannosaurus rex from tooth-marked bones" type="journal article" year="1996">10</bibRefCitation>
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,
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<bibRefCitation id="EFB14B21350EFFB5FA94D9C9FA449AD1" author="Hutchinson, J. & Garcia, M." box="[1394,1410,1677,1690]" journalOrPublisher="Nature" pageId="0" pageNumber="772" pagination="1018 - 1021" part="415" refId="ref4002" refString="11. Hutchinson, J. & Garcia, M. Tyrannosaurus was not a fast runner. Nature 415, 1018 - 1021 (2002)." title="Tyrannosaurus was not a fast runner" type="journal article" year="2002">11</bibRefCitation>
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and phylogeny
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<bibRefCitation id="EFB14B21350EFFB5FC63D9EDFC539AFD" author="Holtz, T. R." box="[901,917,1705,1718]" editor="Tanke, D. & Carpenter, K." journalOrPublisher="Indiana Univ. Press, Bloomington" pageId="0" pageNumber="772" pagination="64 - 83" refId="ref4030" refString="12. Holtz, T. R. in Mesozoic Vertebrate Life (eds Tanke, D. & Carpenter, K.) 64 - 83 (Indiana Univ. Press, Bloomington, 2001)." type="book chapter" volumeTitle="Mesozoic Vertebrate Life" year="2001">12</bibRefCitation>
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,
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<bibRefCitation id="EFB14B21350EFFB5FC7ED9EDFC619AFD" author="Currie, P. J." box="[920,935,1705,1718]" journalOrPublisher="Acta Palaeontol. Pol." pageId="0" pageNumber="772" pagination="191 - 226" part="48" refId="ref4068" refString="13. Currie, P. J. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontol. Pol. 48, 191 - 226 (2003)." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">13</bibRefCitation>
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have been gained. Lacking are empirical data on tyrannosaurid life history such as growth rates, longevity and somatic maturity (adult size) from which the developmental possibilities for how
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<taxonomicName id="4C204D53350EFFB5FC2AD844FBC59B5E" authority="Osborn, 1905" box="[972,1027,1792,1813]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="772" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B954EAC2350EFFB5FC2AD844FBC59B5E" box="[972,1027,1792,1813]" italics="true" pageId="0" pageNumber="772">T. rex</emphasis>
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attained gigantism can be formally tested.
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Recent advances in techniques for determining the ages at death of dinosaurs by using skeletal growth line counts
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<bibRefCitation id="EFB14B21350EFFB5FAECD871FAD79B09" author="Erickson, G. M. & Curry-Rogers, K. & Yerby, S." box="[1290,1297,1845,1858]" journalOrPublisher="Nature" pageId="0" pageNumber="772" pagination="429 - 433" part="412" refId="ref3720" refString="3. Erickson, G. M., Curry-Rogers, K. & Yerby, S. Dinosaur growth patterns and rapid avian growth rates. Nature 412, 429 - 433 (2001)." title="Dinosaur growth patterns and rapid avian growth rates" type="journal article" year="2001">3</bibRefCitation>
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,
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<bibRefCitation id="EFB14B21350EFFB5FAF3D871FAE39B09" author="Chinsamy, A." box="[1301,1317,1845,1858]" journalOrPublisher="Palaeontol. Afr." pageId="0" pageNumber="772" pagination="77 - 82" part="27" refId="ref4104" refString="14. Chinsamy, A. Physiological implications of the bone histology of Syntarsus rhodesiensis (Saurischia: Theropoda). Palaeontol. Afr. 27, 77 - 82 (1993)." title="Physiological implications of the bone histology of Syntarsus rhodesiensis (Saurischia: Theropoda)" type="journal article" year="1993">14</bibRefCitation>
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, coupled with developmental size estimates
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<bibRefCitation id="EFB14B21350EFFB5FBABD815FB939B15" author="Erickson, G. M. & Curry-Rogers, K. & Yerby, S." box="[1101,1109,1873,1886]" journalOrPublisher="Nature" pageId="0" pageNumber="772" pagination="429 - 433" part="412" refId="ref3720" refString="3. Erickson, G. M., Curry-Rogers, K. & Yerby, S. Dinosaur growth patterns and rapid avian growth rates. Nature 412, 429 - 433 (2001)." title="Dinosaur growth patterns and rapid avian growth rates" type="journal article" year="2001">
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, make quantitative growth-curve reconstructions for dinosaurs feasible. These methods have been used to study growth rates in two small theropods, a small and a large ornithischian and a medium-sized and a gigantic sauropodomorph
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<bibRefCitation id="EFB14B21350EFFB5FC82D884FCAA9B86" author="Erickson, G. M. & Curry-Rogers, K. & Yerby, S." box="[868,876,1984,1997]" journalOrPublisher="Nature" pageId="0" pageNumber="772" pagination="429 - 433" part="412" refId="ref3720" refString="3. Erickson, G. M., Curry-Rogers, K. & Yerby, S. Dinosaur growth patterns and rapid avian growth rates. Nature 412, 429 - 433 (2001)." title="Dinosaur growth patterns and rapid avian growth rates" type="journal article" year="2001">
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. These data were used to derive a regression of body mass against growth rate and to generalize broadly about non-avian dinosaur growth
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<bibRefCitation id="EFB14B21350FFFB4FEFCDFEEFEE79CFC" author="Erickson, G. M. & Curry-Rogers, K. & Yerby, S." box="[282,289,170,183]" journalOrPublisher="Nature" pageId="1" pageNumber="773" pagination="429 - 433" part="412" refId="ref3720" refString="3. Erickson, G. M., Curry-Rogers, K. & Yerby, S. Dinosaur growth patterns and rapid avian growth rates. Nature 412, 429 - 433 (2001)." title="Dinosaur growth patterns and rapid avian growth rates" type="journal article" year="2001">
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. However, because of the phylogenetically disparate nature of these data (that is, none are close outgroups to one another) it has not been possible to use them to infer how specific cases of size change occurred within dinosaurian sub-clades such as the
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<taxonomicName id="4C204D53350FFFB4FD29DE45FF2E9D78" authorityName="Osborn" authorityYear="1906" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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. Such an understanding requires multi-species sampling at low taxonomic levels (that is, among closely related species) and access to growth series spanning juvenile through adult stages, a rarity among extinct dinosaurs
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<bibRefCitation id="EFB14B21350FFFB4FE88DE2AFEBB9D30" author="Weishampel, D. B. & Horner, J. R." box="[366,381,366,379]" editor="Carpenter, K. & Hirsch, K. F. & Horner, J. R." journalOrPublisher="Cambridge Univ. Press, Cambridge" pageId="1" pageNumber="773" pagination="229 - 243" refId="ref4138" refString="15. Weishampel, D. B. & Horner, J. R. in Dinosaur Eggs and Babies (eds Carpenter, K., Hirsch, K. F. & Horner, J. R.) 229 - 243 (Cambridge Univ. Press, Cambridge, 1994)." type="book chapter" volumeTitle="Dinosaur Eggs and Babies" year="1994">
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. Furthermore, it requires the capacity to account for growth line losses due to medullar cavity hollowing and cortical remodelling
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<bibRefCitation id="EFB14B21350FFFB4FED8DEE1FE8B9DF9" author="Francillon-Viellot, H." box="[318,333,421,434]" editor="Carter, J. G." journalOrPublisher="Van Nostrand Reinhold, New York" pageId="1" pageNumber="773" pagination="471 - 530" refId="ref4193" refString="16. Francillon-Viellot, H., et al. in Biomineralization: Patterns and Evolutionary Trends (ed. Carter, J. G.) 471 - 530 (Van Nostrand Reinhold, New York, 1990)." type="book chapter" volumeTitle="Biomineralization: Patterns and Evolutionary Trends" year="1990">
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|
||
, two processes that are pervasive in the major weight-bearing bones from large theropods such as tyrannosaurids.
|
||
</paragraph>
|
||
<paragraph id="8B9F36D0350FFFB4FF75DE9BFED09F9B" blockId="1.[122,784,170,2037]" pageId="1" pageNumber="773">
|
||
The sampling problem has been overcome in North American tyrannosaurids. A flurry of recent discoveries has greatly increased the number of substantially complete specimens representing various growth stages available for study. For example, more than 30
|
||
<taxonomicName id="4C204D53350FFFB4FF7EDD14FF099E2E" authority="Osborn, 1905" box="[152,207,592,613]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350FFFB4FF7EDD14FF099E2E" box="[152,207,592,613]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
||
</taxonomicName>
|
||
specimens are known
|
||
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<bibRefCitation id="EFB14B21350FFFB4FE4FDD09FE779E11" author="Brochu, C. A." box="[425,433,589,602]" journalOrPublisher="J. Vert. Paleontol. Mem." pageId="1" pageNumber="773" pagination="1 - 138" part="22" refId="ref3757" refString="4. Brochu, C. A. Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and highresolution computed tomographic analysis of the skull. J. Vert. Paleontol. Mem. 22 (7), 1 - 138 (2003)." title="Osteology of Tyrannosaurus rex: Insights from a nearly complete skeleton and highresolution computed tomographic analysis of the skull" type="journal article" year="2003">4</bibRefCitation>
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,
|
||
<bibRefCitation id="EFB14B21350FFFB4FE52DD09FE059E11" author="Larson, P. & Donnan, K." box="[436,451,589,602]" journalOrPublisher="Invisible Cities Press, Montpelier" pageId="1" pageNumber="773" refId="ref4234" refString="17. Larson, P. & Donnan, K. Rex Appeal (Invisible Cities Press, Montpelier, 2002)." title="Rex Appeal" type="book" year="2002">17</bibRefCitation>
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||
, compared with only 11 reported in 1993 (ref. 18; see Supplementary Information). Recent work has broadened the developmental representation of these animals by showing that several purported ‘dwarf’ tyrannosaur species are juveniles of larger, previously recognized forms such as
|
||
<taxonomicName id="4C204D53350FFFB4FF9CDD98FF769EBA" authority="Osborn, 1905" box="[122,176,732,753]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
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||
<emphasis id="B954EAC2350FFFB4FF9CDD98FF769EBA" box="[122,176,732,753]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
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</taxonomicName>
|
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<bibRefCitation id="EFB14B21350FFFB4FF49DD9CFF799EAE" author="Holtz, T. R." box="[175,191,728,741]" editor="Tanke, D. & Carpenter, K." journalOrPublisher="Indiana Univ. Press, Bloomington" pageId="1" pageNumber="773" pagination="64 - 83" refId="ref4030" refString="12. Holtz, T. R. in Mesozoic Vertebrate Life (eds Tanke, D. & Carpenter, K.) 64 - 83 (Indiana Univ. Press, Bloomington, 2001)." type="book chapter" volumeTitle="Mesozoic Vertebrate Life" year="2001">12</bibRefCitation>
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,
|
||
<bibRefCitation id="EFB14B21350FFFB4FF24DD9CFF179EAE" author="Currie, P. J." box="[194,209,728,741]" journalOrPublisher="Acta Palaeontol. Pol." pageId="1" pageNumber="773" pagination="191 - 226" part="48" refId="ref4068" refString="13. Currie, P. J. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontol. Pol. 48, 191 - 226 (2003)." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">13</bibRefCitation>
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||
,
|
||
<bibRefCitation id="EFB14B21350FFFB4FF35DD9CFF259EAE" author="Currie, P. J. & Hurum, J. H. & Sabath, K." box="[211,227,728,741]" journalOrPublisher="Acta Palaeontol. Pol." pageId="1" pageNumber="773" pagination="227 - 234" part="48" refId="ref4286" refString="19. Currie, P. J., Hurum, J. H. & Sabath, K. Skull structure and evolution in tyrannosaurid dinosaurs. Acta Palaeontol. Pol. 48, 227 - 234 (2003)." title="Skull structure and evolution in tyrannosaurid dinosaurs" type="journal article" year="2003">19</bibRefCitation>
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,
|
||
<bibRefCitation id="EFB14B21350FFFB4FF00DD9CFF319EAE" author="Carr, T. D." box="[230,247,728,741]" journalOrPublisher="J. Vert. Paleontol." pageId="1" pageNumber="773" pagination="497 - 520" part="19" refId="ref4328" refString="20. Carr, T. D. Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria). J. Vert. Paleontol. 19, 497 - 520 (1999)." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="journal article" year="1999">20</bibRefCitation>
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</superScript>
|
||
. Finally, preliminary analyses for this research revealed that several non-weight-bearing bones in tyrannosaurids (for example pubes, fibulae, ribs, gastralia and postorbitals) did not develop hollow medullar cavities and showed negligible intracortical remodelling during their entire life history (
|
||
<figureCitation id="131B2A55350FFFB4FDBEDC0EFD569F2A" box="[600,656,842,865]" captionStart="Figure 1" captionStartId="1.[816,869,1765,1785]" captionTargetBox="[934,1357,755,1730]" captionTargetId="figure@1.[921,1370,742,1742]" captionTargetPageId="1" captionText="Figure 1 Growth-line counts in tyrannosaurids and reptiles of known ages. a, Thinsectioned Gorgosaurus fibula (FMNH PR 2211). The growth record in this element is complete and shows five growth lines (arrows),indicating that it died early in its sixth year of life. b, Haematoxylin-stained fibula from an 8-year-old alligator (Alligator mississippiensis) showing the expected seven growth lines. Inset, gastralia (unstained) from 9-year-old A. mississippiensis showing the expected eight growth lines and the chemical label used to verify the periodicity of ring formation.c, Tyrannosaurus rib (FMNH PR 2081) showing the 15th to 19th growth lines. Inset, external fundamental system16 with nine tightly spaced growth lines, indicating late-adulthood senescence and growthrate attenuation." figureDoi="http://doi.org/10.5281/zenodo.3736477" httpUri="https://zenodo.org/record/3736477/files/figure.png" pageId="1" pageNumber="773">Fig. 1</figureCitation>
|
||
). Like major long bones, these elements are effective for assessing longevity in living reptiles (
|
||
<figureCitation id="131B2A55350FFFB4FEF3DCC6FE979FD3" box="[277,337,898,921]" captionStart="Figure 1" captionStartId="1.[816,869,1765,1785]" captionTargetBox="[934,1357,755,1730]" captionTargetId="figure@1.[921,1370,742,1742]" captionTargetPageId="1" captionText="Figure 1 Growth-line counts in tyrannosaurids and reptiles of known ages. a, Thinsectioned Gorgosaurus fibula (FMNH PR 2211). The growth record in this element is complete and shows five growth lines (arrows),indicating that it died early in its sixth year of life. b, Haematoxylin-stained fibula from an 8-year-old alligator (Alligator mississippiensis) showing the expected seven growth lines. Inset, gastralia (unstained) from 9-year-old A. mississippiensis showing the expected eight growth lines and the chemical label used to verify the periodicity of ring formation.c, Tyrannosaurus rib (FMNH PR 2081) showing the 15th to 19th growth lines. Inset, external fundamental system16 with nine tightly spaced growth lines, indicating late-adulthood senescence and growthrate attenuation." figureDoi="http://doi.org/10.5281/zenodo.3736477" httpUri="https://zenodo.org/record/3736477/files/figure.png" pageId="1" pageNumber="773">Fig. 1</figureCitation>
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)
|
||
<superScript id="7C559B98350FFFB4FEBDDCC4FEBA9FC6" attach="left" box="[347,380,896,909]" fontSize="5" pageId="1" pageNumber="773">
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<bibRefCitation id="EFB14B21350FFFB4FEBDDCC4FEAC9FC6" author="Castanet, J. & Naulleau, G." box="[347,362,896,909]" journalOrPublisher="Zool. Scripta" pageId="1" pageNumber="773" pagination="201 - 208" part="3" refId="ref4361" refString="21. Castanet, J. & Naulleau, G. Donnees experimentales sur la valeur des marques squelettiques comme indicateur de l' age chez Vipera aspis (L.) (Ophidia, Viperidae). Zool. Scripta 3, 201 - 208 (1974)." title="Donnees experimentales sur la valeur des marques squelettiques comme indicateur de l' age chez Vipera aspis (L.) (Ophidia, Viperidae)" type="journal article" year="1974">21</bibRefCitation>
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,
|
||
<bibRefCitation id="EFB14B21350FFFB4FE8ADCC4FEBA9FC6" author="de Buffrenil, V. & Castanet, J." box="[364,380,896,909]" journalOrPublisher="J. Herpetol." pageId="1" pageNumber="773" pagination="414 - 424" part="34" refId="ref4410" refString="22. de Buffrenil, V. & Castanet, J. Age estimation by skeletochronology in the Nile monitor (Varanus niloticus), a highly exploited species. J. Herpetol. 34, 414 - 424 (2000)." title="Age estimation by skeletochronology in the Nile monitor (Varanus niloticus), a highly exploited species" type="journal article" year="2000">22</bibRefCitation>
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||
</superScript>
|
||
and hence provide a viable alternative method for determining the age at death of extinct reptiles such as tyrannosaurids.
|
||
</paragraph>
|
||
<paragraph id="8B9F36D0350FFFB4FF75DC92FDC099B6" blockId="1.[122,784,170,2037]" pageId="1" pageNumber="773">
|
||
Here we exploit these findings to determine the pattern of growth in
|
||
<taxonomicName id="4C204D53350FFFB4FF73DCB6FF0A984C" authority="Osborn, 1905" box="[149,204,1010,1031]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350FFFB4FF73DCB6FF0A984C" box="[149,204,1010,1031]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
||
</taxonomicName>
|
||
and three of its close tyrannosaurid relatives. We then use character optimization methods
|
||
<bibRefCitation id="EFB14B21350FFFB4FE5EDB4FFE019853" author="Maddison, W. P. & Maddison, D. R." box="[440,455,1035,1048]" journalOrPublisher="Sinauer Associates, Sunderland, MA" pageId="1" pageNumber="773" refId="ref4453" refString="23. Maddison, W. P. & Maddison, D. R. MacClade Version 3; Analysis of Phylogeny and Character Evolution (Sinauer Associates, Sunderland, MA, 1992)." title="MacClade Version 3; Analysis of Phylogeny and Character Evolution" type="book" year="1992">
|
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<superScript id="7C559B98350FFFB4FE5EDB4FFE019853" attach="left" box="[440,455,1035,1048]" fontSize="5" pageId="1" pageNumber="773">23</superScript>
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||
</bibRefCitation>
|
||
to infer how
|
||
<taxonomicName id="4C204D53350FFFB4FDAADB4AFD449868" authority="Osborn, 1905" box="[588,642,1038,1059]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350FFFB4FDAADB4AFD449868" box="[588,642,1038,1059]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
||
</taxonomicName>
|
||
attained giant proportions among tyrannosaurids. Finally, this new evidence is used to further our understanding of tyrannosaurid biology. In performing these analyses, we sampled several amedullar bones from adolescent, juvenile, sub-adult and adult representatives of the North American Late Cretaceous tyrannosaurids
|
||
<taxonomicName id="4C204D53350FFFB4FD6FDBDDFF329881" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="sarcophagus">
|
||
<emphasis id="B954EAC2350FFFB4FD6FDBDDFF329881" italics="true" pageId="1" pageNumber="773">Albertosaurus sarcophagus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C204D53350FFFB4FEE0DBF1FE199881" baseAuthorityName="Lambe" baseAuthorityYear="1914" box="[262,479,1205,1226]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis id="B954EAC2350FFFB4FEE0DBF1FE199881" box="[262,479,1205,1226]" italics="true" pageId="1" pageNumber="773">Gorgosaurus libratus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C204D53350FFFB4FE17DBF1FD1B9880" baseAuthorityName="Russell" baseAuthorityYear="1970" box="[497,733,1205,1227]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis id="B954EAC2350FFFB4FE17DBF1FD1B9880" box="[497,733,1205,1227]" italics="true" pageId="1" pageNumber="773">Daspletosaurus torosus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C204D53350FFFB4FF9CDB96FF7598AC" authority="Osborn, 1905" box="[122,179,1234,1255]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350FFFB4FF9CDB96FF7598AC" box="[122,179,1234,1255]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
||
</taxonomicName>
|
||
. Longevity in each of the 20 specimens was assessed from line counts in histological sections by using polarizing, dissecting and reflected-light microscopy (
|
||
<figureCitation id="131B2A55350FFFB4FE57DA4CFE2E9954" box="[433,488,1288,1311]" captionStart="Figure 1" captionStartId="1.[816,869,1765,1785]" captionTargetBox="[934,1357,755,1730]" captionTargetId="figure@1.[921,1370,742,1742]" captionTargetPageId="1" captionText="Figure 1 Growth-line counts in tyrannosaurids and reptiles of known ages. a, Thinsectioned Gorgosaurus fibula (FMNH PR 2211). The growth record in this element is complete and shows five growth lines (arrows),indicating that it died early in its sixth year of life. b, Haematoxylin-stained fibula from an 8-year-old alligator (Alligator mississippiensis) showing the expected seven growth lines. Inset, gastralia (unstained) from 9-year-old A. mississippiensis showing the expected eight growth lines and the chemical label used to verify the periodicity of ring formation.c, Tyrannosaurus rib (FMNH PR 2081) showing the 15th to 19th growth lines. Inset, external fundamental system16 with nine tightly spaced growth lines, indicating late-adulthood senescence and growthrate attenuation." figureDoi="http://doi.org/10.5281/zenodo.3736477" httpUri="https://zenodo.org/record/3736477/files/figure.png" pageId="1" pageNumber="773">Fig. 1</figureCitation>
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||
)
|
||
<superScript id="7C559B98350FFFB4FE14DA42FDCA9958" attach="none" box="[498,524,1286,1299]" fontSize="5" pageId="1" pageNumber="773">
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<bibRefCitation id="EFB14B21350FFFB4FE14DA42FE3F9958" author="Erickson, G. M. & Curry-Rogers, K. & Yerby, S." box="[498,505,1286,1299]" journalOrPublisher="Nature" pageId="1" pageNumber="773" pagination="429 - 433" part="412" refId="ref3720" refString="3. Erickson, G. M., Curry-Rogers, K. & Yerby, S. Dinosaur growth patterns and rapid avian growth rates. Nature 412, 429 - 433 (2001)." title="Dinosaur growth patterns and rapid avian growth rates" type="journal article" year="2001">3</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFB14B21350FFFB4FE1ADA42FDCA9958" author="Chinsamy, A." box="[508,524,1286,1299]" journalOrPublisher="Palaeontol. Afr." pageId="1" pageNumber="773" pagination="77 - 82" part="27" refId="ref4104" refString="14. Chinsamy, A. Physiological implications of the bone histology of Syntarsus rhodesiensis (Saurischia: Theropoda). Palaeontol. Afr. 27, 77 - 82 (1993)." title="Physiological implications of the bone histology of Syntarsus rhodesiensis (Saurischia: Theropoda)" type="journal article" year="1993">14</bibRefCitation>
|
||
</superScript>
|
||
. Conservative estimates of body mass (see Supplementary Information) were made by using femoral circumference measures
|
||
<bibRefCitation id="EFB14B21350FFFB4FE25DA7AFE149900" author="Anderson, J. F. & Hall-Martin, A. & Russell, D. A." box="[451,466,1342,1355]" journalOrPublisher="J. Zool. A" pageId="1" pageNumber="773" pagination="53 - 61" part="207" refId="ref4489" refString="24. Anderson, J. F., Hall-Martin, A. & Russell, D. A. Long bone circumference and weight in mammals, birds, and dinosaurs. J. Zool. A 207, 53 - 61 (1985)." title="Long bone circumference and weight in mammals, birds, and dinosaurs" type="journal article" year="1985">
|
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<superScript id="7C559B98350FFFB4FE25DA7AFE149900" attach="left" box="[451,466,1342,1355]" fontSize="5" pageId="1" pageNumber="773">24</superScript>
|
||
</bibRefCitation>
|
||
. Longevity and size data were plotted and least-squares regression was used to determine the first empirical growth curves for tyrannosaurids
|
||
<bibRefCitation id="EFB14B21350FFFB4FDC5DA32FDED99C8" author="Erickson, G. M. & Curry-Rogers, K. & Yerby, S." box="[547,555,1398,1411]" journalOrPublisher="Nature" pageId="1" pageNumber="773" pagination="429 - 433" part="412" refId="ref3720" refString="3. Erickson, G. M., Curry-Rogers, K. & Yerby, S. Dinosaur growth patterns and rapid avian growth rates. Nature 412, 429 - 433 (2001)." title="Dinosaur growth patterns and rapid avian growth rates" type="journal article" year="2001">
|
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<superScript id="7C559B98350FFFB4FDC5DA32FDED99C8" attach="left" box="[547,555,1398,1411]" fontSize="5" pageId="1" pageNumber="773">3</superScript>
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||
</bibRefCitation>
|
||
. The length and timing of the various developmental stages and the maximal growth rates for each taxon were compared
|
||
<bibRefCitation id="EFB14B21350FFFB4FE4DDAEAFE7C99F0" author="Sussman, M." box="[427,442,1454,1467]" journalOrPublisher="Prentice-Hall, Englewood Cliffs, New Jersey" pageId="1" pageNumber="773" refId="ref4536" refString="25. Sussman, M. Growth and Development (Prentice-Hall, Englewood Cliffs, New Jersey, 1964)." title="Growth and Development" type="book" year="1964">
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<superScript id="7C559B98350FFFB4FE4DDAEAFE7C99F0" attach="left" box="[427,442,1454,1467]" fontSize="5" pageId="1" pageNumber="773">25</superScript>
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||
</bibRefCitation>
|
||
. The results were examined in an evolutionary context
|
||
<bibRefCitation id="EFB14B21350FFFB4FEBEDA8DFEAE999D" author="Maddison, W. P. & Maddison, D. R." box="[344,360,1481,1494]" journalOrPublisher="Sinauer Associates, Sunderland, MA" pageId="1" pageNumber="773" refId="ref4453" refString="23. Maddison, W. P. & Maddison, D. R. MacClade Version 3; Analysis of Phylogeny and Character Evolution (Sinauer Associates, Sunderland, MA, 1992)." title="MacClade Version 3; Analysis of Phylogeny and Character Evolution" type="book" year="1992">
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<superScript id="7C559B98350FFFB4FEBEDA8DFEAE999D" attach="left" box="[344,360,1481,1494]" fontSize="5" pageId="1" pageNumber="773">23</superScript>
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||
</bibRefCitation>
|
||
by using two competing phylogenetic hypotheses for the
|
||
<taxonomicName id="4C204D53350FFFB4FEDEDAA3FE1899B5" authorityName="Osborn" authorityYear="1906" box="[312,478,1511,1534]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
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<superScript id="7C559B98350FFFB4FE38DAA1FDC699B9" attach="left" box="[478,512,1509,1522]" fontSize="5" pageId="1" pageNumber="773">
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<bibRefCitation id="EFB14B21350FFFB4FE38DAA1FE2899B9" author="Holtz, T. R." box="[478,494,1509,1522]" editor="Tanke, D. & Carpenter, K." journalOrPublisher="Indiana Univ. Press, Bloomington" pageId="1" pageNumber="773" pagination="64 - 83" refId="ref4030" refString="12. Holtz, T. R. in Mesozoic Vertebrate Life (eds Tanke, D. & Carpenter, K.) 64 - 83 (Indiana Univ. Press, Bloomington, 2001)." type="book chapter" volumeTitle="Mesozoic Vertebrate Life" year="2001">12</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFB14B21350FFFB4FE17DAA1FDC699B9" author="Currie, P. J." box="[497,512,1509,1522]" journalOrPublisher="Acta Palaeontol. Pol." pageId="1" pageNumber="773" pagination="191 - 226" part="48" refId="ref4068" refString="13. Currie, P. J. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontol. Pol. 48, 191 - 226 (2003)." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">13</bibRefCitation>
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</superScript>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="8B9F36D0350FFFB4FF75D947FA629D04" blockId="1.[122,784,170,2037]" lastBlockId="1.[816,1477,172,698]" pageId="1" pageNumber="773">
|
||
Sampled longevities for
|
||
<taxonomicName id="4C204D53350FFFB4FE6BD941FE019A51" authority="Osborn, 1905" box="[397,455,1541,1562]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350FFFB4FE6BD941FE019A51" box="[397,455,1541,1562]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
||
</taxonomicName>
|
||
ranged from 2 to 28 years and corresponding body mass estimates ranged from 29.9 to 5,654 kg (
|
||
<tableCitation id="C6A2036B350FFFB4FF67D97FFF0C9A19" box="[129,202,1595,1618]" captionStart="Table 1" captionStartId="2.[106,147,175,191]" captionTargetBox="[106,1461,206,655]" captionTargetPageId="2" captionText="Table 1 Specimens, size measures, and longevity data" httpUri="http://table.plazi.org/id/DF5F6658350CFFB7FF8CDFEBFDE69CF4" pageId="1" pageNumber="773" tableUuid="DF5F6658350CFFB7FF8CDFEBFDE69CF4">Table 1</tableCitation>
|
||
). The transition to somatic maturity in this taxon seems to have begun at about 18.5 years of age (
|
||
<figureCitation id="131B2A55350FFFB4FE0CD913FDE39A25" box="[490,549,1623,1646]" captionStart="Figure 2" captionStartId="2.[106,159,1931,1951]" captionTargetBox="[338,1229,1274,1906]" captionTargetId="figure@2.[425,1225,1264,1864]" captionTargetPageId="2" captionText="Figure 2 Logistic growth curves for Tyrannosaurus and three related tyrannosaurids.Note that the exponential stages (the regions of maximal slope) are similar in duration but differ in slope (that is, growth rates). Regression equations (mass in kg, age in years) are as follows: T. rex, mass = {5,551/[1 + e‾0.57(age ‾ 16.1)]} + 5, r 2 = 0.953; D. torosus, mass = {1,728/[1 + e‾0.44(age ‾ 12.1)]} + 5, r 2 = 0.992; G. libratus, mass = {1,234/[1 + e‾0.38(age ‾ 12.4)]} + 5, r 2 = 0.950; A. sarcophagus, mass = {1,218/[1 + e‾0.43(age ‾ 14.1)]} + 5; r 2 = 0.985." figureDoi="http://doi.org/10.5281/zenodo.4011699" httpUri="https://zenodo.org/record/4011699/files/figure.png" pageId="1" pageNumber="773">Fig. 2</figureCitation>
|
||
). At least one individual (exemplified by
|
||
<materialsCitation id="3B483C8D350FFFB4FEC4D930FD769AC1" ID-GBIF-Occurrence="3396395307" box="[290,688,1651,1674]" collectionCode="FMNH" pageId="1" pageNumber="773" specimenCode="FMNH PR2081">FMNH (The Field Museum) PR 2081</materialsCitation>
|
||
), showed evidence for prolonged senescence in the form of conspicuously narrow pericortical growth-line spacing (
|
||
<figureCitation id="131B2A55350FFFB4FDFED9EFFD949A8A" box="[536,594,1707,1730]" captionStart="Figure 1" captionStartId="1.[816,869,1765,1785]" captionTargetBox="[934,1357,755,1730]" captionTargetId="figure@1.[921,1370,742,1742]" captionTargetPageId="1" captionText="Figure 1 Growth-line counts in tyrannosaurids and reptiles of known ages. a, Thinsectioned Gorgosaurus fibula (FMNH PR 2211). The growth record in this element is complete and shows five growth lines (arrows),indicating that it died early in its sixth year of life. b, Haematoxylin-stained fibula from an 8-year-old alligator (Alligator mississippiensis) showing the expected seven growth lines. Inset, gastralia (unstained) from 9-year-old A. mississippiensis showing the expected eight growth lines and the chemical label used to verify the periodicity of ring formation.c, Tyrannosaurus rib (FMNH PR 2081) showing the 15th to 19th growth lines. Inset, external fundamental system16 with nine tightly spaced growth lines, indicating late-adulthood senescence and growthrate attenuation." figureDoi="http://doi.org/10.5281/zenodo.3736477" httpUri="https://zenodo.org/record/3736477/files/figure.png" pageId="1" pageNumber="773">Fig. 1</figureCitation>
|
||
). Maximal growth rates in
|
||
<taxonomicName id="4C204D53350FFFB4FF36D98CFECD9A96" authority="Osborn, 1905" box="[208,267,1736,1757]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350FFFB4FF36D98CFECD9A96" box="[208,267,1736,1757]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
||
</taxonomicName>
|
||
were 2.07 kg d
|
||
<superScript id="7C559B98350FFFB4FE44D985FE7D9A9B" attach="left" box="[418,443,1729,1744]" fontSize="6" pageId="1" pageNumber="773">
|
||
<emphasis id="B954EAC2350FFFB4FE44D985FE779A9B" bold="true" box="[418,433,1729,1744]" pageId="1" pageNumber="773">‾</emphasis>
|
||
<bibRefCitation id="EFB14B21350FFFB4FE54D985FE7D9A9B" author="Gould, S. J." box="[434,443,1729,1744]" journalOrPublisher="Harvard Univ. Press, Cambridge, Massachusetts" pageId="1" pageNumber="773" refId="ref3634" refString="1. Gould, S. J. Ontogeny and Phylogeny (Harvard Univ. Press, Cambridge, Massachusetts, 1977)." title="Ontogeny and Phylogeny" type="book" year="1977">1</bibRefCitation>
|
||
</superScript>
|
||
and such exponential rates were maintained for about 4 years (
|
||
<figureCitation id="131B2A55350FFFB4FE4BD9A6FE2D9AB2" box="[429,491,1762,1785]" captionStart="Figure 2" captionStartId="2.[106,159,1931,1951]" captionTargetBox="[338,1229,1274,1906]" captionTargetId="figure@2.[425,1225,1264,1864]" captionTargetPageId="2" captionText="Figure 2 Logistic growth curves for Tyrannosaurus and three related tyrannosaurids.Note that the exponential stages (the regions of maximal slope) are similar in duration but differ in slope (that is, growth rates). Regression equations (mass in kg, age in years) are as follows: T. rex, mass = {5,551/[1 + e‾0.57(age ‾ 16.1)]} + 5, r 2 = 0.953; D. torosus, mass = {1,728/[1 + e‾0.44(age ‾ 12.1)]} + 5, r 2 = 0.992; G. libratus, mass = {1,234/[1 + e‾0.38(age ‾ 12.4)]} + 5, r 2 = 0.950; A. sarcophagus, mass = {1,218/[1 + e‾0.43(age ‾ 14.1)]} + 5; r 2 = 0.985." figureDoi="http://doi.org/10.5281/zenodo.4011699" httpUri="https://zenodo.org/record/4011699/files/figure.png" pageId="1" pageNumber="773">Fig. 2</figureCitation>
|
||
). The longevity estimates for
|
||
<taxonomicName id="4C204D53350FFFB4FF9CD844FF769B5E" authority="Osborn, 1905" box="[122,176,1792,1813]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350FFFB4FF9CD844FF769B5E" box="[122,176,1792,1813]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
||
</taxonomicName>
|
||
outgroups ranged from 2 to 24 years and corresponding body sizes spanned from 50.3 to 1,791 kg (
|
||
<tableCitation id="C6A2036B350FFFB4FE1FD85EFD809B7A" box="[505,582,1818,1841]" captionStart="Table 1" captionStartId="2.[106,147,175,191]" captionTargetBox="[106,1461,206,655]" captionTargetPageId="2" captionText="Table 1 Specimens, size measures, and longevity data" httpUri="http://table.plazi.org/id/DF5F6658350CFFB7FF8CDFEBFDE69CF4" pageId="1" pageNumber="773" tableUuid="DF5F6658350CFFB7FF8CDFEBFDE69CF4">Table 1</tableCitation>
|
||
). Somatic maturity occurred at between 14 and 16 years in these taxa (
|
||
<figureCitation id="131B2A55350FFFB4FDBBD872FD519B06" box="[605,663,1846,1869]" captionStart="Figure 2" captionStartId="2.[106,159,1931,1951]" captionTargetBox="[338,1229,1274,1906]" captionTargetId="figure@2.[425,1225,1264,1864]" captionTargetPageId="2" captionText="Figure 2 Logistic growth curves for Tyrannosaurus and three related tyrannosaurids.Note that the exponential stages (the regions of maximal slope) are similar in duration but differ in slope (that is, growth rates). Regression equations (mass in kg, age in years) are as follows: T. rex, mass = {5,551/[1 + e‾0.57(age ‾ 16.1)]} + 5, r 2 = 0.953; D. torosus, mass = {1,728/[1 + e‾0.44(age ‾ 12.1)]} + 5, r 2 = 0.992; G. libratus, mass = {1,234/[1 + e‾0.38(age ‾ 12.4)]} + 5, r 2 = 0.950; A. sarcophagus, mass = {1,218/[1 + e‾0.43(age ‾ 14.1)]} + 5; r 2 = 0.985." figureDoi="http://doi.org/10.5281/zenodo.4011699" httpUri="https://zenodo.org/record/4011699/files/figure.png" pageId="1" pageNumber="773">Fig. 2</figureCitation>
|
||
). Like
|
||
<taxonomicName id="4C204D53350FFFB4FD30D873FCCC9B07" authority="Osborn, 1905" box="[726,778,1847,1868]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350FFFB4FD30D873FCCC9B07" box="[726,778,1847,1868]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
||
</taxonomicName>
|
||
, at least some exceptionally large individuals of
|
||
<taxonomicName id="4C204D53350FFFB4FDB5D817FD229B23" authorityName="Osborn" authorityYear="1905" box="[595,740,1875,1896]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="sarcophagus">
|
||
<emphasis id="B954EAC2350FFFB4FDB5D817FD229B23" box="[595,740,1875,1896]" italics="true" pageId="1" pageNumber="773">A. sarcophagus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C204D53350FFFB4FF9CD82BFF1A9BCF" baseAuthorityName="Russell" baseAuthorityYear="1970" box="[122,220,1903,1924]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis id="B954EAC2350FFFB4FF9CD82BFF1A9BCF" box="[122,220,1903,1924]" italics="true" pageId="1" pageNumber="773">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
showed narrow pericortical growth-line spacing indicative of the onset of senescence. The maximal growth rates for the three smaller tyrannosaurid taxa ranged from 0.31 to 0.48 kg d
|
||
<superScript id="7C559B98350FFFB4FD17D8E4FCCC9BE4" attach="left" box="[753,778,1952,1967]" fontSize="6" pageId="1" pageNumber="773">
|
||
<emphasis id="B954EAC2350FFFB4FD17D8E4FCC69BE4" bold="true" box="[753,768,1952,1967]" pageId="1" pageNumber="773">‾</emphasis>
|
||
<bibRefCitation id="EFB14B21350FFFB4FCE7D8E4FCCC9BE4" author="Gould, S. J." box="[769,778,1952,1967]" journalOrPublisher="Harvard Univ. Press, Cambridge, Massachusetts" pageId="1" pageNumber="773" refId="ref3634" refString="1. Gould, S. J. Ontogeny and Phylogeny (Harvard Univ. Press, Cambridge, Massachusetts, 1977)." title="Ontogeny and Phylogeny" type="book" year="1977">1</bibRefCitation>
|
||
</superScript>
|
||
; such exponential stage rates were also maintained for about 4 years (
|
||
<figureCitation id="131B2A55350FFFB4FF64D89AFF039BBF" box="[130,197,2014,2037]" captionStart="Figure 2" captionStartId="2.[106,159,1931,1951]" captionTargetBox="[338,1229,1274,1906]" captionTargetId="figure@2.[425,1225,1264,1864]" captionTargetPageId="2" captionText="Figure 2 Logistic growth curves for Tyrannosaurus and three related tyrannosaurids.Note that the exponential stages (the regions of maximal slope) are similar in duration but differ in slope (that is, growth rates). Regression equations (mass in kg, age in years) are as follows: T. rex, mass = {5,551/[1 + e‾0.57(age ‾ 16.1)]} + 5, r 2 = 0.953; D. torosus, mass = {1,728/[1 + e‾0.44(age ‾ 12.1)]} + 5, r 2 = 0.992; G. libratus, mass = {1,234/[1 + e‾0.38(age ‾ 12.4)]} + 5, r 2 = 0.950; A. sarcophagus, mass = {1,218/[1 + e‾0.43(age ‾ 14.1)]} + 5; r 2 = 0.985." figureDoi="http://doi.org/10.5281/zenodo.4011699" httpUri="https://zenodo.org/record/4011699/files/figure.png" pageId="1" pageNumber="773">Fig. 2</figureCitation>
|
||
). Optimization of growth rates onto the two current phylogenetic hypotheses of tyrannosaurid relationships suggests that a 1.5-fold acceleration in maximal growth rate might diagnose
|
||
<taxonomicName id="4C204D53350FFFB4FCD6DFA0FC119CB0" authorityName="Matthew & Brown" authorityYear="1922" box="[816,983,228,251]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="subFamily" subFamily="Tyrannosaurinae">Tyrannosaurinae</taxonomicName>
|
||
(the clade comprising
|
||
<emphasis id="B954EAC2350FFFB4FB58DFA1FC409D5D" italics="true" pageId="1" pageNumber="773">
|
||
<taxonomicName id="4C204D53350FFFB4FB58DFA1FA969CB1" authorityName="Russell" authorityYear="1970" box="[1214,1360,229,250]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="genus">Daspletosaurus</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C204D53350FFFB4FA65DFA1FC409D5D" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||
</emphasis>
|
||
<superScript id="7C559B98350FFFB4FC60DFBAFC6E9D40" attach="left" box="[902,936,254,267]" fontSize="5" pageId="1" pageNumber="773">
|
||
<bibRefCitation id="EFB14B21350FFFB4FC60DFBAFC539D40" author="Currie, P. J." box="[902,917,254,267]" journalOrPublisher="Acta Palaeontol. Pol." pageId="1" pageNumber="773" pagination="191 - 226" part="48" refId="ref4068" refString="13. Currie, P. J. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontol. Pol. 48, 191 - 226 (2003)." title="Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada" type="journal article" year="2003">13</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFB14B21350FFFB4FC7EDFBAFC6E9D40" author="Currie, P. J. & Hurum, J. H. & Sabath, K." box="[920,936,254,267]" journalOrPublisher="Acta Palaeontol. Pol." pageId="1" pageNumber="773" pagination="227 - 234" part="48" refId="ref4286" refString="19. Currie, P. J., Hurum, J. H. & Sabath, K. Skull structure and evolution in tyrannosaurid dinosaurs. Acta Palaeontol. Pol. 48, 227 - 234 (2003)." title="Skull structure and evolution in tyrannosaurid dinosaurs" type="journal article" year="2003">19</bibRefCitation>
|
||
</superScript>
|
||
,
|
||
<figureCitation id="131B2A55350FFFB4FC50DE44FC349D5C" box="[950,1010,256,279]" captionStart="Figure 2" captionStartId="2.[106,159,1931,1951]" captionTargetBox="[338,1229,1274,1906]" captionTargetId="figure@2.[425,1225,1264,1864]" captionTargetPageId="2" captionText="Figure 2 Logistic growth curves for Tyrannosaurus and three related tyrannosaurids.Note that the exponential stages (the regions of maximal slope) are similar in duration but differ in slope (that is, growth rates). Regression equations (mass in kg, age in years) are as follows: T. rex, mass = {5,551/[1 + e‾0.57(age ‾ 16.1)]} + 5, r 2 = 0.953; D. torosus, mass = {1,728/[1 + e‾0.44(age ‾ 12.1)]} + 5, r 2 = 0.992; G. libratus, mass = {1,234/[1 + e‾0.38(age ‾ 12.4)]} + 5, r 2 = 0.950; A. sarcophagus, mass = {1,218/[1 + e‾0.43(age ‾ 14.1)]} + 5; r 2 = 0.985." figureDoi="http://doi.org/10.5281/zenodo.4011699" httpUri="https://zenodo.org/record/4011699/files/figure.png" pageId="1" pageNumber="773">Fig. 2</figureCitation>
|
||
). A second substantial increase in growth rate optimizes as a physiological autapomorphy of
|
||
<taxonomicName id="4C204D53350FFFB4FAE5DE59FA549D79" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1283,1426,285,306]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="genus">
|
||
<emphasis id="B954EAC2350FFFB4FAE5DE59FA549D79" box="[1283,1426,285,306]" italics="true" pageId="1" pageNumber="773">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
irrespective of phylogenetic hypothesis and optimization criterion.
|
||
</paragraph>
|
||
<paragraph id="8B9F36D0350FFFB7FCAEDE11FE5D98F7" blockId="1.[816,1477,172,698]" lastPageId="2" lastPageNumber="774" pageId="1" pageNumber="773">
|
||
<taxonomicName id="4C204D53350FFFB4FCAEDE11FCB99D21" authority="Osborn, 1905" box="[840,895,341,362]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350FFFB4FCAEDE11FCB99D21" box="[840,895,341,362]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
||
</taxonomicName>
|
||
is notable for its great size, which is at least 15-fold greater than the largest living terrestrial carnivorous animals today and second only to
|
||
<taxonomicName id="4C204D53350FFFB4FC2DDEC8FBA79DEA" authorityName="Coria & Salgado" authorityYear="1995" box="[971,1121,396,417]" class="Reptilia" family="Carcharodontosauridae" genus="Giganotosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="genus">
|
||
<emphasis id="B954EAC2350FFFB4FC2DDEC8FBA79DEA" box="[971,1121,396,417]" italics="true" pageId="1" pageNumber="773">Giganotosaurus</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation id="EFB14B21350FFFB4FB87DECDFBB69DDD" author="Coria, R. A. & Salgado, L." box="[1121,1136,393,406]" journalOrPublisher="Nature" pageId="1" pageNumber="773" pagination="224 - 226" part="377" refId="ref4557" refString="26. Coria, R. A. & Salgado, L. A new giant carnivorous dinosaur from the Cretaceous of Patagonia. Nature 377, 224 - 226 (1995)." title="A new giant carnivorous dinosaur from the Cretaceous of Patagonia" type="journal article" year="1995">
|
||
<superScript id="7C559B98350FFFB4FB87DECDFBB69DDD" attach="left" box="[1121,1136,393,406]" fontSize="5" pageId="1" pageNumber="773">26</superScript>
|
||
</bibRefCitation>
|
||
among theropod dinosaurs. How did it attain such great proportions within the
|
||
<taxonomicName id="4C204D53350FFFB4FAF0DEE3FA7A9DF5" authorityName="Osborn" authorityYear="1906" box="[1302,1468,423,446]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
? From the two competing hypotheses of tyrannosaurid phylogeny it is most parsimonious to conclude that
|
||
<taxonomicName id="4C204D53350FFFB4FB55DEA4FB2C9DBE" authority="Osborn, 1905" box="[1203,1258,480,501]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350FFFB4FB55DEA4FB2C9DBE" box="[1203,1258,480,501]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
||
</taxonomicName>
|
||
acquired the majority of its giant proportions after diverging from the common ancestor of itself and
|
||
<taxonomicName id="4C204D53350FFFB4FC49DD5CFBD79E66" baseAuthorityName="Russell" baseAuthorityYear="1970" box="[943,1041,536,557]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis id="B954EAC2350FFFB4FC49DD5CFBD79E66" box="[943,1041,536,557]" italics="true" pageId="1" pageNumber="773">D. torosus</emphasis>
|
||
</taxonomicName>
|
||
, a species with an optimized body mass of about 1,800 kg. Direct comparison between the tyrannosaurid growth curves shows that the transition to the exponential and stationary phases of development occurred about 2–4 years later in
|
||
<taxonomicName id="4C204D53350FFFB4FCD6DDCCFCA39ED6" authority="Osborn, 1905" box="[816,869,648,669]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350FFFB4FCD6DDCCFCA39ED6" box="[816,869,648,669]" italics="true" pageId="1" pageNumber="773">T. rex</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="131B2A55350FFFB4FC97DDC3FC6A9ED6" box="[881,940,647,670]" captionStart="Figure 2" captionStartId="2.[106,159,1931,1951]" captionTargetBox="[338,1229,1274,1906]" captionTargetId="figure@2.[425,1225,1264,1864]" captionTargetPageId="2" captionText="Figure 2 Logistic growth curves for Tyrannosaurus and three related tyrannosaurids.Note that the exponential stages (the regions of maximal slope) are similar in duration but differ in slope (that is, growth rates). Regression equations (mass in kg, age in years) are as follows: T. rex, mass = {5,551/[1 + e‾0.57(age ‾ 16.1)]} + 5, r 2 = 0.953; D. torosus, mass = {1,728/[1 + e‾0.44(age ‾ 12.1)]} + 5, r 2 = 0.992; G. libratus, mass = {1,234/[1 + e‾0.38(age ‾ 12.4)]} + 5, r 2 = 0.950; A. sarcophagus, mass = {1,218/[1 + e‾0.43(age ‾ 14.1)]} + 5; r 2 = 0.985." figureDoi="http://doi.org/10.5281/zenodo.4011699" httpUri="https://zenodo.org/record/4011699/files/figure.png" pageId="1" pageNumber="773">Fig. 2</figureCitation>
|
||
). However, such temporal post-displacement had little to do with the evolution of its gigantism because the exponential stage, during which most body size is accrued
|
||
<bibRefCitation id="EFB14B21350CFFB7FDD3DC45FD829F45" author="Sussman, M." box="[565,580,769,782]" journalOrPublisher="Prentice-Hall, Englewood Cliffs, New Jersey" pageId="2" pageNumber="774" refId="ref4536" refString="25. Sussman, M. Growth and Development (Prentice-Hall, Englewood Cliffs, New Jersey, 1964)." title="Growth and Development" type="book" year="1964">25</bibRefCitation>
|
||
, was not extended beyond the ancestral, 4-year condition observed in other tyrannosaurids. Rather, the key developmental modification that propelled
|
||
<taxonomicName id="4C204D53350CFFB7FF8CDC1CFF609F26" authority="Osborn, 1905" box="[106,166,856,877]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="774" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350CFFB7FF8CDC1CFF609F26" box="[106,166,856,877]" italics="true" pageId="2" pageNumber="774">T. rex</emphasis>
|
||
</taxonomicName>
|
||
to giant proportions was primarily through evolutionary acceleration in the exponential stage growth rate and the transition zones bounding it. This is reflected in the regions of maximal slope on the growth curves depicted in
|
||
<figureCitation id="131B2A55350CFFB7FE52DCEEFE2D9F8B" box="[436,491,938,961]" captionStart="Figure 2" captionStartId="2.[106,159,1931,1951]" captionTargetBox="[338,1229,1274,1906]" captionTargetId="figure@2.[425,1225,1264,1864]" captionTargetPageId="2" captionText="Figure 2 Logistic growth curves for Tyrannosaurus and three related tyrannosaurids.Note that the exponential stages (the regions of maximal slope) are similar in duration but differ in slope (that is, growth rates). Regression equations (mass in kg, age in years) are as follows: T. rex, mass = {5,551/[1 + e‾0.57(age ‾ 16.1)]} + 5, r 2 = 0.953; D. torosus, mass = {1,728/[1 + e‾0.44(age ‾ 12.1)]} + 5, r 2 = 0.992; G. libratus, mass = {1,234/[1 + e‾0.38(age ‾ 12.4)]} + 5, r 2 = 0.950; A. sarcophagus, mass = {1,218/[1 + e‾0.43(age ‾ 14.1)]} + 5; r 2 = 0.985." figureDoi="http://doi.org/10.5281/zenodo.4011699" httpUri="https://zenodo.org/record/4011699/files/figure.png" pageId="2" pageNumber="774">Fig. 2</figureCitation>
|
||
and holds true regardless of which evolutionary hypothesis is correct and how the maximum growth rates are optimized. Notably, this method of attaining gigantism contrasts with that in the largest crocodilians and lizards, where ancestral growth rates were retained and the exponential stages lengthened
|
||
<bibRefCitation id="EFB14B21350CFFB7FEF0DB77FEE3980B" author="Erickson, G. M. & de Ricqles, A. & de Buffrenil, V. & Molnar, R. E. & Bayless, M. A." box="[278,293,1075,1088]" journalOrPublisher="J. Vert. Paleontol." pageId="2" pageNumber="774" pagination="966 - 970" part="23" refId="ref4591" refString="27. Erickson, G. M., de Ricqles, A., de Buffrenil, V., Molnar, R. E. & Bayless, M. A. Vermiform bones and the evolution of gigantism in Megalania - how a reptilian fox became a lion. J. Vert. Paleontol. 23, 966 - 970 (2003)." title="Vermiform bones and the evolution of gigantism in Megalania - how a reptilian fox became a lion" type="journal article" year="2003">27</bibRefCitation>
|
||
. How other dinosaurs attained gigantism within their respective sub-clades will serve as an interesting line of inquiry in the future. Does the same pattern of acceleratory growth seen here characterize the means by which all or most members of the
|
||
<taxonomicName id="4C204D53350CFFB7FF8CDBE1FF1E98F7" box="[106,216,1189,1212]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="774" phylum="Chordata" rank="order">Dinosauria</taxonomicName>
|
||
attained great size?
|
||
</paragraph>
|
||
<caption id="DF5F6658350FFFB4FCD6D9A1FC689BBF" ID-DOI="http://doi.org/10.5281/zenodo.3736477" ID-Zenodo-Dep="3736477" httpUri="https://zenodo.org/record/3736477/files/figure.png" pageId="1" pageNumber="773" startId="1.[816,869,1765,1785]" targetBox="[934,1357,755,1730]" targetPageId="1">
|
||
<paragraph id="8B9F36D0350FFFB4FCD6D9A1FC689BBF" blockId="1.[816,1477,1765,2036]" pageId="1" pageNumber="773">
|
||
<emphasis id="B954EAC2350FFFB4FCD6D9A1FCB29AB2" bold="true" box="[816,884,1765,1785]" pageId="1" pageNumber="773">Figure 1</emphasis>
|
||
Growth-line counts in tyrannosaurids and reptiles of known ages.
|
||
<emphasis id="B954EAC2350FFFB4FA88D9A1FABE9AB2" bold="true" box="[1390,1400,1765,1785]" pageId="1" pageNumber="773">a</emphasis>
|
||
, Thinsectioned
|
||
<taxonomicName id="4C204D53350FFFB4FC9BD844FC1B9B5F" box="[893,989,1792,1812]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="genus">
|
||
<emphasis id="B954EAC2350FFFB4FC9BD844FC1B9B5F" box="[893,989,1792,1812]" italics="true" pageId="1" pageNumber="773">Gorgosaurus</emphasis>
|
||
</taxonomicName>
|
||
fibula (FMNH PR 2211). The growth record in this element is complete and shows five growth lines (arrows),indicating that it died early in its sixth year of life.
|
||
<emphasis id="B954EAC2350FFFB4FC82D87CFCA99B07" bold="true" box="[868,879,1848,1868]" pageId="1" pageNumber="773">b</emphasis>
|
||
, Haematoxylin-stained fibula from an 8-year-old alligator (
|
||
<taxonomicName id="4C204D53350FFFB4FACDD87CFC6F9B23" class="Reptilia" family="Tyrannosauridae" genus="Alligator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="mississippiensis">
|
||
<emphasis id="B954EAC2350FFFB4FACDD87CFC6F9B23" italics="true" pageId="1" pageNumber="773">Alligator mississippiensis</emphasis>
|
||
</taxonomicName>
|
||
) showing the expected seven growth lines. Inset, gastralia (unstained) from 9-year-old
|
||
<taxonomicName id="4C204D53350FFFB4FC4AD834FBFD9BCF" box="[940,1083,1904,1924]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="species" species="mississippiensis">
|
||
<emphasis id="B954EAC2350FFFB4FC4AD834FBFD9BCF" box="[940,1083,1904,1924]" italics="true" pageId="1" pageNumber="773">A. mississippiensis</emphasis>
|
||
</taxonomicName>
|
||
showing the expected eight growth lines and the chemical label used to verify the periodicity of ring formation.
|
||
<emphasis id="B954EAC2350FFFB4FB13D8C8FB399BEB" bold="true" box="[1269,1279,1932,1952]" pageId="1" pageNumber="773">c</emphasis>
|
||
,
|
||
<taxonomicName id="4C204D53350FFFB4FAEED8C8FAB19BEB" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1288,1399,1932,1952]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="773" phylum="Chordata" rank="genus">
|
||
<emphasis id="B954EAC2350FFFB4FAEED8C8FAB19BEB" box="[1288,1399,1932,1952]" italics="true" pageId="1" pageNumber="773">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
rib (
|
||
<materialsCitation id="3B483C8D350FFFB4FA70D8C8FCB59BF7" ID-GBIF-Occurrence="3396395305" collectionCode="FMNH" pageId="1" pageNumber="773" specimenCode="FMNH PR2081">FMNH PR 2081</materialsCitation>
|
||
) showing the 15th to 19th growth lines. Inset, external fundamental system
|
||
<bibRefCitation id="EFB14B21350FFFB4FA55D8E2FA069BFA" author="Francillon-Viellot, H." box="[1459,1472,1958,1969]" editor="Carter, J. G." journalOrPublisher="Van Nostrand Reinhold, New York" pageId="1" pageNumber="773" pagination="471 - 530" refId="ref4193" refString="16. Francillon-Viellot, H., et al. in Biomineralization: Patterns and Evolutionary Trends (ed. Carter, J. G.) 471 - 530 (Van Nostrand Reinhold, New York, 1990)." type="book chapter" volumeTitle="Biomineralization: Patterns and Evolutionary Trends" year="1990">
|
||
<superScript id="7C559B98350FFFB4FA55D8E2FA069BFA" attach="left" box="[1459,1472,1958,1969]" fontSize="5" pageId="1" pageNumber="773">16</superScript>
|
||
</bibRefCitation>
|
||
with nine tightly spaced growth lines, indicating late-adulthood senescence and growthrate attenuation.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF5F6658350CFFB7FF8CDFEBFDE69CF4" ID-Table-UUID="DF5F6658350CFFB7FF8CDFEBFDE69CF4" box="[106,544,175,191]" httpUri="http://table.plazi.org/id/DF5F6658350CFFB7FF8CDFEBFDE69CF4" pageId="2" pageNumber="774" startId="2.[106,147,175,191]" targetBox="[106,1461,206,655]" targetIsTable="true" targetPageId="2">
|
||
<paragraph id="8B9F36D0350CFFB7FF8CDFEBFDE69CF4" box="[106,544,175,191]" pageId="2" pageNumber="774">
|
||
<tableCitation id="C6A2036B350CFFB7FF8CDFEBFF679CF4" box="[106,161,175,191]" captionStart="Table 1" captionStartId="2.[106,147,175,191]" captionTargetBox="[106,1461,206,655]" captionTargetPageId="2" captionText="Table 1 Specimens, size measures, and longevity data" httpUri="http://table.plazi.org/id/DF5F6658350CFFB7FF8CDFEBFDE69CF4" pageId="2" pageNumber="774" tableUuid="DF5F6658350CFFB7FF8CDFEBFDE69CF4">Table 1</tableCitation>
|
||
<emphasis id="B954EAC2350CFFB7FF40DFEBFDE69CF4" bold="true" box="[166,544,175,191]" pageId="2" pageNumber="774">Specimens, size measures, and longevity data</emphasis>
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8B9F36D0350CFFB7FF8CDF8AFAD89EC4" pageId="2" pageNumber="774">
|
||
<table id="F920C470350C004AFF8CDF8AFA739EC4" box="[106,1461,206,655]" gridcols="6" gridrows="21" pageId="2" pageNumber="774">
|
||
<tr id="35103492350C004AFF8CDF8AFA739CA5" box="[106,1461,206,238]" gridrow="0" pageId="2" pageNumber="774">
|
||
<th id="76C15DEE350C004AFF8CDF8AFA739CA5" box="[106,1461,206,238]" colspan="6" colspanRight="5" gridcol="0" gridrow="0" pageId="2" pageNumber="774">Taxon Specimen no. Elements examined Femoral length (cm) Body mass (kg) Longevity (years); no. of growth lines in EFS ...................................................................................................................................................................................................................................................................................................................................................................</th>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDFBCFA739D43" box="[106,1461,248,264]" gridrow="1" pageId="2" pageNumber="774">
|
||
<th id="76C15DEE350C004AFF8CDFBCFEF09D43" box="[106,310,248,264]" gridcol="0" gridrow="1" pageId="2" pageNumber="774">
|
||
<taxonomicName id="4C204D53350CFFB7FF8CDFBCFF349D43" authority="Osborn, 1905" box="[106,242,248,264]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="774" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350CFFB7FF8CDFBCFF349D43" box="[106,242,248,264]" italics="true" pageId="2" pageNumber="774">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="76C15DEE350C004AFE85DFBCFDEC9D43" box="[355,554,248,264]" gridcol="1" gridrow="1" pageId="2" pageNumber="774">
|
||
<materialsCitation id="3B483C8D350CFFB7FE6ADFBCFDC79D43" ID-GBIF-Occurrence="3396395301" box="[396,513,248,264]" collectionCode="FMNH" pageId="2" pageNumber="774" specimenCode="FMNH PR2081">FMNH PR 2081</materialsCitation>
|
||
</td>
|
||
<td id="76C15DEE350C004AFD96DFBCFD169D43" box="[624,720,248,264]" gridcol="2" gridrow="1" pageId="2" pageNumber="774">G, R, F, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDFBCFC409D43" box="[825,902,248,264]" gridcol="3" gridrow="1" pageId="2" pageNumber="774">134.5</td>
|
||
<td id="76C15DEE350C004AFC1DDFBCFBE79D43" box="[1019,1057,248,264]" gridcol="4" gridrow="1" pageId="2" pageNumber="774">5654</td>
|
||
<td id="76C15DEE350C004AFAE4DFBCFA739D43" box="[1282,1461,248,264]" gridcol="5" gridrow="1" pageId="2" pageNumber="774">28; 9</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDE48FA739D57" box="[106,1461,268,284]" gridrow="2" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DE48FDEC9D57" box="[355,554,268,284]" gridcol="1" gridrow="2" pageId="2" pageNumber="774">
|
||
<materialsCitation id="3B483C8D350CFFB7FE76DE48FE3B9D57" ID-GBIF-Occurrence="3396395302" box="[400,509,268,284]" collectionCode="TMP" pageId="2" pageNumber="774" specimenCode="TMP 81.12.1">RTMP 81.12.1</materialsCitation>
|
||
</td>
|
||
<td id="76C15DEE350C004AFD96DE48FD169D57" box="[624,720,268,284]" gridcol="2" gridrow="2" pageId="2" pageNumber="774">R, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDE48FC409D57" box="[825,902,268,284]" gridcol="3" gridrow="2" pageId="2" pageNumber="774">128.4 est.</td>
|
||
<td id="76C15DEE350C004AFC1DDE48FBE79D57" box="[1019,1057,268,284]" gridcol="4" gridrow="2" pageId="2" pageNumber="774">5040</td>
|
||
<td id="76C15DEE350C004AFAE4DE48FA739D57" box="[1282,1461,268,284]" gridcol="5" gridrow="2" pageId="2" pageNumber="774">22</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDE65FA739D7A" box="[106,1461,289,305]" gridrow="3" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DE65FDEC9D7A" box="[355,554,289,305]" gridcol="1" gridrow="3" pageId="2" pageNumber="774">
|
||
<materialsCitation id="3B483C8D350CFFB7FE73DE65FE329D7A" ID-GBIF-Occurrence="3396395304" box="[405,500,289,305]" collectionCode="TMP" pageId="2" pageNumber="774" specimenCode="TMP 81.6.1">RTMP 81.6.</materialsCitation>
|
||
1
|
||
</td>
|
||
<td id="76C15DEE350C004AFD96DE65FD169D7A" box="[624,720,289,305]" gridcol="2" gridrow="3" pageId="2" pageNumber="774">G, R, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDE65FC409D7A" box="[825,902,289,305]" gridcol="3" gridrow="3" pageId="2" pageNumber="774">120</td>
|
||
<td id="76C15DEE350C004AFC1DDE65FBE79D7A" box="[1019,1057,289,305]" gridcol="4" gridrow="3" pageId="2" pageNumber="774">3230</td>
|
||
<td id="76C15DEE350C004AFAE4DE65FA739D7A" box="[1282,1461,289,305]" gridcol="5" gridrow="3" pageId="2" pageNumber="774">18</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDE72FA739D0D" box="[106,1461,310,326]" gridrow="4" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DE72FDEC9D0D" box="[355,554,310,326]" gridcol="1" gridrow="4" pageId="2" pageNumber="774">
|
||
<materialsCitation id="3B483C8D350CFFB7FE69DE72FE3D9D0D" ID-GBIF-Occurrence="3396395308" box="[399,507,310,326]" collectionCode="ICM" pageId="2" pageNumber="774" specimenCode="ICM 2001.90.1">ICM 2001.90.</materialsCitation>
|
||
1
|
||
</td>
|
||
<td id="76C15DEE350C004AFD96DE72FD169D0D" box="[624,720,310,326]" gridcol="2" gridrow="4" pageId="2" pageNumber="774">G, R</td>
|
||
<td id="76C15DEE350C004AFCDFDE72FC409D0D" box="[825,902,310,326]" gridcol="3" gridrow="4" pageId="2" pageNumber="774">116.8 est.</td>
|
||
<td id="76C15DEE350C004AFC1DDE72FBE79D0D" box="[1019,1057,310,326]" gridcol="4" gridrow="4" pageId="2" pageNumber="774">2984</td>
|
||
<td id="76C15DEE350C004AFAE4DE72FA739D0D" box="[1282,1461,310,326]" gridcol="5" gridrow="4" pageId="2" pageNumber="774">16</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDE0EFA739D11" box="[106,1461,330,346]" gridrow="5" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DE0EFDEC9D11" box="[355,554,330,346]" gridcol="1" gridrow="5" pageId="2" pageNumber="774">
|
||
<materialsCitation id="3B483C8D350CFFB7FE73DE0EFE3E9D11" ID-GBIF-Occurrence="3396395306" box="[405,504,330,346]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/ContactSheet.php?QueryName=DetailedQuery&StartAt=1&QueryPage=%2Fdinosaur-institute%2Findex.php&Restriction=%28exists+%28SecDepartment_tab+where+%28SecDepartment+contains+%27Dinosaur+Institute%27%29%29%29&col_SpeSpecimenNoInteger=23845&LimitPerPage=20&Search=Search& " pageId="2" pageNumber="774" specimenCode="LACM 23845">LACM 23845</materialsCitation>
|
||
</td>
|
||
<td id="76C15DEE350C004AFD96DE0EFD169D11" box="[624,720,330,346]" gridcol="2" gridrow="5" pageId="2" pageNumber="774">P, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDE0EFC409D11" box="[825,902,330,346]" gridcol="3" gridrow="5" pageId="2" pageNumber="774">98.9</td>
|
||
<td id="76C15DEE350C004AFC1DDE0EFBE79D11" box="[1019,1057,330,346]" gridcol="4" gridrow="5" pageId="2" pageNumber="774">1807</td>
|
||
<td id="76C15DEE350C004AFAE4DE0EFA739D11" box="[1282,1461,330,346]" gridcol="5" gridrow="5" pageId="2" pageNumber="774">14</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDE1BFA739D24" box="[106,1461,351,367]" gridrow="6" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DE1BFDEC9D24" box="[355,554,351,367]" gridcol="1" gridrow="6" pageId="2" pageNumber="774">
|
||
<materialsCitation id="3B483C8D350CFFB7FE75DE1BFE3C9D24" ID-GBIF-Occurrence="3396395303" box="[403,506,351,367]" collectionCode="AMNH" httpUri="http://research.amnh.org/paleontology/search.php?action=detail&specimen_id=89189" pageId="2" pageNumber="774" specimenCode="AMNH 30564">AMNH 30564</materialsCitation>
|
||
</td>
|
||
<td id="76C15DEE350C004AFD96DE1BFD169D24" box="[624,720,351,367]" gridcol="2" gridrow="6" pageId="2" pageNumber="774">G, R, F</td>
|
||
<td id="76C15DEE350C004AFCDFDE1BFC409D24" box="[825,902,351,367]" gridcol="3" gridrow="6" pageId="2" pageNumber="774">98</td>
|
||
<td id="76C15DEE350C004AFC1DDE1BFBE79D24" box="[1019,1057,351,367]" gridcol="4" gridrow="6" pageId="2" pageNumber="774">1761</td>
|
||
<td id="76C15DEE350C004AFAE4DE1BFA739D24" box="[1282,1461,351,367]" gridcol="5" gridrow="6" pageId="2" pageNumber="774">15</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDE37FA739DC8" box="[106,1461,371,387]" gridrow="7" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DE37FDEC9DC8" box="[355,554,371,387]" gridcol="1" gridrow="7" pageId="2" pageNumber="774">
|
||
<materialsCitation id="3B483C8D350CFFB7FE73DE37FE3E9DC8" ID-GBIF-Occurrence="3396395309" box="[405,504,371,387]" collectionCode="LACM" httpUri="https://collections.nhm.org/dinosaur-institute/ContactSheet.php?QueryName=DetailedQuery&StartAt=1&QueryPage=%2Fdinosaur-institute%2Findex.php&Restriction=%28exists+%28SecDepartment_tab+where+%28SecDepartment+contains+%27Dinosaur+Institute%27%29%29%29&col_SpeSpecimenNoInteger=23845&LimitPerPage=20&Search=Search& " pageId="2" pageNumber="774" specimenCode="LACM 28471">LACM 28471</materialsCitation>
|
||
</td>
|
||
<td id="76C15DEE350C004AFD96DE37FD169DC8" box="[624,720,371,387]" gridcol="2" gridrow="7" pageId="2" pageNumber="774">R, C</td>
|
||
<td id="76C15DEE350C004AFCDFDE37FC409DC8" box="[825,902,371,387]" gridcol="3" gridrow="7" pageId="2" pageNumber="774">25.2 est.</td>
|
||
<td id="76C15DEE350C004AFC1DDE37FBE79DC8" box="[1019,1057,371,387]" gridcol="4" gridrow="7" pageId="2" pageNumber="774">29.9</td>
|
||
<td id="76C15DEE350C004AFAE4DE37FA739DC8" box="[1282,1461,371,387]" gridcol="5" gridrow="7" pageId="2" pageNumber="774">2</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDECCFA739DD3" box="[106,1461,392,408]" gridrow="8" pageId="2" pageNumber="774">
|
||
<th id="76C15DEE350C004AFF8CDECCFEF09DD3" box="[106,310,392,408]" gridcol="0" gridrow="8" pageId="2" pageNumber="774">
|
||
<taxonomicName id="4C204D53350CFFB7FF8CDECCFEC29DD3" baseAuthorityName="Lambe" baseAuthorityYear="1914" box="[106,260,392,408]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="774" phylum="Chordata" rank="species" species="libratus">
|
||
<emphasis id="B954EAC2350CFFB7FF8CDECCFEC29DD3" box="[106,260,392,408]" italics="true" pageId="2" pageNumber="774">Gorgosaurus libratus</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="76C15DEE350C004AFE85DECCFDEC9DD3" box="[355,554,392,408]" gridcol="1" gridrow="8" pageId="2" pageNumber="774">RTMP 94.12.602</td>
|
||
<td id="76C15DEE350C004AFD96DECCFD169DD3" box="[624,720,392,408]" gridcol="2" gridrow="8" pageId="2" pageNumber="774">R, G, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDECCFC409DD3" box="[825,902,392,408]" gridcol="3" gridrow="8" pageId="2" pageNumber="774">91.6</td>
|
||
<td id="76C15DEE350C004AFC1DDECCFBE79DD3" box="[1019,1057,392,408]" gridcol="4" gridrow="8" pageId="2" pageNumber="774">1105</td>
|
||
<td id="76C15DEE350C004AFAE4DECCFA739DD3" box="[1282,1461,392,408]" gridcol="5" gridrow="8" pageId="2" pageNumber="774">18</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDED8FA739DE7" box="[106,1461,412,428]" gridrow="9" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DED8FDEC9DE7" box="[355,554,412,428]" gridcol="1" gridrow="9" pageId="2" pageNumber="774">RTMP 73.30.1</td>
|
||
<td id="76C15DEE350C004AFD96DED8FD169DE7" box="[624,720,412,428]" gridcol="2" gridrow="9" pageId="2" pageNumber="774">F, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDED8FC409DE7" box="[825,902,412,428]" gridcol="3" gridrow="9" pageId="2" pageNumber="774">80.4 est.</td>
|
||
<td id="76C15DEE350C004AFC1DDED8FBE79DE7" box="[1019,1057,412,428]" gridcol="4" gridrow="9" pageId="2" pageNumber="774">747</td>
|
||
<td id="76C15DEE350C004AFAE4DED8FA739DE7" box="[1282,1461,412,428]" gridcol="5" gridrow="9" pageId="2" pageNumber="774">14</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDEF5FA739D8A" box="[106,1461,433,449]" gridrow="10" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DEF5FDEC9D8A" box="[355,554,433,449]" gridcol="1" gridrow="10" pageId="2" pageNumber="774">RTMP 99.33.1</td>
|
||
<td id="76C15DEE350C004AFD96DEF5FD169D8A" box="[624,720,433,449]" gridcol="2" gridrow="10" pageId="2" pageNumber="774">G, F, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDEF5FC409D8A" box="[825,902,433,449]" gridcol="3" gridrow="10" pageId="2" pageNumber="774">75</td>
|
||
<td id="76C15DEE350C004AFC1DDEF5FBE79D8A" box="[1019,1057,433,449]" gridcol="4" gridrow="10" pageId="2" pageNumber="774">607</td>
|
||
<td id="76C15DEE350C004AFAE4DEF5FA739D8A" box="[1282,1461,433,449]" gridcol="5" gridrow="10" pageId="2" pageNumber="774">14</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDE82FA739D9D" box="[106,1461,454,470]" gridrow="11" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DE82FDEC9D9D" box="[355,554,454,470]" gridcol="1" gridrow="11" pageId="2" pageNumber="774">RTMP 86.144.1</td>
|
||
<td id="76C15DEE350C004AFD96DE82FD169D9D" box="[624,720,454,470]" gridcol="2" gridrow="11" pageId="2" pageNumber="774">R, F, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDE82FC409D9D" box="[825,902,454,470]" gridcol="3" gridrow="11" pageId="2" pageNumber="774">54.2</td>
|
||
<td id="76C15DEE350C004AFC1DDE82FBE79D9D" box="[1019,1057,454,470]" gridcol="4" gridrow="11" pageId="2" pageNumber="774">229</td>
|
||
<td id="76C15DEE350C004AFAE4DE82FA739D9D" box="[1282,1461,454,470]" gridcol="5" gridrow="11" pageId="2" pageNumber="774">7</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDE9EFA739DA1" box="[106,1461,474,490]" gridrow="12" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DE9EFDEC9DA1" box="[355,554,474,490]" gridcol="1" gridrow="12" pageId="2" pageNumber="774">FMNH PR 2211</td>
|
||
<td id="76C15DEE350C004AFD96DE9EFD169DA1" box="[624,720,474,490]" gridcol="2" gridrow="12" pageId="2" pageNumber="774">G, R, F, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDE9EFC409DA1" box="[825,902,474,490]" gridcol="3" gridrow="12" pageId="2" pageNumber="774">44.5</td>
|
||
<td id="76C15DEE350C004AFC1DDE9EFBE79DA1" box="[1019,1057,474,490]" gridcol="4" gridrow="12" pageId="2" pageNumber="774">127</td>
|
||
<td id="76C15DEE350C004AFAE4DE9EFA739DA1" box="[1282,1461,474,490]" gridcol="5" gridrow="12" pageId="2" pageNumber="774">5</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDEABFA739DB4" box="[106,1461,495,511]" gridrow="13" pageId="2" pageNumber="774">
|
||
<th id="76C15DEE350C004AFF8CDEABFEF09DB4" box="[106,310,495,511]" gridcol="0" gridrow="13" pageId="2" pageNumber="774">
|
||
<taxonomicName id="4C204D53350CFFB7FF8CDEABFEF09DB4" authorityName="Osborn" authorityYear="1905" box="[106,310,495,511]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="774" phylum="Chordata" rank="species" species="sarcophagus">
|
||
<emphasis id="B954EAC2350CFFB7FF8CDEABFEF09DB4" box="[106,310,495,511]" italics="true" pageId="2" pageNumber="774">Albertosaurus sarcophagus</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="76C15DEE350C004AFE85DEABFDEC9DB4" box="[355,554,495,511]" gridcol="1" gridrow="13" pageId="2" pageNumber="774">RTMP 81.10.1</td>
|
||
<td id="76C15DEE350C004AFD96DEABFD169DB4" box="[624,720,495,511]" gridcol="2" gridrow="13" pageId="2" pageNumber="774">G, R, F, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDEABFC409DB4" box="[825,902,495,511]" gridcol="3" gridrow="13" pageId="2" pageNumber="774">89.5</td>
|
||
<td id="76C15DEE350C004AFC1DDEABFBE79DB4" box="[1019,1057,495,511]" gridcol="4" gridrow="13" pageId="2" pageNumber="774">1142</td>
|
||
<td id="76C15DEE350C004AFAE4DEABFA739DB4" box="[1282,1461,495,511]" gridcol="5" gridrow="13" pageId="2" pageNumber="774">24; 2</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDD47FA739E58" box="[106,1461,515,531]" gridrow="14" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DD47FDEC9E58" box="[355,554,515,531]" gridcol="1" gridrow="14" pageId="2" pageNumber="774">AMNH 5432</td>
|
||
<td id="76C15DEE350C004AFD96DD47FD169E58" box="[624,720,515,531]" gridcol="2" gridrow="14" pageId="2" pageNumber="774">P, F</td>
|
||
<td id="76C15DEE350C004AFCDFDD47FC409E58" box="[825,902,515,531]" gridcol="3" gridrow="14" pageId="2" pageNumber="774">99.3 est.</td>
|
||
<td id="76C15DEE350C004AFC1DDD47FBE79E58" box="[1019,1057,515,531]" gridcol="4" gridrow="14" pageId="2" pageNumber="774">1282</td>
|
||
<td id="76C15DEE350C004AFAE4DD47FA739E58" box="[1282,1461,515,531]" gridcol="5" gridrow="14" pageId="2" pageNumber="774">22</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDD5CFA739E63" box="[106,1461,536,552]" gridrow="15" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DD5CFDEC9E63" box="[355,554,536,552]" gridcol="1" gridrow="15" pageId="2" pageNumber="774">USNM 12814/AMNH 5428</td>
|
||
<td id="76C15DEE350C004AFD96DD5CFD169E63" box="[624,720,536,552]" gridcol="2" gridrow="15" pageId="2" pageNumber="774">R</td>
|
||
<td id="76C15DEE350C004AFCDFDD5CFC409E63" box="[825,902,536,552]" gridcol="3" gridrow="15" pageId="2" pageNumber="774">86.0 est.</td>
|
||
<td id="76C15DEE350C004AFC1DDD5CFBE79E63" box="[1019,1057,536,552]" gridcol="4" gridrow="15" pageId="2" pageNumber="774">1013</td>
|
||
<td id="76C15DEE350C004AFAE4DD5CFA739E63" box="[1282,1461,536,552]" gridcol="5" gridrow="15" pageId="2" pageNumber="774">18</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDD69FA739E76" box="[106,1461,557,573]" gridrow="16" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DD69FDEC9E76" box="[355,554,557,573]" gridcol="1" gridrow="16" pageId="2" pageNumber="774">RTMP 86.64.01</td>
|
||
<td id="76C15DEE350C004AFD96DD69FD169E76" box="[624,720,557,573]" gridcol="2" gridrow="16" pageId="2" pageNumber="774">R, F, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDD69FC409E76" box="[825,902,557,573]" gridcol="3" gridrow="16" pageId="2" pageNumber="774">78.2 est.</td>
|
||
<td id="76C15DEE350C004AFC1DDD69FBE79E76" box="[1019,1057,557,573]" gridcol="4" gridrow="16" pageId="2" pageNumber="774">762</td>
|
||
<td id="76C15DEE350C004AFAE4DD69FA739E76" box="[1282,1461,557,573]" gridcol="5" gridrow="16" pageId="2" pageNumber="774">15</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDD05FA739E1A" box="[106,1461,577,593]" gridrow="17" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DD05FDEC9E1A" box="[355,554,577,593]" gridcol="1" gridrow="17" pageId="2" pageNumber="774">RTMP 2002.45.46</td>
|
||
<td id="76C15DEE350C004AFD96DD05FD169E1A" box="[624,720,577,593]" gridcol="2" gridrow="17" pageId="2" pageNumber="774">F, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDD05FC409E1A" box="[825,902,577,593]" gridcol="3" gridrow="17" pageId="2" pageNumber="774">31.6 est.</td>
|
||
<td id="76C15DEE350C004AFC1DDD05FBE79E1A" box="[1019,1057,577,593]" gridcol="4" gridrow="17" pageId="2" pageNumber="774">50.3</td>
|
||
<td id="76C15DEE350C004AFAE4DD05FA739E1A" box="[1282,1461,577,593]" gridcol="5" gridrow="17" pageId="2" pageNumber="774">2</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDD12FA739E2D" box="[106,1461,598,614]" gridrow="18" pageId="2" pageNumber="774">
|
||
<th id="76C15DEE350C004AFF8CDD12FEF09E2D" box="[106,310,598,614]" gridcol="0" gridrow="18" pageId="2" pageNumber="774">
|
||
<taxonomicName id="4C204D53350CFFB7FF8CDD12FEDF9E2D" baseAuthorityName="Russell" baseAuthorityYear="1970" box="[106,281,598,614]" class="Reptilia" family="Tyrannosauridae" genus="Daspletosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="774" phylum="Chordata" rank="species" species="torosus">
|
||
<emphasis id="B954EAC2350CFFB7FF8CDD12FEDF9E2D" box="[106,281,598,614]" italics="true" pageId="2" pageNumber="774">Daspletosaurus torosus</emphasis>
|
||
</taxonomicName>
|
||
</th>
|
||
<td id="76C15DEE350C004AFE85DD12FDEC9E2D" box="[355,554,598,614]" gridcol="1" gridrow="18" pageId="2" pageNumber="774">FMNH PR 308</td>
|
||
<td id="76C15DEE350C004AFD96DD12FD169E2D" box="[624,720,598,614]" gridcol="2" gridrow="18" pageId="2" pageNumber="774">G, R</td>
|
||
<td id="76C15DEE350C004AFCDFDD12FC409E2D" box="[825,902,598,614]" gridcol="3" gridrow="18" pageId="2" pageNumber="774">96.0 est.</td>
|
||
<td id="76C15DEE350C004AFC1DDD12FBE79E2D" box="[1019,1057,598,614]" gridcol="4" gridrow="18" pageId="2" pageNumber="774">1791</td>
|
||
<td id="76C15DEE350C004AFAE4DD12FA739E2D" box="[1282,1461,598,614]" gridcol="5" gridrow="18" pageId="2" pageNumber="774">21; 5</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDD2EFA739E31" box="[106,1461,618,634]" gridrow="19" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DD2EFDEC9E31" box="[355,554,618,634]" gridcol="1" gridrow="19" pageId="2" pageNumber="774">AMNH 5438</td>
|
||
<td id="76C15DEE350C004AFD96DD2EFD169E31" box="[624,720,618,634]" gridcol="2" gridrow="19" pageId="2" pageNumber="774">R</td>
|
||
<td id="76C15DEE350C004AFCDFDD2EFC409E31" box="[825,902,618,634]" gridcol="3" gridrow="19" pageId="2" pageNumber="774">84.1 est.</td>
|
||
<td id="76C15DEE350C004AFC1DDD2EFBE79E31" box="[1019,1057,618,634]" gridcol="4" gridrow="19" pageId="2" pageNumber="774">1518</td>
|
||
<td id="76C15DEE350C004AFAE4DD2EFA739E31" box="[1282,1461,618,634]" gridcol="5" gridrow="19" pageId="2" pageNumber="774">17</td>
|
||
</tr>
|
||
<tr id="35103492350C004AFF8CDD3BFA739EC4" box="[106,1461,639,655]" gridrow="20" pageId="2" pageNumber="774" rowspan-0="1">
|
||
<td id="76C15DEE350C004AFE85DD3BFDEC9EC4" box="[355,554,639,655]" gridcol="1" gridrow="20" pageId="2" pageNumber="774">RTMP 94.143.1</td>
|
||
<td id="76C15DEE350C004AFD96DD3BFD169EC4" box="[624,720,639,655]" gridcol="2" gridrow="20" pageId="2" pageNumber="774">G, OLB</td>
|
||
<td id="76C15DEE350C004AFCDFDD3BFC409EC4" box="[825,902,639,655]" gridcol="3" gridrow="20" pageId="2" pageNumber="774">62.6 est.</td>
|
||
<td id="76C15DEE350C004AFC1DDD3BFBE79EC4" box="[1019,1057,639,655]" gridcol="4" gridrow="20" pageId="2" pageNumber="774">496</td>
|
||
<td id="76C15DEE350C004AFAE4DD3BFA739EC4" box="[1282,1461,639,655]" gridcol="5" gridrow="20" pageId="2" pageNumber="774">10</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph id="8B9F36D0350CFFB7FF8CDDC8FA759ED0" box="[106,1459,652,667]" pageId="2" pageNumber="774">...................................................................................................................................................................................................................................................................................................................................................................</paragraph>
|
||
<paragraph id="8B9F36D0350CFFB7FF8CDDE4FAED9E89" pageId="2" pageNumber="774">
|
||
<tableNote id="76C6375E350CFFB7FF8CDDE4FAED9E89" pageId="2" pageNumber="774" targetBox="[106,1461,206,655]" targetPageId="2">
|
||
FMNH, The Field Museum; RTMP, Royal Tyrrell Museum of Palaeontology; ICM, Indianapolis Children’s Museum; LACM, Los Angeles County Museum; AMNH, American Museum of Natural History; USNM, United States National Museum. R, rib; G, gastralia; F, fibula; P, pubis; C, dermal skull bones; OLB, other long bones; est., estimated; EFS, external fundamental system
|
||
<bibRefCitation id="EFB14B21350CFFB7FAFADDF6FAE19EF1" author="Francillon-Viellot, H." box="[1308,1319,690,698]" editor="Carter, J. G." journalOrPublisher="Van Nostrand Reinhold, New York" pageId="2" pageNumber="774" pagination="471 - 530" refId="ref4193" refString="16. Francillon-Viellot, H., et al. in Biomineralization: Patterns and Evolutionary Trends (ed. Carter, J. G.) 471 - 530 (Van Nostrand Reinhold, New York, 1990)." type="book chapter" volumeTitle="Biomineralization: Patterns and Evolutionary Trends" year="1990">16</bibRefCitation>
|
||
.
|
||
</tableNote>
|
||
</paragraph>
|
||
<paragraph id="8B9F36D0350CFFB6FCDEDC47FE419F37" lastBlockId="3.[122,785,172,893]" lastPageId="3" lastPageNumber="775" pageId="2" pageNumber="774">
|
||
Besides revealing how the evolution of
|
||
<taxonomicName id="4C204D53350CFFB7FB3BDC40FADF9F52" authority="Osborn, 1905" box="[1245,1305,772,793]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="774" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350CFFB7FB3BDC40FADF9F52" box="[1245,1305,772,793]" italics="true" pageId="2" pageNumber="774">T. rex</emphasis>
|
||
</taxonomicName>
|
||
gigantism was obtained, the data garnered here provide for a more comprehensive understanding of tyrannosaurid biology. For instance the presence of thin, tightly packed growth lines late in development (
|
||
<figureCitation id="131B2A55350CFFB7FA96DC12FA6D9F26" box="[1392,1451,854,877]" captionStart="Figure 1" captionStartId="1.[816,869,1765,1785]" captionTargetBox="[934,1357,755,1730]" captionTargetId="figure@1.[921,1370,742,1742]" captionTargetPageId="1" captionText="Figure 1 Growth-line counts in tyrannosaurids and reptiles of known ages. a, Thinsectioned Gorgosaurus fibula (FMNH PR 2211). The growth record in this element is complete and shows five growth lines (arrows),indicating that it died early in its sixth year of life. b, Haematoxylin-stained fibula from an 8-year-old alligator (Alligator mississippiensis) showing the expected seven growth lines. Inset, gastralia (unstained) from 9-year-old A. mississippiensis showing the expected eight growth lines and the chemical label used to verify the periodicity of ring formation.c, Tyrannosaurus rib (FMNH PR 2081) showing the 15th to 19th growth lines. Inset, external fundamental system16 with nine tightly spaced growth lines, indicating late-adulthood senescence and growthrate attenuation." figureDoi="http://doi.org/10.5281/zenodo.3736477" httpUri="https://zenodo.org/record/3736477/files/figure.png" pageId="2" pageNumber="774">Fig. 1</figureCitation>
|
||
) shows that these animals, like nearly all (if not all) dinosaurs, had determinate growth
|
||
<bibRefCitation id="EFB14B21350CFFB7FC02DCC8FC2D9FD2" author="Erickson, G. M. & Curry-Rogers, K. & Yerby, S." box="[996,1003,908,921]" journalOrPublisher="Nature" pageId="2" pageNumber="774" pagination="429 - 433" part="412" refId="ref3720" refString="3. Erickson, G. M., Curry-Rogers, K. & Yerby, S. Dinosaur growth patterns and rapid avian growth rates. Nature 412, 429 - 433 (2001)." title="Dinosaur growth patterns and rapid avian growth rates" type="journal article" year="2001">3</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFB14B21350CFFB7FC08DCC8FC389FD2" author="Chinsamy, A." box="[1006,1022,908,921]" journalOrPublisher="Palaeontol. Afr." pageId="2" pageNumber="774" pagination="77 - 82" part="27" refId="ref4104" refString="14. Chinsamy, A. Physiological implications of the bone histology of Syntarsus rhodesiensis (Saurischia: Theropoda). Palaeontol. Afr. 27, 77 - 82 (1993)." title="Physiological implications of the bone histology of Syntarsus rhodesiensis (Saurischia: Theropoda)" type="journal article" year="1993">14</bibRefCitation>
|
||
. They would not have gained an appreciably greater size than the largest specimens studied here and could spend nearly 30% of their lives as full-grown adults (
|
||
<figureCitation id="131B2A55350CFFB7FB08DC82FAED9F96" box="[1262,1323,966,989]" captionStart="Figure 2" captionStartId="2.[106,159,1931,1951]" captionTargetBox="[338,1229,1274,1906]" captionTargetId="figure@2.[425,1225,1264,1864]" captionTargetPageId="2" captionText="Figure 2 Logistic growth curves for Tyrannosaurus and three related tyrannosaurids.Note that the exponential stages (the regions of maximal slope) are similar in duration but differ in slope (that is, growth rates). Regression equations (mass in kg, age in years) are as follows: T. rex, mass = {5,551/[1 + e‾0.57(age ‾ 16.1)]} + 5, r 2 = 0.953; D. torosus, mass = {1,728/[1 + e‾0.44(age ‾ 12.1)]} + 5, r 2 = 0.992; G. libratus, mass = {1,234/[1 + e‾0.38(age ‾ 12.4)]} + 5, r 2 = 0.950; A. sarcophagus, mass = {1,218/[1 + e‾0.43(age ‾ 14.1)]} + 5; r 2 = 0.985." figureDoi="http://doi.org/10.5281/zenodo.4011699" httpUri="https://zenodo.org/record/4011699/files/figure.png" pageId="2" pageNumber="774">Fig. 2</figureCitation>
|
||
). In addition, the maximal growth rates for these tyrannosaurid species are only 33–52% of the rates expected for non-avian dinosaurs of their size when compared with the more broadly sampled data of Erickson
|
||
<emphasis id="B954EAC2350CFFB7FCC6DB73FC899800" box="[800,847,1078,1100]" italics="true" pageId="2" pageNumber="774">et al.</emphasis>
|
||
<bibRefCitation id="EFB14B21350CFFB7FCA8DB77FC90980B" author="Erickson, G. M. & Curry-Rogers, K. & Yerby, S." box="[846,854,1075,1088]" journalOrPublisher="Nature" pageId="2" pageNumber="774" pagination="429 - 433" part="412" refId="ref3720" refString="3. Erickson, G. M., Curry-Rogers, K. & Yerby, S. Dinosaur growth patterns and rapid avian growth rates. Nature 412, 429 - 433 (2001)." title="Dinosaur growth patterns and rapid avian growth rates" type="journal article" year="2001">3</bibRefCitation>
|
||
. This provides the first evidence of its kind pointing to major differences in whole body growth rates among a non-avian dinosaur sub-clade. Such findings are not unexpected because similar patterns (for example primates within Eutheria) occur within living vertebrate groups
|
||
<bibRefCitation id="EFB14B21350CFFB7FC3FDBE7FC2F98FB" author="Case, T. E." box="[985,1001,1187,1200]" journalOrPublisher="Q. Rev. Biol." pageId="2" pageNumber="774" pagination="243 - 282" part="53" refId="ref4658" refString="28. Case, T. E. On the evolution and adaptive significance of post-natal growth rates in the terrestrial vertebrates. Q. Rev. Biol. 53, 243 - 282 (1978)." title="On the evolution and adaptive significance of post-natal growth rates in the terrestrial vertebrates" type="journal article" year="1978">28</bibRefCitation>
|
||
. Our findings also have a bearing on the biomechanical capacities of tyrannosaurids.
|
||
<taxonomicName id="4C204D53350DFFB6FDCCDFEAFD989C88" authority="Osborn, 1905" box="[554,606,174,195]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="775" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis id="B954EAC2350DFFB6FDCCDFEAFD989C88" box="[554,606,174,195]" italics="true" pageId="3" pageNumber="775">T. rex</emphasis>
|
||
</taxonomicName>
|
||
’s capacity for ‘fast running’ was biomechanially infeasible after a body mass of about 1,000 kg was attained
|
||
<bibRefCitation id="EFB14B21350DFFB6FEADDFA6FE9C9CA4" author="Hutchinson, J. & Garcia, M." box="[331,346,226,239]" journalOrPublisher="Nature" pageId="3" pageNumber="775" pagination="1018 - 1021" part="415" refId="ref4002" refString="11. Hutchinson, J. & Garcia, M. Tyrannosaurus was not a fast runner. Nature 415, 1018 - 1021 (2002)." title="Tyrannosaurus was not a fast runner" type="journal article" year="2002">
|
||
<superScript id="7C559B98350DFFB6FEADDFA6FE9C9CA4" attach="right" box="[331,346,226,239]" fontSize="5" pageId="3" pageNumber="775">11</superScript>
|
||
</bibRefCitation>
|
||
. This corresponds to a juvenile-sized animal just 13 years of age on the basis of our longevity data and conservative estimates of body mass (
|
||
<figureCitation id="131B2A55350DFFB6FDE7DE58FD879D78" box="[513,577,284,307]" captionStart="Figure 2" captionStartId="2.[106,159,1931,1951]" captionTargetBox="[338,1229,1274,1906]" captionTargetId="figure@2.[425,1225,1264,1864]" captionTargetPageId="2" captionText="Figure 2 Logistic growth curves for Tyrannosaurus and three related tyrannosaurids.Note that the exponential stages (the regions of maximal slope) are similar in duration but differ in slope (that is, growth rates). Regression equations (mass in kg, age in years) are as follows: T. rex, mass = {5,551/[1 + e‾0.57(age ‾ 16.1)]} + 5, r 2 = 0.953; D. torosus, mass = {1,728/[1 + e‾0.44(age ‾ 12.1)]} + 5, r 2 = 0.992; G. libratus, mass = {1,234/[1 + e‾0.38(age ‾ 12.4)]} + 5, r 2 = 0.950; A. sarcophagus, mass = {1,218/[1 + e‾0.43(age ‾ 14.1)]} + 5; r 2 = 0.985." figureDoi="http://doi.org/10.5281/zenodo.4011699" httpUri="https://zenodo.org/record/4011699/files/figure.png" pageId="3" pageNumber="775">Fig. 2</figureCitation>
|
||
). If we assume that the same relationship held true for the smaller tyrannosaurid species studied here, such locomotory limitations would not have emerged until these animals were much closer to adult size (
|
||
<figureCitation id="131B2A55350DFFB6FD2EDE34FCC29DCD" box="[712,772,368,391]" captionStart="Figure 2" captionStartId="2.[106,159,1931,1951]" captionTargetBox="[338,1229,1274,1906]" captionTargetId="figure@2.[425,1225,1264,1864]" captionTargetPageId="2" captionText="Figure 2 Logistic growth curves for Tyrannosaurus and three related tyrannosaurids.Note that the exponential stages (the regions of maximal slope) are similar in duration but differ in slope (that is, growth rates). Regression equations (mass in kg, age in years) are as follows: T. rex, mass = {5,551/[1 + e‾0.57(age ‾ 16.1)]} + 5, r 2 = 0.953; D. torosus, mass = {1,728/[1 + e‾0.44(age ‾ 12.1)]} + 5, r 2 = 0.992; G. libratus, mass = {1,234/[1 + e‾0.38(age ‾ 12.4)]} + 5, r 2 = 0.950; A. sarcophagus, mass = {1,218/[1 + e‾0.43(age ‾ 14.1)]} + 5; r 2 = 0.985." figureDoi="http://doi.org/10.5281/zenodo.4011699" httpUri="https://zenodo.org/record/4011699/files/figure.png" pageId="3" pageNumber="775">Fig. 2</figureCitation>
|
||
). Finally, a glimpse into the potential population age structure for a dinosaur is also afforded from these data. Currie
|
||
<bibRefCitation id="EFB14B21350DFFB6FD81DEE2FDA99DF9" author="Currie, P. J." box="[615,623,422,434]" journalOrPublisher="GAIA" pageId="3" pageNumber="775" pagination="271 - 277" part="15" refId="ref3877" refString="7. Currie, P. J. Possible evidence of gregarious behavior in tyrannosaurids. GAIA 15, 271 - 277 (1998)." title="Possible evidence of gregarious behavior in tyrannosaurids" type="journal article" year="1998">
|
||
<superScript id="7C559B98350DFFB6FD81DEE2FDA99DF9" attach="left" box="[615,623,422,434]" fontSize="5" pageId="3" pageNumber="775">7</superScript>
|
||
</bibRefCitation>
|
||
has described a catastrophic death assemblage consisting of eight or nine
|
||
<taxonomicName id="4C204D53350DFFB6FD1CDE80FF289DBE" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="775" phylum="Chordata" rank="species" species="sarcophagus">
|
||
<emphasis id="B954EAC2350DFFB6FD1CDE80FF289DBE" italics="true" pageId="3" pageNumber="775">A. sarcophagus</emphasis>
|
||
</taxonomicName>
|
||
specimens thought to represent an entire pack or a subset of one. On the basis of femoral lengths, the age and developmental stage of each animal can now be estimated. The group seems to have consisted of two or three older adults ~21 or more years of age, one ~17-year-old young adult, four ~12–17- year-old sub-adults that were undergoing exponential stage growth at the time of death, and one ~10-year-old juvenile that was beginning the transition to exponential stage growth. A reopening of the site has revealed at least one more specimen (RTMP (Royal Tyrrell Museum of Palaeontology) 2002.45.46) shown here to be only 2 years old (
|
||
<tableCitation id="C6A2036B350DFFB6FEFBDDB2FEA09F46" box="[285,358,758,781]" captionStart="Table 1" captionStartId="2.[106,147,175,191]" captionTargetBox="[106,1461,206,655]" captionTargetPageId="2" captionText="Table 1 Specimens, size measures, and longevity data" httpUri="http://table.plazi.org/id/DF5F6658350CFFB7FF8CDFEBFDE69CF4" pageId="3" pageNumber="775" tableUuid="DF5F6658350CFFB7FF8CDFEBFDE69CF4">Table 1</tableCitation>
|
||
). This indicates that
|
||
<taxonomicName id="4C204D53350DFFB6FDD5DDB3FD079F47" authorityName="Osborn" authorityYear="1905" box="[563,705,759,780]" class="Reptilia" family="Tyrannosauridae" genus="Albertosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="775" phylum="Chordata" rank="species" species="sarcophagus">
|
||
<emphasis id="B954EAC2350DFFB6FDD5DDB3FD079F47" box="[563,705,759,780]" italics="true" pageId="3" pageNumber="775">A. sarcophagus</emphasis>
|
||
</taxonomicName>
|
||
groups, whether temporary or permanent, might have been composed of individuals spanning the age spectrum from adolescents to very old, senescent adults, a finding consistent with trackway evidence for other theropod dinosaurs
|
||
<bibRefCitation id="EFB14B21350DFFB6FE9FDC20FE479F3B" author="Currie, P. J." box="[377,385,868,880]" journalOrPublisher="GAIA" pageId="3" pageNumber="775" pagination="271 - 277" part="15" refId="ref3877" refString="7. Currie, P. J. Possible evidence of gregarious behavior in tyrannosaurids. GAIA 15, 271 - 277 (1998)." title="Possible evidence of gregarious behavior in tyrannosaurids" type="journal article" year="1998">
|
||
<superScript id="7C559B98350DFFB6FE9FDC20FE479F3B" attach="left" box="[377,385,868,880]" fontSize="5" pageId="3" pageNumber="775">7</superScript>
|
||
</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |