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<mods:title id="32555C0383BFAC671066A2CC5115BDC0">Age and growth dynamics of Tyrannosaurus rex</mods:title>
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<mods:title id="203C9033A5C3CD8009ACCDC485B80720">Age and growth dynamics of Tyrannosaurus rex</mods:title>
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<mods:namePart id="03380C8C69CB32A2BBBC89C8B0FB660E">Horner, John R.</mods:namePart>
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<mods:namePart id="064E553CA74DFD3F56FF4C7AA610E660">Horner, John R.</mods:namePart>
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<mods:namePart id="DB8547AA7B788328704C0BB0EC3B140A">Padian, Kevin</mods:namePart>
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<treatment id="03D3878CFF96FFB2CF4BFAEA3CB057AE" ID-DOI="http://doi.org/10.5281/zenodo.3728604" ID-GBIF-Taxon="187974824" ID-Zenodo-Dep="3728604" LSID="urn:lsid:plazi:treatment:03D3878CFF96FFB2CF4BFAEA3CB057AE" httpUri="http://treatment.plazi.org/id/03D3878CFF96FFB2CF4BFAEA3CB057AE" lastPageId="4" lastPageNumber="1879" pageId="0" pageNumber="1875" scope_genus="Age">
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<subSubSection id="C3606511FF96FFB6CF4BFAEA3CBC55FA" pageId="0" pageNumber="1875" type="nomenclature">
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<paragraph id="8BC5369AFF96FFB6CF4BFAEA3CBC55FA" blockId="0.[844,1482,891,1922]" pageId="0" pageNumber="1875">
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Nearly all
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<taxonomicName id="4C7A4D19FF96FFB6CFFCFAEA3B265766" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[978,1039,1289,1310]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1875" phylum="Chordata" rank="species" species="rex">
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<taxonomicName id="4C7A4D19FF96FFB6CFFCFAEA3B265766" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[978,1039,1289,1310]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1875" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B90EEA88FF96FFB6CFFCFAEA3B265766" box="[978,1039,1289,1310]" italics="true" pageId="0" pageNumber="1875">T. rex</emphasis>
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</taxonomicName>
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specimens known are presumed on gross morphology to be adult or sub-adult. The smallest of this sample,
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@ -83,27 +83,14 @@ Padian
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</paragraph>
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<paragraph id="8BC5369AFF96FFB6CF62F80A3AE05A62" blockId="0.[844,1481,1990,2074]" pageId="0" pageNumber="1875">
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To assess age and growth dynamics in
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<taxonomicName id="4C7A4D19FF96FFB6C8E2F80A3A2C5583" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1228,1285,2025,2043]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1875" phylum="Chordata" rank="species" species="rex">
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<taxonomicName id="4C7A4D19FF96FFB6C8E2F80A3A2C5583" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1228,1285,2025,2043]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="1875" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B90EEA88FF96FFB6C8E2F80A3A2C5583" box="[1228,1285,2025,2043]" italics="true" pageId="0" pageNumber="1875">T. rex</emphasis>
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</taxonomicName>
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, we took transverse thin sections at mid-shaft of the tibia, femur and fibula (and in
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</paragraph>
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<caption id="DF056612FF97FFB7CC52FF7D3FCD5A6E" ID-DOI="http://doi.org/10.5281/zenodo.3726772" ID-Table-UUID="DF056612FF97FFB7CC52FF7D3FCD5A6E" ID-Zenodo-Dep="3726772" httpUri="http://table.plazi.org/id/DF056612FF97FFB7CC52FF7D3FCD5A6E" pageId="1" pageNumber="1876" targetBox="[256,719,171,2070]" targetIsTable="true" targetPageId="1">
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<paragraph id="8BC5369AFF97FFB7CC52FF7D3FCD5A6E" pageId="1" pageNumber="1876">
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, been Tcor calcu-;) has of mm bone (Methods bone which of. from Tnon circumference cavity for, marrow retrocalculated
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<emphasis id="B90EEA88FF97FFB7CC52FDFA3FB85052" box="[124,145,537,554]" italics="true" pageId="1" pageNumber="1876">C</emphasis>
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;) mm ((radius LAGs bone of medullary of length, number,
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<emphasis id="B90EEA88FF97FFB7CC52FCFB3FB8515F" box="[124,145,792,807]" italics="true" pageId="1" pageNumber="1876">L</emphasis>
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MR,; fibula; LAGs, fib; preserved retroc femur; LAGs Tlag). 2, § fem in without see; counted tibia explanation, tib thickness be; could sections for cortical that; thin, LAGs incremental labelled Tnon of); v (of preserved number and.; study numbers, parabolic this LAGs LAGs in actual) catalogue with iv (; analysed; with thickness thickness penultimate
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<emphasis id="B90EEA88FF97FFB7CC4EF9913F5C54F7" box="[96,117,1650,1679]" italics="true" pageId="1" pageNumber="1876">rex</emphasis>
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, MOR cortical) cortical iii the (
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<emphasis id="B90EEA88FF97FFB7CC4EF9773F5C555E" box="[96,117,1684,1830]" italics="true" pageId="1" pageNumber="1876">Tyrannosaurus</emphasis>
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; from, Tlag minus maximum are; of thickness radius) ii numbers (Specimens; total mean cortical =). i 1) (Specimen eroded: Table (total lation
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</paragraph>
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</caption>
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<paragraph id="8BC5369AFF97FFB7CF3EFF7D3C915047" blockId="1.[784,1422,158,1140]" pageId="1" pageNumber="1876">
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other long bones such as humerus, ulna and metatarsals for comparison as available) in seven specimens (see
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<tableCitation id="C6F80321FF97FFB7C891FF5E3BD652A8" box="[1215,1279,189,208]" captionStart="Table 2" captionStartId="2.[158,219,158,179]" captionTargetBox="[158,1448,290,730]" captionTargetPageId="2" captionText="Table 2. Widths of LAG intervals for each specimen and section of Tyrannosaurus rex examined. (Element identifications are keyed to table 1. Intervals are listed from the outermost cortex inward, in millimetres; numbers in parentheses indicate cortical intervals in which LAGs were obscured from view and could not be counted (see text for extrapolations). inc, incomplete LAG intervals.)" httpUri="http://table.plazi.org/id/DF056612FF94FFB4CCB0FF7D3ECA537E" pageId="1" pageNumber="1876" tableUuid="DF056612FF94FFB4CCB0FF7D3ECA537E">table 1</tableCitation>
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<tableCitation id="C6F80321FF97FFB7C891FF5E3BD652A8" box="[1215,1279,189,208]" captionStart="Table 1" captionStartId="1.[96,117,2009,2070]" captionTargetBox="[256,719,171,2070]" captionTargetPageId="1" captionText="Table 1. Specimens of Tyrannosaurus rex analysed in this study. (Specimen numbers are from the MOR, with catalogue numbers of labelled thin sections; tib, tibia; fem, femur; fib, fibula; L, length of bone (mm); C, circumference of bone (mm); Tcor, total cortical thickness; Tlag, cortical thickness with LAGs preserved; Tnon, cortical thickness without LAGs preserved; MR, medullary radius (marrow cavity from which bone has been eroded) = total radius minus cortical thickness; actual LAGs, number of LAGs that could be counted in Tlag; retroc LAGs, number of LAGs retrocalculated for Tnon. Methods of calcu- lation : (i) mean; (ii) maximum; (iii) penultimate; (iv) parabolic; and (v) incremental; for explanation see § 2.)" httpUri="http://table.plazi.org/id/DF056612FF97FFB7CC4EF83A3FCD51B6" pageId="1" pageNumber="1876" tableUuid="DF056612FF97FFB7CC4EF83A3FCD51B6">table 1</tableCitation>
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). Although no single bone infallibly records growth dynamics, the tibial midshaft has a rounder cross-section (and therefore is less subject to cortical drift than other bones); its large size and rapid growth delays secondary erosion–reconstruction cycles of the cortex that obliterate primary bone records; and its record of LAGs agrees better with those of other less altered bones (
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<bibRefCitation id="EFEB4B6BFF97FFB7C8BCFE963A7B53F0" author="Horner, J. R. & de Ricqles, A. & Padian, K." box="[1170,1362,372,392]" journalOrPublisher="Paleobiology" pageId="1" pageNumber="1876" pagination="295 - 304" part="25" refId="ref5768" refString="Horner, J. R., de Ricqles, A. & Padian, K. 1999 Variation in dinosaur skeletochronology indicators: implications for age assessment and physiology. Paleobiology 25, 295 - 304." title="Variation in dinosaur skeletochronology indicators: implications for age assessment and physiology" type="journal article" year="1999">
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Horner
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@ -118,7 +105,7 @@ We measured or estimated the length, circumference and cortical thickness of eac
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). Histological observations were made through a Nikon petrographic Microscope; LAG intervals were measured and digitized with a Nikon DS-L1 camera to a resolution of 10 цm. By extrapolating these patterns into the marrow cavity, which formerly contained cortical bone now destroyed by erosion and remodelling, we could estimate the number of lines that had been erased. We assumed that the LAGs are annual (
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<bibRefCitation id="EFEB4B6BFF97FFB7CFC5FC9F3BA451F7" author="Chinsamy, A." box="[1003,1165,892,911]" journalOrPublisher="Mod. Geol." pageId="1" pageNumber="1876" pagination="319 - 329" part="18" refId="ref5560" refString="Chinsamy, A. 1993 Bone histology and growth trajectory of the prosauropod dinosaur Massospondylus carinatus Owen. Mod. Geol. 18, 319 - 329." title="Bone histology and growth trajectory of the prosauropod dinosaur Massospondylus carinatus Owen" type="journal article" year="1993">Chinsamy 1993</bibRefCitation>
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), and that the diameter of the hatchling
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<taxonomicName id="4C7A4D19FF97FFB7CFBAFC783CE251D5" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[916,971,923,941]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1876" phylum="Chordata" rank="species" species="rex">
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<taxonomicName id="4C7A4D19FF97FFB7CFBAFC783CE251D5" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[916,971,923,941]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1876" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B90EEA88FF97FFB7CFBAFC783CE251D5" box="[916,971,923,941]" italics="true" pageId="1" pageNumber="1876">T. rex</emphasis>
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</taxonomicName>
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tibia was
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@ -296,10 +283,25 @@ All bones sampled have LAGs throughout the outer cortex and inner cortex (when n
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</taxonomicName>
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(0.17 mm) in the collections of the MOR.
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</paragraph>
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<caption id="DF056612FF97FFB7CC4EF83A3FCD51B6" ID-Table-UUID="DF056612FF97FFB7CC4EF83A3FCD51B6" box="[96,230,158,2070]" httpUri="http://table.plazi.org/id/DF056612FF97FFB7CC4EF83A3FCD51B6" pageId="1" pageNumber="1876" startId="1.[96,117,2009,2070]" targetBox="[256,719,171,2070]" targetIsTable="true" targetPageId="1">
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<paragraph id="8BC5369AFF97FFB7CC4EF83A3F5C57E8" blockId="1.[96,230,158,2070]" box="[96,117,1421,2070]" pageId="1" pageNumber="1876">
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Table 1. Specimens of
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<emphasis id="B90EEA88FF97FFB7CC4EF9773F5C555E" box="[96,117,1684,1830]" italics="true" pageId="1" pageNumber="1876">Tyrannosaurus</emphasis>
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<emphasis id="B90EEA88FF97FFB7CC4EF9913F5C54F7" box="[96,117,1650,1679]" italics="true" pageId="1" pageNumber="1876">rex</emphasis>
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analysed in this study.
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</paragraph>
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<paragraph id="8BC5369AFF97FFB7CC52F7ED3FCD51B6" blockId="1.[96,230,158,2070]" box="[124,230,158,2070]" pageId="1" pageNumber="1876">
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(Specimen numbers are from the MOR, with catalogue numbers of labelled thin sections; tib, tibia; fem, femur; fib, fibula;
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<emphasis id="B90EEA88FF97FFB7CC52FCFB3FB8515F" box="[124,145,792,807]" italics="true" pageId="1" pageNumber="1876">L</emphasis>
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||||
, length of bone (mm);
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<emphasis id="B90EEA88FF97FFB7CC52FDFA3FB85052" box="[124,145,537,554]" italics="true" pageId="1" pageNumber="1876">C</emphasis>
|
||||
, circumference of bone (mm); Tcor, total cortical thickness; Tlag, cortical thickness with LAGs preserved; Tnon, cortical thickness without LAGs preserved; MR, medullary radius (marrow cavity from which bone has been eroded) = total radius minus cortical thickness; actual LAGs, number of LAGs that could be counted in Tlag; retroc LAGs, number of LAGs retrocalculated for Tnon. Methods of calcu- lation: (i) mean; (ii) maximum; (iii) penultimate; (iv) parabolic; and (v) incremental; for explanation see § 2.)
|
||||
</paragraph>
|
||||
</caption>
|
||||
<caption id="DF056612FF94FFB4CCB0FF7D3ECA537E" ID-Table-UUID="DF056612FF94FFB4CCB0FF7D3ECA537E" httpUri="http://table.plazi.org/id/DF056612FF94FFB4CCB0FF7D3ECA537E" pageId="2" pageNumber="1877" startId="2.[158,219,158,179]" targetBox="[158,1448,290,730]" targetIsTable="true" targetPageId="2">
|
||||
<paragraph id="8BC5369AFF94FFB4CCB0FF7D3B5A52CB" blockId="2.[158,1482,158,262]" box="[158,1139,158,179]" pageId="2" pageNumber="1877">
|
||||
Table 2. Widths of LAG intervals for each specimen and section of
|
||||
<taxonomicName id="4C7A4D19FF94FFB4CF60FF7D3B2B52CB" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[846,1026,158,179]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1877" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF94FFB4CF60FF7D3B2B52CB" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[846,1026,158,179]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1877" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF94FFB4CF60FF7D3B2B52CB" box="[846,1026,158,179]" italics="true" pageId="2" pageNumber="1877">Tyrannosaurus rex</emphasis>
|
||||
</taxonomicName>
|
||||
examined.
|
||||
|
|
@ -407,7 +409,7 @@ The most completely preserved sequence of LAGs from the tibia is of
|
|||
</paragraph>
|
||||
<paragraph id="8BC5369AFF94FFB5CF4BF8053A4C57F1" blockId="2.[844,1482,837,2075]" lastBlockId="3.[784,1422,1175,2075]" lastPageId="3" lastPageNumber="1878" pageId="2" pageNumber="1877">
|
||||
If
|
||||
<taxonomicName id="4C7A4D19FF94FFB4CF55F8053C915583" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[891,952,2022,2043]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1877" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF94FFB4CF55F8053C915583" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[891,952,2022,2043]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1877" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF94FFB4CF55F8053C915583" box="[891,952,2022,2043]" italics="true" pageId="2" pageNumber="1877">T. rex</emphasis>
|
||||
</taxonomicName>
|
||||
’s growth was slowing at a consistent rate, at what age did it virtually stop growing? To extrapolate this, we applied a factor of 0.8, the inverse of the incremental factor of 1.25, to the preserved series of LAG intervals, calibrated from the last three preserved intervals. At this rate of growth decrease, within 5 years
|
||||
|
|
@ -423,7 +425,7 @@ would have deposited 0.5 mm of bone centrifugally per year, and after 8 years th
|
|||
<caption id="DF056612FF95FFB5CC4CFCAC3CBB563C" ID-DOI="http://doi.org/10.5281/zenodo.3726774" ID-Zenodo-Dep="3726774" httpUri="https://zenodo.org/record/3726774/files/figure.png" pageId="3" pageNumber="1878" startId="3.[98,166,847,868]" targetBox="[189,1336,160,827]" targetPageId="3">
|
||||
<paragraph id="8BC5369AFF95FFB5CC4CFCAC3CBB563C" blockId="3.[98,1415,847,1092]" pageId="3" pageNumber="1878">
|
||||
Figure 1. Bone histology of
|
||||
<taxonomicName id="4C7A4D19FF95FFB5CDAFFCB33D1C511D" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[385,565,848,869]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1878" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF95FFB5CDAFFCB33D1C511D" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[385,565,848,869]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1878" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF95FFB5CDAFFCB33D1C511D" box="[385,565,848,869]" italics="true" pageId="3" pageNumber="1878">Tyrannosaurus rex</emphasis>
|
||||
</taxonomicName>
|
||||
. Transverse thin sections of long bone mid-shafts. (
|
||||
|
|
@ -463,13 +465,13 @@ Castanet
|
|||
</caption>
|
||||
<paragraph id="8BC5369AFF95FFB5CF07FA723A6E5538" blockId="3.[784,1422,1175,2075]" pageId="3" pageNumber="1878">
|
||||
According to these estimates, a virtually fully grown
|
||||
<taxonomicName id="4C7A4D19FF95FFB5C95BFA713C0657BF" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1878" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF95FFB5C95BFA713C0657BF" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1878" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF95FFB5C95BFA713C0657BF" italics="true" pageId="3" pageNumber="1878">T. rex</emphasis>
|
||||
</taxonomicName>
|
||||
would have been between 15 and 18 years old. (However, as the femur of
|
||||
<materialsCitation id="3B123CC7FF95FFB5CFD2FA333B54579E" ID-GBIF-Occurrence="3352123314" box="[1020,1149,1488,1510]" collectionCode="MOR" pageId="3" pageNumber="1878" specimenCode="MOR 1152">MOR 1152</materialsCitation>
|
||||
shows, growth can drop more precipitously than an incremental estimate suggests.) We have not found in any sectioned
|
||||
<taxonomicName id="4C7A4D19FF95FFB5C885F9F33BC2545D" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1195,1259,1552,1573]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1878" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF95FFB5C885F9F33BC2545D" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1195,1259,1552,1573]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1878" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF95FFB5C885F9F33BC2545D" box="[1195,1259,1552,1573]" italics="true" pageId="3" pageNumber="1878">T. rex</emphasis>
|
||||
</taxonomicName>
|
||||
tibia or femur evidence of an outer acellular, nearly avascular layer similar to the external fundamental system (EFS) (
|
||||
|
|
@ -486,7 +488,7 @@ Horner
|
|||
)
|
||||
</taxonomicName>
|
||||
.) The absence of an EFS in preserved specimens of
|
||||
<taxonomicName id="4C7A4D19FF95FFB5C835F92F3B755499" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1051,1116,1740,1761]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1878" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF95FFB5C835F92F3B755499" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1051,1116,1740,1761]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1878" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF95FFB5C835F92F3B755499" box="[1051,1116,1740,1761]" italics="true" pageId="3" pageNumber="1878">T. rex</emphasis>
|
||||
</taxonomicName>
|
||||
could suggest that none of the animals had stopped growing, although growth was greatly slowing. Without an EFS, the near cessation of growth seems the most practical indicator of full size.
|
||||
|
|
@ -499,7 +501,7 @@ To test this hypothesis, we examined sections of a femur and fibula of
|
|||
is complete, however. Its transverse radius of 22.55 mm was measured from the perimeter to a small erosion room representing the centroid of the bone. It comprises an outer, highly compressed section of 0.44 mm, 17 LAG intervals in a section of 11.08 mm, and an inner cortical area of 11.03 mm, where the record of primary bone structure has been completely erased by secondary osteons. Using the maximum and penultimate interval retrocalculations, the inner cortex could have been deposited in 7.5–10 years. However, the LAG intervals increase centripetally at a rate of
|
||||
<emphasis id="B90EEA88FF92FFB2CEA4FE963D8853F2" box="[650,673,373,394]" italics="true" pageId="4" pageNumber="1879">ca</emphasis>
|
||||
. 1.17 mm, which incremental rate yields no more than 5 years to complete the inner cortex. According to this calculation, the age of the specimen is 22–27 years, plus the time represented by the outer, highly compressed section. Because the last three LAG intervals of the femur drop precipitously in width to less than 1 mm, we infer that the fibular LAG may represent those 3 years. The most likely estimate of the age of this fully grown
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CD41FD893E865007" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[367,431,618,639]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CD41FD893E865007" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[367,431,618,639]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF92FFB2CD41FD893E865007" box="[367,431,618,639]" italics="true" pageId="4" pageNumber="1879">T. rex</emphasis>
|
||||
</taxonomicName>
|
||||
, therefore, was 22–25 years, and full growth was reached at least 3 years before death. This estimate could be too high if the fibula grew at substantially higher rates earlier in life.
|
||||
|
|
@ -508,7 +510,7 @@ is complete, however. Its transverse radius of 22.55 mm was measured from the pe
|
|||
<caption id="DF056612FF95FFB5CC4CF89A3E5B5A62" ID-DOI="http://doi.org/10.5281/zenodo.3726776" ID-Zenodo-Dep="3726776" httpUri="https://zenodo.org/record/3726776/files/figure.png" pageId="3" pageNumber="1878" startId="3.[98,166,1913,1934]" targetBox="[127,744,1180,1907]" targetPageId="3">
|
||||
<paragraph id="8BC5369AFF95FFB5CC4CF89A3E5B5A62" blockId="3.[98,730,1913,2074]" pageId="3" pageNumber="1878">
|
||||
Figure 2. Growth trajectories of individual specimens of
|
||||
<taxonomicName id="4C7A4D19FF95FFB5CC4CF8753E3F55D3" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[98,278,1942,1963]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1878" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF95FFB5CC4CF8753E3F55D3" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[98,278,1942,1963]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1878" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF95FFB5CC4CF8753E3F55D3" box="[98,278,1942,1963]" italics="true" pageId="3" pageNumber="1878">Tyrannosaurus rex</emphasis>
|
||||
</taxonomicName>
|
||||
. These are expressed as an annual percentage of attainment of full size (cortical radius of long bones). The variation results from the fact that different bones grow at different rates. The early trajectories of bone growth are estimated (see table 1).
|
||||
|
|
@ -522,7 +524,7 @@ Figure 2. Growth trajectories of individual specimens of
|
|||
</paragraph>
|
||||
<paragraph id="8BC5369AFF92FFB2CCB0FCDC3EA356BB" blockId="4.[158,796,798,2075]" pageId="4" pageNumber="1879">
|
||||
These results have several implications for the assessment of age and growth in large dinosaurs. Not all bones provide the same signal of age. Larger or more robust specimens are not necessarily older than smaller more gracile ones. The substantial variation seen in
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CDC9FC593D0351B7" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[487,554,954,975]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CDC9FC593D0351B7" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[487,554,954,975]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF92FFB2CDC9FC593D0351B7" box="[487,554,954,975]" italics="true" pageId="4" pageNumber="1879">T. rex</emphasis>
|
||||
</taxonomicName>
|
||||
skeletons cannot yet reliably be attributed to either age, sexual dimorphism or anagenetic evolution through their known temporal range. Erosion and reconstruction of cortical bone obscure primary tissues useful in skeletochronology. Various methods can retrocalculate destroyed tissue, but none is
|
||||
|
|
@ -533,13 +535,13 @@ better than any other. Independent evidence of growth dynamics is needed from ea
|
|||
Three out of the seven specimens that we analysed appear to have effectively ceased active growth 2 or 3 years before death, although their cortical radius continued to increase annually by 0.5–0.7% (e.g.
|
||||
<materialsCitation id="3B123CC7FF92FFB2CE61FACB3DE55745" ID-GBIF-Occurrence="3352123306" box="[591,716,1319,1341]" collectionCode="MOR" pageId="4" pageNumber="1879" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||||
). Four other specimens appear to have still been growing, but LAG interval decreases suggest that each of them would have reached effectively full size in another 1 to 3 years. Evidence from the femora and tibiae suggest, therefore, that
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CCFEFA223E2657AE" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[208,271,1473,1494]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CCFEFA223E2657AE" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[208,271,1473,1494]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF92FFB2CCFEFA223E2657AE" box="[208,271,1473,1494]" italics="true" pageId="4" pageNumber="1879">T. rex</emphasis>
|
||||
</taxonomicName>
|
||||
reached full size by 16±3 years, but we caution that our sample is small and individual variation may have affected age at maturity. The fibula of
|
||||
<materialsCitation id="3B123CC7FF92FFB2CE10FA1D3D95546B" ID-GBIF-Occurrence="3352123319" box="[574,700,1533,1555]" collectionCode="MOR" pageId="4" pageNumber="1879" specimenCode="MOR 1152">MOR 1152</materialsCitation>
|
||||
suggests a higher range of 22–27 years, although the rate of its inner cortical growth may be underestimated. In any case, within about two decades
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CD95F9B93ED05417" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[443,505,1626,1647]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CD95F9B93ED05417" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[443,505,1626,1647]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF92FFB2CD95F9B93ED05417" box="[443,505,1626,1647]" italics="true" pageId="4" pageNumber="1879">T. rex</emphasis>
|
||||
</taxonomicName>
|
||||
appears to have effectively stopped growing.
|
||||
|
|
@ -560,7 +562,7 @@ Laws
|
|||
1975
|
||||
</bibRefCitation>
|
||||
). Estimates for the mass of an adult
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CCB0F8F23FF2555E" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[158,219,1809,1830]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CCB0F8F23FF2555E" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[158,219,1809,1830]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF92FFB2CCB0F8F23FF2555E" box="[158,219,1809,1830]" italics="true" pageId="4" pageNumber="1879">T. rex</emphasis>
|
||||
</taxonomicName>
|
||||
range from 5000 to 8000 kg (
|
||||
|
|
@ -574,11 +576,11 @@ Laws
|
|||
), with some outliers reported up to 7300 kg (
|
||||
<bibRefCitation id="EFEB4B6BFF92FFB2CE55F8AD3C26551C" author="Nowak, R. M." box="[635,783,1870,1892]" journalOrPublisher="Baltimore, MD: Johns Hopkins University Press" pageId="4" pageNumber="1879" refId="ref5945" refString="Nowak, R. M. 1999 Walker's mammals of the aeorld, vol. II. Baltimore, MD: Johns Hopkins University Press." title="Walker's mammals of the aeorld, vol. II" type="book" year="1999">Nowak 1999</bibRefCitation>
|
||||
), so the body masses of
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CD81F88E3EDD55FA" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[431,500,1901,1922]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CD81F88E3EDD55FA" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[431,500,1901,1922]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF92FFB2CD81F88E3EDD55FA" box="[431,500,1901,1922]" italics="true" pageId="4" pageNumber="1879">T. rex</emphasis>
|
||||
</taxonomicName>
|
||||
and African elephants at full size are similar. It follows that the dinosaur and the elephant would have grown at roughly similar rates, consistent with the well vascularized fibro-lamellar tissue that
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CCB0F8043FF75584" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[158,222,2023,2044]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7A4D19FF92FFB2CCB0F8043FF75584" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[158,222,2023,2044]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B90EEA88FF92FFB2CCB0F8043FF75584" box="[158,222,2023,2044]" italics="true" pageId="4" pageNumber="1879">T. rex</emphasis>
|
||||
</taxonomicName>
|
||||
deposited (
|
||||
|
|
@ -613,7 +615,7 @@ Sampson
|
|||
2003
|
||||
</bibRefCitation>
|
||||
). Estimates of habitat partitioning and living space requirements have been based on the assumption that tyrannosaurids were ectothermic, because ecological models based on extrapolations from much smaller living guilds do not work if tyrannosaurids were endothermic. Our study indicates that
|
||||
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<taxonomicName id="4C7A4D19FF92FFB2CF62FDA83CA25018" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[844,907,587,608]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1879" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B90EEA88FF92FFB2CF62FDA83CA25018" box="[844,907,587,608]" italics="true" pageId="4" pageNumber="1879">T. rex</emphasis>
|
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|
||||
grew quickly to adult size, and its growth dynamics suggest high basal metabolic rates to sustain this growth (
|
||||
|
|
|
|||
|
|
@ -1,60 +1,60 @@
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The large theropod
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<taxonomicName id="4C7CADB73624FF6C8639FCDFFBCCFCAF" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[815,1141,794,832]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="166" phylum="Chordata" rank="species" species="rex">
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<emphasis id="B9080A263624FF6C8639FCDFFBCCFCAF" box="[815,1141,794,832]" italics="true" pageId="0" pageNumber="166">Tyrannosaurus rex</emphasis>
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</taxonomicName>
|
||||
is the archetype carnivorous dinosaur ever since it was named in 1905 (
|
||||
|
|
@ -69,7 +69,7 @@ Even its name, “tyrant-lizard king,” invoked it as a top predator. Recent ch
|
|||
;
|
||||
<bibRefCitation id="EFEDABC53624FF6C81F2FC0EF9D2FC1E" author="Calvo, J. O. & Coria, R." box="[1252,1643,971,1009]" editor="Perez-Moreno, B. R & Holtz, T. J. & Sanz, J. L. & Moratalla, J." firstAuthor="Calvo" journalOrPublisher="Gaia: Revista de Geociencias, Museu National de Historia Natural, Lisbon" pageId="0" pageNumber="166" pagination="117 - 122" part="15" refId="ref8976" refString="Calvo, J. O., and Coria, R. 1998. Rew specimen of Giganotosaurus carolinii (Coria & Salgado, 1995), supports it as the largest theropod ever found. P. 117 - 122 in Perez-Moreno, B. R, Holtz, T. J., Sanz, J. L., and Moratalla, J. (eds.). Aspects of Theropod Paleobiology. Gaia: Revista de Geociencias, Museu National de Historia Natural, Lisbon, 15." title="Rew specimen of Giganotosaurus carolinii (Coria & Salgado, 1995), supports it as the largest theropod ever found" type="book chapter" volumeTitle="Aspects of Theropod Paleobiology" year="1998">Calvo and Coria 1998</bibRefCitation>
|
||||
) have not diminished its popularity. Osborn reasoned that
|
||||
<taxonomicName id="4C7CADB73624FF6C8026FBC0F98CFBC4" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1328,1589,1029,1067]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="166" phylum="Chordata" rank="genus">
|
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<taxonomicName id="4C7CADB73624FF6C8026FBC0F98CFBC4" authorityName="Osborn" authorityYear="1905" box="[1328,1589,1029,1067]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="166" phylum="Chordata" rank="genus">
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<emphasis id="B9080A263624FF6C8026FBC0F98CFBC4" box="[1328,1589,1029,1067]" italics="true" pageId="0" pageNumber="166">Tyrannosaurus</emphasis>
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</taxonomicName>
|
||||
was a predator on the basis of its teeth. Coprolitic material, some containing fossilized soft tissue, supports a carnivorous diet (e.g.,
|
||||
|
|
@ -86,7 +86,7 @@ was a predator on the basis of its teeth. Coprolitic material, some containing f
|
|||
Although
|
||||
<bibRefCitation id="EFEDABC53624FF6C87C1FAE9FC6EFABD" author="Osborn, H. F." box="[727,983,1324,1362]" firstAuthor="Osborn" journalOrPublisher="Bulletin of the American Museum of Natural History" pageId="0" pageNumber="166" pagination="259 - 265" part="21" refId="ref10204" refString="Osborn, H. F. 1905. Tyrannosaurus and other Cretaceous carnivorous dinosaurs. Bulletin of the American Museum of Natural History 21: 259 - 265." title="Tyrannosaurus and other Cretaceous carnivorous dinosaurs" type="journal article" year="1905">Osborn (1905</bibRefCitation>
|
||||
, 1912) was clear that he considered
|
||||
<taxonomicName id="4C7CADB73624FF6C836EFAE9FD84FA62" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="166" phylum="Chordata" rank="genus">
|
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<taxonomicName id="4C7CADB73624FF6C836EFAE9FD84FA62" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="166" phylum="Chordata" rank="genus">
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<emphasis id="B9080A263624FF6C836EFAE9FD84FA62" italics="true" pageId="0" pageNumber="166">Tyrannosaurus</emphasis>
|
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</taxonomicName>
|
||||
a predator,
|
||||
|
|
@ -96,7 +96,7 @@ dissented, considering tyrannosaurids to be scavengers instead, primarily becaus
|
|||
, when the hypothesis was reintroduced by
|
||||
<bibRefCitation id="EFEDABC53624FF6C80BCF9D2FD82F997" author="Horner, J. R. & Lessem, D." firstAuthor="Horner" journalOrPublisher="Simon & Schuster, New York" pageId="0" pageNumber="166" refId="ref9705" refString="Horner, J. R., and Lessem, D. 1993. The Complete T. rex. Simon & Schuster, New York." title="The Complete T. rex" type="book" year="1993">Horner and Lessem (1993)</bibRefCitation>
|
||||
. By and large, though,
|
||||
<taxonomicName id="4C7CADB73624FF6C86C9F997FB55F997" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[991,1260,1618,1656]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="166" phylum="Chordata" rank="genus">
|
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<taxonomicName id="4C7CADB73624FF6C86C9F997FB55F997" authorityName="Osborn" authorityYear="1905" box="[991,1260,1618,1656]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="166" phylum="Chordata" rank="genus">
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<emphasis id="B9080A263624FF6C86C9F997FB55F997" box="[991,1260,1618,1656]" italics="true" pageId="0" pageNumber="166">Tyrannosaurus</emphasis>
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</taxonomicName>
|
||||
has been considered an active predator, although the mode of attack remains controversial and includes flank bite and run (
|
||||
|
|
@ -123,7 +123,7 @@ considered head shaking as a means of flesh removal from small prey and direct r
|
|||
organ before the mouth was engaged (
|
||||
<figureCitation id="1347CAB13624FF6C8116F64CFB22F640" box="[1024,1179,2441,2479]" captionStart="Fig. 10.1" captionText="Figure 10.1. Comparison of maximum forelimb motion in 3 well-known theropods. None of the dinosaurs can reach its manus to its mouth as a result of constraints in the shoulder (see Carpenter 2002). Note that Tyrannosaurus has the greatest range or retraction. Not to scale. " figureDoi="http://doi.org/10.5281/zenodo.3942815" httpUri="https://zenodo.org/record/3942815/files/figure.png" pageId="0" pageNumber="166" targetBox="[465,2349,252,2501]" targetPageId="1">Fig. 10.1</figureCitation>
|
||||
). Does this mean, however, that the forelimbs of
|
||||
<taxonomicName id="4C7CADB73624FF6C87B3F603FC15F606" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[677,940,2502,2537]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="166" phylum="Chordata" rank="genus">
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<taxonomicName id="4C7CADB73624FF6C87B3F603FC15F606" authorityName="Osborn" authorityYear="1905" box="[677,940,2502,2537]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="166" phylum="Chordata" rank="genus">
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<emphasis id="B9080A263624FF6C87B3F603FC15F606" box="[677,940,2502,2537]" italics="true" pageId="0" pageNumber="166">Tyrannosaurus</emphasis>
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</taxonomicName>
|
||||
were as useless as portrayed? Paul (1988, p. 320) considers such the question irrelevant: “the reduced size of the forelimb shows they were not important to their owners, so they should not be important to us.” This position, supported by Lockley et al. (this volume), is based on an unsubstantiated assumption (“were not important to their owners”), which is just as useless as the untestable speculations of Lockley et al. (this volume). In point of fact,
|
||||
|
|
@ -145,7 +145,7 @@ of the dinosaurs can reach its manus to its mouth as a result of constraints in
|
|||
shoulder
|
||||
<emphasis id="B9080A263624FF6C8D5BF500F7C9F4F2" italics="true" pageId="0" pageNumber="166">(see Carpenter 2002). Note</emphasis>
|
||||
that
|
||||
<taxonomicName id="4C7CADB73624FF6C8DC4F53FF7EDF4BD" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="166" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73624FF6C8DC4F53FF7EDF4BD" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="0" pageNumber="166" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
has the greatest range or retraction.
|
||||
<emphasis id="B9080A263624FF6C82D7F45DF7BBF454" box="[1985,2050,2968,3003]" italics="true" pageId="0" pageNumber="166">Not</emphasis>
|
||||
to scale.
|
||||
|
|
@ -154,13 +154,13 @@ to scale.
|
|||
<paragraph id="8BC3D6343626FF6E812AFF2CF74AFE16" blockId="2.[997,2358,226,799]" pageId="2" pageNumber="168">
|
||||
New material has led to our reassessing the forelimb of
|
||||
<emphasis id="B9080A263626FF6E8D34FF2CF68CFEE0" box="[2082,2357,233,271]" italics="true" pageId="2" pageNumber="168">
|
||||
<taxonomicName id="4C7CADB73626FF6E8D34FF2CF697FEE0" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[2082,2350,233,271]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73626FF6E8D34FF2CF697FEE0" authorityName="Osborn" authorityYear="1905" box="[2082,2350,233,271]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
,
|
||||
</emphasis>
|
||||
and a stronger case is made for forelimb use during predation. Some of this new material displays pathologies, which is important because they are a reflection of lifestyle behaviors (
|
||||
<bibRefCitation id="EFEDABC53626FF6E8327FE5CF7FDFE50" author="Rothschild, B. M. & Martin, L. D." box="[1585,2116,409,447]" firstAuthor="Rothschild" journalOrPublisher="CRC Press, Boca Raton, FL" pageId="2" pageNumber="168" refId="ref10469" refString="Rothschild, B. M., and Martin, L. D. 1993. Paleopathology: Disease in the Fossil Record. CRC Press, Boca Raton, FL." title="Paleopathology: Disease in the Fossil Record" type="book" year="1993">Rothschild and Martin 1993</bibRefCitation>
|
||||
). As Paul has noted (this volume),
|
||||
<taxonomicName id="4C7CADB73626FF6E8076FE16F9D0FE16" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1376,1641,467,505]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73626FF6E8076FE16F9D0FE16" authorityName="Osborn" authorityYear="1905" box="[1376,1641,467,505]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
must have had a rough, active life.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
|
|
@ -202,13 +202,10 @@ The materials used in this study are as follows: scapula of
|
|||
</paragraph>
|
||||
</subSubSection>
|
||||
<subSubSection id="C36685BF3626FF7884C4FC61F9B8FB28" lastPageId="20" lastPageNumber="186" pageId="2" pageNumber="168" type="description">
|
||||
<paragraph id="8BC3D6343626FF6E84C4FC61FC31FB8F" blockId="2.[463,904,932,1120]" pageId="2" pageNumber="168">
|
||||
<heading id="D08B61583626FF6E84C4FC61FC99FBF2" bold="true" fontSize="19" level="2" pageId="2" pageNumber="168" reason="0">New Information on the Pectoral</heading>
|
||||
<heading id="D08B61583626FF6E84C6FBEFFC31FB8F" bold="true" box="[464,904,1066,1120]" centered="true" fontSize="19" level="1" pageId="2" pageNumber="168" reason="0">Girdle and Forelimb</heading>
|
||||
</paragraph>
|
||||
<paragraph id="8BC3D6343626FF6E84C4FC61FC31FB8F" blockId="2.[463,904,932,1120]" pageId="2" pageNumber="168">New Information on the Pectoral Girdle and Forelimb</paragraph>
|
||||
<paragraph id="8BC3D6343626FF6E86F5FC74F759FA9C" blockId="2.[994,2357,938,1398]" pageId="2" pageNumber="168">
|
||||
The pectoral girdle and forelimb of
|
||||
<taxonomicName id="4C7CADB73626FF6E8366FC74F8C4FC38" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1648,1917,945,983]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB73626FF6E8366FC74F8C4FC38" authorityName="Osborn" authorityYear="1905" box="[1648,1917,945,983]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A263626FF6E8366FC74F8C4FC38" box="[1648,1917,945,983]" italics="true" pageId="2" pageNumber="168">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
have been described by
|
||||
|
|
@ -226,7 +223,7 @@ as described by
|
|||
.
|
||||
<bibRefCitation id="EFEDABC53626FF6E8301FAD7F8A6FAD7" author="Brochu, C. A." box="[1559,1823,1298,1336]" firstAuthor="Brochu" journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="2" pageNumber="168" refId="ref8873" refString="Brochu, C. A. 2002. Osteology of Tyrannosaurus rex: Insights from a Nearly Complete Skeleton and High-Resolution Computed Tomographic Analysis of the Skull. Journal of Vertebrate Paleontology Memoir" title="Osteology of Tyrannosaurus rex: Insights from a Nearly Complete Skeleton and High-Resolution Computed Tomographic Analysis of the Skull" type="book" year="2002">Brochu (2002)</bibRefCitation>
|
||||
has discussed the possibility of ossified sternal plates in
|
||||
<taxonomicName id="4C7CADB73626FF6E80C7FA88F967FA9C" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1489,1758,1357,1395]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB73626FF6E80C7FA88F967FA9C" authorityName="Osborn" authorityYear="1905" box="[1489,1758,1357,1395]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A263626FF6E80C7FA88F967FA9C" box="[1489,1758,1357,1395]" italics="true" pageId="2" pageNumber="168">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
but came to no conclusion.
|
||||
|
|
@ -239,7 +236,7 @@ but came to no conclusion.
|
|||
<paragraph id="8BC3D6343626FF6E86F6F9EBF822F55D" blockId="2.[985,2354,1575,3034]" pageId="2" pageNumber="168">
|
||||
The furcula of
|
||||
<emphasis id="B9080A263626FF6E801FF9EBF9A6F9BB" box="[1289,1567,1582,1620]" italics="true" pageId="2" pageNumber="168">
|
||||
<taxonomicName id="4C7CADB73626FF6E801FF9EBF9A0F9BB" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1289,1561,1582,1620]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73626FF6E801FF9EBF9A0F9BB" authorityName="Osborn" authorityYear="1905" box="[1289,1561,1582,1620]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
,
|
||||
</emphasis>
|
||||
the presence of which was predicted by
|
||||
|
|
@ -258,7 +255,7 @@ the presence of which was predicted by
|
|||
<bibRefCitation id="EFEDABC53626FF6E8D2FF7C0F753F7C4" author="Hui, C. A." box="[2105,2282,2053,2091]" firstAuthor="Hui" journalOrPublisher="Journal of Morphology" pageId="2" pageNumber="168" pagination="284 - 293" part="251" refId="ref9732" refString="Hui, C. A. 2002. Avian furcula morphology may indicate relationships of flight requirements among birds. Journal of Morphology 251: 284 - 293." title="Avian furcula morphology may indicate relationships of flight requirements among birds" type="journal article" year="2002">Hui 2002</bibRefCitation>
|
||||
). In
|
||||
<emphasis id="B9080A263626FF6E86C8F785FB48F789" box="[990,1265,2112,2150]" italics="true" pageId="2" pageNumber="168">
|
||||
<taxonomicName id="4C7CADB73626FF6E86C8F785FB52F789" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[990,1259,2112,2150]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73626FF6E86C8F785FB52F789" authorityName="Osborn" authorityYear="1905" box="[990,1259,2112,2150]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
,
|
||||
</emphasis>
|
||||
however, the furcula is clearly designed to resist extreme lateral forces: (1) the ramfare anteroposteriorly flattened (cf.
|
||||
|
|
@ -276,7 +273,7 @@ nearly uniform throughout its length (see
|
|||
, fig. 16.8;
|
||||
<bibRefCitation id="EFEDABC53626FF6E8D4EF5D3FB15F598" author="Larson, P & Rigby, J. K." editor="Carpenter, K." firstAuthor="Larson" journalOrPublisher="Indiana University Press, Bloomington" pageId="2" pageNumber="168" pagination="247 - 255" refId="ref9889" refString="Larson, P, and Rigby, J. K. 2005. Furcula of Tyrannosaurus rex. P. 247 - 255 in Carpenter, K. (ed.). The Carnivorous Dinosaurs. Indiana University Press, Bloomington." title="Furcula of Tyrannosaurus rex" type="book" volumeTitle="The Carnivorous Dinosaurs" year="2005">Larson and Rigby 2005</bibRefCitation>
|
||||
, fig. 12.3). Therefore, the great depth of the
|
||||
<taxonomicName id="4C7CADB73626FF6E82C2F594F758F598" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[2004,2273,2641,2679]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB73626FF6E82C2F594F758F598" authorityName="Osborn" authorityYear="1905" box="[2004,2273,2641,2679]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A263626FF6E82C2F594F758F598" box="[2004,2273,2641,2679]" italics="true" pageId="2" pageNumber="168">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
furcula
|
||||
|
|
@ -285,7 +282,7 @@ unusual and is clearly adapted to resist stress.
|
|||
</paragraph>
|
||||
<paragraph id="8BC3D6343626FF6E813DF502F690F438" blockId="2.[985,2354,1575,3034]" pageId="2" pageNumber="168">
|
||||
Three of the 5 known
|
||||
<taxonomicName id="4C7CADB73626FF6E80FEF502F94CF502" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1512,1781,2759,2797]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB73626FF6E80FEF502F94CF502" authorityName="Osborn" authorityYear="1905" box="[1512,1781,2759,2797]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A263626FF6E80FEF502F94CF502" box="[1512,1781,2759,2797]" italics="true" pageId="2" pageNumber="168">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
furculae are pathologic (
|
||||
|
|
@ -313,7 +310,7 @@ F]).
|
|||
Figure
|
||||
<emphasis id="B9080A263627FF6F82D5FA91F7A8FA98" bold="true" box="[1987,2065,1364,1399]" pageId="3" pageNumber="169">10.2</emphasis>
|
||||
. Furcula of
|
||||
<taxonomicName id="4C7CADB73627FF6F825AFA4CF712FA43" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1868,2219,1417,1452]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="169" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73627FF6F825AFA4CF712FA43" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1868,2219,1417,1452]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="169" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||||
<emphasis id="B9080A263627FF6F8DA3FA4CF81FFA0E" italics="true" pageId="3" pageNumber="169">include</emphasis>
|
||||
<emphasis id="B9080A263627FF6F82B9FA7BF792FA0E" bold="true" box="[1967,2091,1470,1505]" pageId="3" pageNumber="169">several</emphasis>
|
||||
with
|
||||
|
|
@ -435,7 +432,7 @@ was used to calculate the force. The normal portion of the furcula corresponding
|
|||
</paragraph>
|
||||
<paragraph id="8BC3D6343620FF6B86C5F596F924FDC9" blockId="4.[979,2342,2635,3036]" lastBlockId="7.[462,1829,212,1200]" lastPageId="7" lastPageNumber="173" pageId="4" pageNumber="170">
|
||||
The scapula or
|
||||
<taxonomicName id="4C7CADB73620FF6881FAF596FAF4F596" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1260,1357,2643,2681]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="170" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7CADB73620FF6881FAF596FAF4F596" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1260,1357,2643,2681]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="170" phylum="Chordata" rank="species" species="rex">
|
||||
T.
|
||||
<emphasis id="B9080A263620FF688001F596FAF4F596" box="[1303,1357,2643,2681]" italics="true" pageId="4" pageNumber="170">rex</emphasis>
|
||||
</taxonomicName>
|
||||
|
|
@ -458,7 +455,7 @@ and shows a small facet for the epicleidium near the scapulocoracoid suture (
|
|||
A,
|
||||
<figureCitation id="1347CAB13623FF6B80BAFF1EF9E8FEEE" box="[1452,1617,219,257]" captionStart="Fig. 10.6" captionText="Figure 10.6. Position of the furcula relative to the scapula-coracoids as seen in a mounted skeleton of Tyrannosaurus (cast of BHI3033). Human (Neal L. Larson) for scale. " figureDoi="http://doi.org/10.5281/zenodo.3942825" httpUri="https://zenodo.org/record/3942825/files/figure.png" pageId="7" pageNumber="173" targetBox="[429,1868,1369,3017]" targetPageId="6">Fig. 10.6</figureCitation>
|
||||
), a position supported by a nearly uncrushed
|
||||
<taxonomicName id="4C7CADB73623FF6B812AFED0FAFCFED4" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1084,1349,277,315]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="173" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB73623FF6B812AFED0FAFCFED4" authorityName="Osborn" authorityYear="1905" box="[1084,1349,277,315]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="173" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A263623FF6B812AFED0FAFCFED4" box="[1084,1349,277,315]" italics="true" pageId="7" pageNumber="173">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
chest region found in situ that
|
||||
|
|
@ -477,7 +474,7 @@ of the pectoral
|
|||
<emphasis id="B9080A263621FF698DD8FB70F68BFB37" box="[2254,2354,1205,1240]" italics="true" pageId="5" pageNumber="171">girdle</emphasis>
|
||||
and forelimb of
|
||||
<emphasis id="B9080A263621FF698D98FB2FF745FAAD" italics="true" pageId="5" pageNumber="171">
|
||||
<taxonomicName id="4C7CADB73621FF698D98FB2FF7A6FAAD" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="171" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73621FF698D98FB2FF7A6FAAD" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="171" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
showing (A)
|
||||
</emphasis>
|
||||
the distribution of force from the
|
||||
|
|
@ -566,7 +563,7 @@ to
|
|||
<emphasis id="B9080A263623FF6B8DEDFECAF82FFE73" italics="true" pageId="7" pageNumber="173">the scapula-coracoids as seen in</emphasis>
|
||||
<emphasis id="B9080A263623FF6B82B5FEBCF80EFE73" bold="true" box="[1955,1975,377,412]" pageId="7" pageNumber="173">a</emphasis>
|
||||
mounted skeleton of
|
||||
<taxonomicName id="4C7CADB73623FF6B826CFE6AF728FE3D" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1914,2193,431,466]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="173" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB73623FF6B826CFE6AF728FE3D" authorityName="Osborn" authorityYear="1905" box="[1914,2193,431,466]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="7" pageNumber="173" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A263623FF6B826CFE6AF728FE3D" bold="true" box="[1914,2193,431,466]" pageId="7" pageNumber="173">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
<emphasis id="B9080A263623FF6B8D8BFE6AF710FE3D" box="[2205,2217,431,466]" italics="true" pageId="7" pageNumber="173">(</emphasis>
|
||||
|
|
@ -587,7 +584,7 @@ also shows an unusual pathology of the glenoid, which is partially collapsed as
|
|||
). The rotation is less near the acromion and greater near the glenoid, suggesting that great rotational forces were applied to the coracoid in a posteroventral direction, thereby partially collapsing the glenoid. Although the damage may have resulted from a fall onto the chest, the direction of rotation also corresponds to the vector for the M. coracobrachialis brevis ventralis (although the terminology is retained for
|
||||
<emphasis id="B9080A263623FF6B8319FB42F99BFB42" box="[1551,1570,1159,1197]" italics="true" pageId="7" pageNumber="173">“</emphasis>
|
||||
dorsal” versus “ventral” muscles, we are aware that the more vertical position of the humerus in
|
||||
<taxonomicName id="4C7CADB7362CFF648184FE82FA2CFE82" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1170,1429,327,365]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="174" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362CFF648184FE82FA2CFE82" authorityName="Osborn" authorityYear="1905" box="[1170,1429,327,365]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="8" pageNumber="174" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
indicates a need for modified terminology). It is therefore possible that the damage occurred when the individual was young and the forelimbs were pulling struggling prey toward the chest. Unfortunately, so little of the skeleton was recovered (see N. L. Larson this volume) that the extent and location of damage to other bones
|
||||
<emphasis id="B9080A26362CFF64828FFDF4F80BFDB8" box="[1945,1970,561,599]" italics="true" pageId="8" pageNumber="174">is</emphasis>
|
||||
unknown (e.g., right scapula-coracoid, humerus, gastralia). The distribution of pathologies elsewhere on the skeleton might resolve between the 2 possibilities.
|
||||
|
|
@ -698,7 +695,7 @@ and
|
|||
Figure 10.9. Muscle maps for humerus
|
||||
<emphasis id="B9080A26362DFF658D49FEC2F704FEB1" italics="true" pageId="9" pageNumber="175">
|
||||
in
|
||||
<taxonomicName id="4C7CADB7362DFF658D9BFEC2F841FEB1" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362DFF658D9BFEC2F841FEB1" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362DFF658D9BFEC2F841FEB1" bold="true" pageId="9" pageNumber="175">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
,
|
||||
|
|
@ -718,7 +715,7 @@ Muscle map based on
|
|||
A,
|
||||
<emphasis id="B9080A26362DFF658D42FDCBF7B9FD89" italics="true" pageId="9" pageNumber="175">
|
||||
F)
|
||||
<taxonomicName id="4C7CADB7362DFF658D94FDCBF843FD89" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362DFF658D94FDCBF843FD89" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
.
|
||||
</emphasis>
|
||||
Map for
|
||||
|
|
@ -729,7 +726,7 @@ Map for
|
|||
G
|
||||
<emphasis id="B9080A26362DFF658D01FDBDF79BFD74" box="[2071,2082,632,667]" italics="true" pageId="9" pageNumber="175">)</emphasis>
|
||||
and predicted for
|
||||
<taxonomicName id="4C7CADB7362DFF6582AAFD69F768FD20" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1980,2257,684,719]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362DFF6582AAFD69F768FD20" authorityName="Osborn" authorityYear="1905" box="[1980,2257,684,719]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<emphasis id="B9080A26362DFF658DC9FD69F753FD20" box="[2271,2282,684,719]" italics="true" pageId="9" pageNumber="175">(</emphasis>
|
||||
C,
|
||||
<emphasis id="B9080A26362DFF658C0FFD69F684FD20" box="[2329,2365,684,719]" italics="true" pageId="9" pageNumber="175">D)</emphasis>
|
||||
|
|
@ -746,7 +743,7 @@ and
|
|||
<emphasis id="B9080A26362DFF658DF4FC8EF850FC4C" italics="true" pageId="9" pageNumber="175">predicted</emphasis>
|
||||
for
|
||||
<emphasis id="B9080A26362DFF658D22FC45F7A9FC37" italics="true" pageId="9" pageNumber="175">
|
||||
<taxonomicName id="4C7CADB7362DFF658D22FC45F80CFC37" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362DFF658D22FC45F80CFC37" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
(E, J)
|
||||
</emphasis>
|
||||
based on deformation of
|
||||
|
|
@ -769,13 +766,13 @@ C,
|
|||
a better match for the
|
||||
<emphasis id="B9080A26362DFF658261FB31F877FAF8" box="[1911,1998,1268,1303]" italics="true" pageId="9" pageNumber="175">scars</emphasis>
|
||||
of
|
||||
<taxonomicName id="4C7CADB7362DFF658D1AFB31F698FAF8" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[2060,2337,1268,1303]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362DFF658D1AFB31F698FAF8" authorityName="Osborn" authorityYear="1905" box="[2060,2337,1268,1303]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<emphasis id="B9080A26362DFF65826CFAEDF6ABFAA4" box="[1914,2322,1320,1355]" italics="true" pageId="9" pageNumber="175">(A, F). This prediction is</emphasis>
|
||||
also
|
||||
<emphasis id="B9080A26362DFF6582D0FA98F7C7FA6F" box="[1990,2174,1373,1408]" italics="true" pageId="9" pageNumber="175">supported</emphasis>
|
||||
by the pattern of avulsion seen
|
||||
<emphasis id="B9080A26362DFF65826EFA02F80EFA05" box="[1912,1975,1479,1514]" italics="true" pageId="9" pageNumber="175">in a</emphasis>
|
||||
<taxonomicName id="4C7CADB7362DFF6582D0FA02F762FA05" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1990,2267,1479,1514]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362DFF6582D0FA02F762FA05" authorityName="Osborn" authorityYear="1905" box="[1990,2267,1479,1514]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
humerus
|
||||
<emphasis id="B9080A26362DFF6582E7FA38F86CF9BA" italics="true" pageId="9" pageNumber="175">(K, L). See Figure 10.13</emphasis>
|
||||
and text for further explanation.
|
||||
|
|
@ -823,7 +820,7 @@ noted that the incident ofavulsion
|
|||
<emphasis id="B9080A26362EFF6682B8FA82F87EFA82" box="[1966,1991,1351,1389]" italics="true" pageId="10" pageNumber="176">is</emphasis>
|
||||
highest in younger human individuals. Regardless of the cause, these pathologies support the hypothesis of forelimb use in
|
||||
<emphasis id="B9080A26362EFF6680F5FA79F956FA0D" box="[1507,1775,1468,1506]" italics="true" pageId="10" pageNumber="176">
|
||||
<taxonomicName id="4C7CADB7362EFF6680F5FA79F953FA0D" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1507,1770,1468,1506]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="176" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362EFF6680F5FA79F953FA0D" authorityName="Osborn" authorityYear="1905" box="[1507,1770,1468,1506]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="176" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
.
|
||||
</emphasis>
|
||||
</paragraph>
|
||||
|
|
@ -834,7 +831,7 @@ highest in younger human individuals. Regardless of the cause, these pathologies
|
|||
</paragraph>
|
||||
<paragraph id="8BC3D634362EFF6686C6F95BF8B4F650" blockId="10.[970,2337,1686,2675]" pageId="10" pageNumber="176">
|
||||
The incomplete manus of
|
||||
<taxonomicName id="4C7CADB7362EFF6680AEF95BF979F92B" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1464,1728,1694,1732]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="176" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362EFF6680AEF95BF979F92B" authorityName="Osborn" authorityYear="1905" box="[1464,1728,1694,1732]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="176" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
was described by
|
||||
<bibRefCitation id="EFEDABC5362EFF668D07F95BFB05F910" author="Carpenter, K. & Smith, M." editor="Tanke, D. & Carpenter, K." firstAuthor="Carpenter" journalOrPublisher="Indiana University Press, Bloomington" pageId="10" pageNumber="176" pagination="90 - 116" refId="ref9216" refString="Carpenter, K., and Smith, M. 2001. Forelimb osteology and biomechanics of Tyrannosaurus rex. P. 90 - 116 in Tanke, D., and Carpenter, K. (eds.). Mesozoic Vertebrate Life. Indiana University Press, Bloomington." title="Forelimb osteology and biomechanics of Tyrannosaurus rex" type="book chapter" volumeTitle="Mesozoic Vertebrate Life" year="2001">Carpenter and Smith (2001)</bibRefCitation>
|
||||
. New specimens,
|
||||
|
|
@ -864,7 +861,7 @@ is from a larger individual than
|
|||
</paragraph>
|
||||
<paragraph id="8BC3D634362EFF66810AF610F950F59F" blockId="10.[970,2337,1686,2675]" pageId="10" pageNumber="176">
|
||||
All of the new information on the forelimb was used to create a new view of the pectoral girdle and forelimb of
|
||||
<taxonomicName id="4C7CADB7362EFF6683C8F5CAF852F5DA" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1758,2027,2575,2613]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="176" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362EFF6683C8F5CAF852F5DA" authorityName="Osborn" authorityYear="1905" box="[1758,2027,2575,2613]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="10" pageNumber="176" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362EFF6683C8F5CAF852F5DA" box="[1758,2027,2575,2613]" italics="true" pageId="10" pageNumber="176">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
(
|
||||
|
|
@ -897,7 +894,7 @@ Distal
|
|||
</emphasis>
|
||||
of
|
||||
<emphasis id="B9080A26362FFF678D7CFF3AF7B2FE63" italics="true" pageId="11" pageNumber="177">
|
||||
<taxonomicName id="4C7CADB7362FFF678D7CFF3AF857FEB8" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="177" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362FFF678D7CFF3AF857FEB8" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="11" pageNumber="177" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
in multiple views: proximal
|
||||
</emphasis>
|
||||
or dorsal
|
||||
|
|
@ -943,7 +940,7 @@ centimeters.
|
|||
Although we have attempted to minimize our assumptions, some are required as a result of the nature of fossilized remains. We assume the following: (1) The power output of muscles of extinct tetrapods is comparable to that of extant tetrapods—that is, muscles were not weaker or stronger than they are today. (2) Scars on fossilized bone that are clearly not pathological (e.g., lack of obvious remodeling; see
|
||||
<bibRefCitation id="EFEDABC5362FFF678166F576F9C9F537" author="Rothschild, B. M. & Martin, L. D." box="[1136,1648,2738,2777]" firstAuthor="Rothschild" journalOrPublisher="CRC Press, Boca Raton, FL" pageId="11" pageNumber="177" refId="ref10469" refString="Rothschild, B. M., and Martin, L. D. 1993. Paleopathology: Disease in the Fossil Record. CRC Press, Boca Raton, FL." title="Paleopathology: Disease in the Fossil Record" type="book" year="1993">Rothschild and Martin 1993</bibRefCitation>
|
||||
) indicate insertion or origin for muscles or for ligaments (but see below). (3) Homologous origin-insertion scars, as determined by extant phylogenetic bracketing, identify each muscle (but see below). (4) The extent of the ioint surface can be determined from the smooth surfaces of the joints, which are separated from the diaphysis by a rim or abrupt textural transition, and denote the area capped by joint cartilage. (5) The movement of the joints was less than the area covered by the cartilage cap (based on dissections of birds) (Carpenter 2002). With these basic assumptions, we reanalyze the forelimb of
|
||||
<taxonomicName id="4C7CADB73629FF618366F9ACFD8AF925" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB73629FF618366F9ACFD8AF925" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A263629FF618366F9ACFD8AF925" italics="true" pageId="13" pageNumber="779">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
below.
|
||||
|
|
@ -976,7 +973,7 @@ Figure 10.12. Articulated pectoral
|
|||
and
|
||||
<emphasis id="B9080A263628FF608603F818FDBFF7D9" italics="true" pageId="12" pageNumber="178">forelimb</emphasis>
|
||||
of
|
||||
<taxonomicName id="4C7CADB73628FF608750F7D6FCE2F7D9" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[582,859,2067,2102]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="178" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73628FF608750F7D6FCE2F7D9" authorityName="Osborn" authorityYear="1905" box="[582,859,2067,2102]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="12" pageNumber="178" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<emphasis id="B9080A263628FF6084ADF78DFE60F784" box="[443,473,2120,2155]" italics="true" pageId="12" pageNumber="178">in</emphasis>
|
||||
lateral
|
||||
<emphasis id="B9080A263628FF608776F78DFDD5F784" box="[608,620,2120,2155]" italics="true" pageId="12" pageNumber="178">(</emphasis>
|
||||
|
|
@ -996,7 +993,7 @@ SC
|
|||
to
|
||||
<emphasis id="B9080A263629FF618D22F993F6B6F996" box="[2100,2319,1622,1657]" italics="true" pageId="13" pageNumber="779">approximate</emphasis>
|
||||
that of
|
||||
<taxonomicName id="4C7CADB73629FF6182F5F94EF745F941" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[2019,2300,1675,1710]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73629FF6182F5F94EF745F941" authorityName="Osborn" authorityYear="1905" box="[2019,2300,1675,1710]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
. Wavy vertical
|
||||
<emphasis id="B9080A263629FF618D45F905F718F90C" box="[2131,2209,1728,1763]" italics="true" pageId="13" pageNumber="779">lines</emphasis>
|
||||
show direction and degree of
|
||||
|
|
@ -1007,13 +1004,13 @@ of the
|
|||
<emphasis id="B9080A263629FF6182C7F851F7E2F858" box="[2001,2139,1940,1975]" italics="true" pageId="13" pageNumber="779">position</emphasis>
|
||||
and shape of various muscles on
|
||||
<emphasis id="B9080A263629FF618DF2F80FF6A2F802" box="[2276,2331,1994,2029]" italics="true" pageId="13" pageNumber="779">the</emphasis>
|
||||
<taxonomicName id="4C7CADB73629FF618277F83BF7CFF7CE" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1889,2166,2046,2081]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73629FF618277F83BF7CFF7CE" authorityName="Osborn" authorityYear="1905" box="[1889,2166,2046,2081]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<emphasis id="B9080A263629FF618D96F83BF770F7CE" box="[2176,2249,2046,2081]" italics="true" pageId="13" pageNumber="779">SC (</emphasis>
|
||||
see text
|
||||
<emphasis id="B9080A263629FF6182B6F7F1F869F7B8" box="[1952,2000,2100,2135]" italics="true" pageId="13" pageNumber="779">). (</emphasis>
|
||||
A
|
||||
<emphasis id="B9080A263629FF6182FCF7F1F84FF7B8" box="[2026,2038,2100,2135]" italics="true" pageId="13" pageNumber="779">)</emphasis>
|
||||
<taxonomicName id="4C7CADB73629FF618D12F7F1F6A0F7B8" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[2052,2329,2100,2135]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73629FF618D12F7F1F6A0F7B8" authorityName="Osborn" authorityYear="1905" box="[2052,2329,2100,2135]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<emphasis id="B9080A263629FF618248F7ACF73CF763" box="[1886,2181,2153,2188]" italics="true" pageId="13" pageNumber="779">scapula-coracoid</emphasis>
|
||||
used
|
||||
<emphasis id="B9080A263629FF618DF8F7ACF6AAF763" bold="true" box="[2286,2323,2153,2188]" pageId="13" pageNumber="779">as</emphasis>
|
||||
|
|
@ -1041,7 +1038,7 @@ morphed into
|
|||
<emphasis id="B9080A263629FF61824BF56EF82AF53F" box="[1885,1939,2731,2768]" italics="true" pageId="13" pageNumber="779">the</emphasis>
|
||||
acromion process of
|
||||
<emphasis id="B9080A263629FF61824BF525F82AF4EA" box="[1885,1939,2784,2821]" italics="true" pageId="13" pageNumber="779">the</emphasis>
|
||||
<taxonomicName id="4C7CADB73629FF6182B4F527F70EF4EA" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1954,2231,2786,2821]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73629FF6182B4F527F70EF4EA" authorityName="Osborn" authorityYear="1905" box="[1954,2231,2786,2821]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
scapula. Location of
|
||||
<emphasis id="B9080A263629FF618DD4F4D0F824F49F" italics="true" pageId="13" pageNumber="779">actual</emphasis>
|
||||
muscle scars
|
||||
|
|
@ -1126,7 +1123,7 @@ the more vertical
|
|||
<emphasis id="B9080A26362AFF6287BDF795FC8EF79C" box="[683,823,2128,2163]" italics="true" pageId="14" pageNumber="180">position</emphasis>
|
||||
of the humerus
|
||||
<emphasis id="B9080A26362AFF6287B0F740FD7DF747" box="[678,708,2181,2216]" italics="true" pageId="14" pageNumber="180">in</emphasis>
|
||||
<taxonomicName id="4C7CADB7362AFF6287C2F740FD80F731" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362AFF6287C2F740FD80F731" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362AFF6287C2F740FD80F731" italics="true" pageId="14" pageNumber="180">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
indicates a need for modified terminology.
|
||||
|
|
@ -1134,25 +1131,25 @@ indicates a need for modified terminology.
|
|||
</caption>
|
||||
<paragraph id="8BC3D6343629FF618706F91BF93CF6BF" blockId="13.[444,1809,1457,2975]" pageId="13" pageNumber="779">
|
||||
A testable method of predicting musculature patterns for the forelimb elements of
|
||||
<taxonomicName id="4C7CADB73629FF61878FF8DCFC1DF8D0" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[665,932,1817,1855]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73629FF61878FF8DCFC1DF8D0" authorityName="Osborn" authorityYear="1905" box="[665,932,1817,1855]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
is presented that uses extant phylogenetic bracketing, which is based on comparable elements of the crocodilian and bird. Previously,
|
||||
<bibRefCitation id="EFEDABC53629FF618799F84BFB36F85B" author="Carpenter, K. & Smith, M." box="[655,1167,1934,1972]" editor="Tanke, D. & Carpenter, K." firstAuthor="Carpenter" journalOrPublisher="Indiana University Press, Bloomington" pageId="13" pageNumber="779" pagination="90 - 116" refId="ref9216" refString="Carpenter, K., and Smith, M. 2001. Forelimb osteology and biomechanics of Tyrannosaurus rex. P. 90 - 116 in Tanke, D., and Carpenter, K. (eds.). Mesozoic Vertebrate Life. Indiana University Press, Bloomington." title="Forelimb osteology and biomechanics of Tyrannosaurus rex" type="book chapter" volumeTitle="Mesozoic Vertebrate Life" year="2001">Carpenter and Smith (2001)</bibRefCitation>
|
||||
had presented muscle maps for the forelimb of
|
||||
<emphasis id="B9080A263629FF618782F80CFC15F800" box="[660,940,1993,2031]" italics="true" pageId="13" pageNumber="779">
|
||||
<taxonomicName id="4C7CADB73629FF618782F80CFC1CF800" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[660,933,1993,2031]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73629FF618782F80CFC1CF800" authorityName="Osborn" authorityYear="1905" box="[660,933,1993,2031]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
,
|
||||
</emphasis>
|
||||
the results of which were partially criticized by
|
||||
<bibRefCitation id="EFEDABC53629FF6184A9F7C1FD74F7C5" author="Brochu, C. A." box="[447,717,2052,2090]" firstAuthor="Brochu" journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="13" pageNumber="779" refId="ref8873" refString="Brochu, C. A. 2002. Osteology of Tyrannosaurus rex: Insights from a Nearly Complete Skeleton and High-Resolution Computed Tomographic Analysis of the Skull. Journal of Vertebrate Paleontology Memoir" title="Osteology of Tyrannosaurus rex: Insights from a Nearly Complete Skeleton and High-Resolution Computed Tomographic Analysis of the Skull" type="book" year="2002">Brochu (2002)</bibRefCitation>
|
||||
. The forelimb map was admittedly influenced too greatly by the phylogenetic placement of
|
||||
<taxonomicName id="4C7CADB73629FF618139F7FAFA81F78A" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1071,1336,2111,2149]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73629FF618139F7FAFA81F78A" authorityName="Osborn" authorityYear="1905" box="[1071,1336,2111,2149]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
as closer to modern birds than to modern crocodilians (e.g.,
|
||||
<bibRefCitation id="EFEDABC53629FF61815FF7BFFA80F74F" author="Brochu, C. A." box="[1097,1337,2170,2208]" firstAuthor="Brochu" journalOrPublisher="Journal of Vertebrate Paleontology Memoir" pageId="13" pageNumber="779" refId="ref8873" refString="Brochu, C. A. 2002. Osteology of Tyrannosaurus rex: Insights from a Nearly Complete Skeleton and High-Resolution Computed Tomographic Analysis of the Skull. Journal of Vertebrate Paleontology Memoir" title="Osteology of Tyrannosaurus rex: Insights from a Nearly Complete Skeleton and High-Resolution Computed Tomographic Analysis of the Skull" type="book" year="2002">Brochu 2002</bibRefCitation>
|
||||
). It was believed that the muscle patterns should reflect this phylogenetic closeness, thus resulting in a loss of objectivity from the start (the loss of objectivity in theropod studies was subsequently criticized by Carpenter 2002, pp. 72-73).
|
||||
</paragraph>
|
||||
<paragraph id="8BC3D6343629FF628719F6A1F76DFCFC" blockId="13.[444,1809,1457,2975]" lastBlockId="14.[976,2348,215,3024]" lastPageId="14" lastPageNumber="180" pageId="13" pageNumber="779">
|
||||
The scapula and coracoid of
|
||||
<taxonomicName id="4C7CADB73629FF618132F6A1FA9FF665" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1060,1318,2404,2442]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73629FF618132F6A1FA9FF665" authorityName="Osborn" authorityYear="1905" box="[1060,1318,2404,2442]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
(and dinosaurs in general) are the most difficult elements on which to map muscles because few muscle scars are present and the shape of the coracoid differs; these issues hamper the application of phylogenetic bracketing. Nevertheless, the scapula and coracoid are crucial in muscular reconstructions because the muscle origin patterns affect the muscle patterns for the rest of the forelimb. Although muscle scars remain the chief means for mapping muscles, supplemental information is needed in the case of the scapula and coracoid. This supplemental information, as a testable hypothesis, is obtained by deforming the scapulocoracoids of both a crocodile
|
||||
<emphasis id="B9080A263629FF61814EF4B0FBDDF474" box="[1112,1124,2933,2971]" italics="true" pageId="13" pageNumber="779">(</emphasis>
|
||||
<taxonomicName id="4C7CADB73629FF618172F4B0FB41F474" box="[1124,1272,2933,2971]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Alligator</taxonomicName>
|
||||
|
|
@ -1164,26 +1161,26 @@ and bird
|
|||
)
|
||||
</emphasis>
|
||||
to approximate that of
|
||||
<taxonomicName id="4C7CADB7362AFF6281C6FF18FA6DFEEC" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1232,1492,221,259]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362AFF6281C6FF18FA6DFEEC" authorityName="Osborn" authorityYear="1905" box="[1232,1492,221,259]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
(
|
||||
<figureCitation id="1347CAB1362AFF6280E3FF18F915FEEC" box="[1525,1708,221,259]" captionStart="Fig. 10.13" captionText="Figure 10.13. Deformation of a crocodilian and avian scapula and coracoid (SC) to approximate that of Tyrannosaurus. Wavy vertical lines show direction and degree of morphing. This method allows for the prediction of the position and shape of various muscles on the Tyrannosaurus SC (see text). (A) Tyrannosaurus scapula-coracoid used as the end point to morphing of the (B) crocodilian and (E) avian pectoral girdles. Note that 2 possible scenarios (C, D) occur in the morphing of the crocodilian SC depending on what portions of the crocodilian scapula is morphed into the acromion process of the Tyrannosaurus scapula. Location of actual muscle scars (G). Abbreviations: bb, M. biceps brachii; c-M. costocoracoideus; cbd, M. coracobrachialis brevis dorsalis; cbv, M. coracobrachialis brevis ventralis; ce, M. coracobrachialis externus; ch, M. coracohumeralis; dc, M. deltoideus clavicularis; ds, M. deltoideus scapularis; I, M. levator scapulae; rs, M. rhomboideus superficialis; sb, M. supracoracoideus brevis; sc, M. scapulohumeralis cranialis; scd, scapulohumeralis caudalis; se, M. subscapularis externus; si, M. supracoracoideus intermedius; svc, M. subscapularis ventralis cranialis; svt, M. serratus ventralis thoracis; t, M. trapezius; tbclps, M. triceps brachii; caput longus pars scapularis; tll, M. triceps longus lateralis; tm, M. terres major. Figure (B) and terminology adapted from Meers (2003); figure (E) and terminology adapted from Yasuda (2002). Although the terminology is retained for “dorsal" versus "ventral" muscles (e.g., m.c.b. ventralis), the more vertical position of the humerus in Tyrannosaurus indicates a need for modified terminology. " figureDoi="http://doi.org/10.5281/zenodo.3942839" httpUri="https://zenodo.org/record/3942839/files/figure.png" pageId="14" pageNumber="180" targetBox="[456,2335,199,1371]" targetPageId="13">Fig. 10.13</figureCitation>
|
||||
). The technique does not involve morphing of one scapulocoracoid to another because the scapulocoracoid of
|
||||
<taxonomicName id="4C7CADB7362AFF628111FE97FAB3FE97" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1031,1290,338,376]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362AFF628111FE97FAB3FE97" authorityName="Osborn" authorityYear="1905" box="[1031,1290,338,376]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362AFF628111FE97FAB3FE97" box="[1031,1290,338,376]" italics="true" pageId="14" pageNumber="180">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
is not used as one end point. Although the technique uses a Cartesian grid, it does not attempt to explain homologous points of 2 forms in the manner used by
|
||||
<bibRefCitation id="EFEDABC5362AFF6280EDFE02F876FE02" author="Thompson, D'Arcy W." box="[1531,1999,455,493]" firstAuthor="D'Arcy Thompson" journalOrPublisher="Cambridge University Press, Cambridge" pageId="14" pageNumber="180" refId="ref10744" refString="Thompson, D'Arcy W. 1961. On Growth and Form. Cambridge University Press, Cambridge." title="On Growth and Form" type="book" year="1961">D’Arcy Thompson (1961)</bibRefCitation>
|
||||
. Instead, the technique attempts to predict the muscle origin and insertion patterns of the scapulocoracoid of
|
||||
<taxonomicName id="4C7CADB7362AFF628029FDF8F9F5FD8C" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1343,1612,573,611]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362AFF628029FDF8F9F5FD8C" authorityName="Osborn" authorityYear="1905" box="[1343,1612,573,611]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362AFF628029FDF8F9F5FD8C" box="[1343,1612,573,611]" italics="true" pageId="14" pageNumber="180">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
as it would be if the scapulocoracoid of the crocodile or bird were deformed into that of
|
||||
<taxonomicName id="4C7CADB7362AFF628264FDBDF739FD71" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1906,2176,632,670]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362AFF628264FDBDF739FD71" authorityName="Osborn" authorityYear="1905" box="[1906,2176,632,670]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
. The results can then be used to determine which of the 2 muscle patterns, crocodilian or avian,
|
||||
<emphasis id="B9080A26362AFF6281E9FD28FAA1FCFC" box="[1279,1304,749,787]" italics="true" pageId="14" pageNumber="180">is</emphasis>
|
||||
most like that seen on the bones of
|
||||
<emphasis id="B9080A26362AFF6282A8FD28F76DFCFC" box="[1982,2260,749,787]" italics="true" pageId="14" pageNumber="180">
|
||||
<taxonomicName id="4C7CADB7362AFF6282A8FD28F776FCFC" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1982,2255,749,787]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362AFF6282A8FD28F776FCFC" authorityName="Osborn" authorityYear="1905" box="[1982,2255,749,787]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
.
|
||||
</emphasis>
|
||||
</paragraph>
|
||||
|
|
@ -1198,13 +1195,13 @@ B, E). Minimum thickness of the scapular neck was selected to standardize the sc
|
|||
,
|
||||
</emphasis>
|
||||
and
|
||||
<taxonomicName id="4C7CADB7362AFF6281EFFC1DFA47FC11" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1273,1534,984,1022]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362AFF6281EFFC1DFA47FC11" authorityName="Osborn" authorityYear="1905" box="[1273,1534,984,1022]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362AFF6281EFFC1DFA47FC11" box="[1273,1534,984,1022]" italics="true" pageId="14" pageNumber="180">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
(
|
||||
<figureCitation id="1347CAB1362AFF62830CFC1DF965FC11" box="[1562,1756,984,1022]" captionStart="Fig. 10.13" captionText="Figure 10.13. Deformation of a crocodilian and avian scapula and coracoid (SC) to approximate that of Tyrannosaurus. Wavy vertical lines show direction and degree of morphing. This method allows for the prediction of the position and shape of various muscles on the Tyrannosaurus SC (see text). (A) Tyrannosaurus scapula-coracoid used as the end point to morphing of the (B) crocodilian and (E) avian pectoral girdles. Note that 2 possible scenarios (C, D) occur in the morphing of the crocodilian SC depending on what portions of the crocodilian scapula is morphed into the acromion process of the Tyrannosaurus scapula. Location of actual muscle scars (G). Abbreviations: bb, M. biceps brachii; c-M. costocoracoideus; cbd, M. coracobrachialis brevis dorsalis; cbv, M. coracobrachialis brevis ventralis; ce, M. coracobrachialis externus; ch, M. coracohumeralis; dc, M. deltoideus clavicularis; ds, M. deltoideus scapularis; I, M. levator scapulae; rs, M. rhomboideus superficialis; sb, M. supracoracoideus brevis; sc, M. scapulohumeralis cranialis; scd, scapulohumeralis caudalis; se, M. subscapularis externus; si, M. supracoracoideus intermedius; svc, M. subscapularis ventralis cranialis; svt, M. serratus ventralis thoracis; t, M. trapezius; tbclps, M. triceps brachii; caput longus pars scapularis; tll, M. triceps longus lateralis; tm, M. terres major. Figure (B) and terminology adapted from Meers (2003); figure (E) and terminology adapted from Yasuda (2002). Although the terminology is retained for “dorsal" versus "ventral" muscles (e.g., m.c.b. ventralis), the more vertical position of the humerus in Tyrannosaurus indicates a need for modified terminology. " figureDoi="http://doi.org/10.5281/zenodo.3942839" httpUri="https://zenodo.org/record/3942839/files/figure.png" pageId="14" pageNumber="180" targetBox="[456,2335,199,1371]" targetPageId="13">Figs. 10.13</figureCitation>
|
||||
A, B, E) because of the peculiar shape of the avian coracoid precluded scapula-coracoid length as the standard. The Mesh Warp feature of Corel PhotoPaint 7 was used to deform the images using a 10 by 10 grid. By manually moving each intersect ofthe gridlines (node), a small area of the scapulocoracoid surrounding each node could be deformed. The deformation was smooth, meaning that no sharp angles and lines resulted, thus approximating changes in a biological structure. Nodes were moved until the outline of the scapulocoracoid closely approximated the that of
|
||||
<taxonomicName id="4C7CADB7362AFF62807DFA6AF9C9FA3A" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1387,1648,1455,1493]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362AFF62807DFA6AF9C9FA3A" authorityName="Osborn" authorityYear="1905" box="[1387,1648,1455,1493]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
(cf.
|
||||
<figureCitation id="1347CAB1362AFF6283ABFA6AF8D0FA3A" box="[1725,1897,1455,1493]" captionStart="Fig. 10.13" captionText="Figure 10.13. Deformation of a crocodilian and avian scapula and coracoid (SC) to approximate that of Tyrannosaurus. Wavy vertical lines show direction and degree of morphing. This method allows for the prediction of the position and shape of various muscles on the Tyrannosaurus SC (see text). (A) Tyrannosaurus scapula-coracoid used as the end point to morphing of the (B) crocodilian and (E) avian pectoral girdles. Note that 2 possible scenarios (C, D) occur in the morphing of the crocodilian SC depending on what portions of the crocodilian scapula is morphed into the acromion process of the Tyrannosaurus scapula. Location of actual muscle scars (G). Abbreviations: bb, M. biceps brachii; c-M. costocoracoideus; cbd, M. coracobrachialis brevis dorsalis; cbv, M. coracobrachialis brevis ventralis; ce, M. coracobrachialis externus; ch, M. coracohumeralis; dc, M. deltoideus clavicularis; ds, M. deltoideus scapularis; I, M. levator scapulae; rs, M. rhomboideus superficialis; sb, M. supracoracoideus brevis; sc, M. scapulohumeralis cranialis; scd, scapulohumeralis caudalis; se, M. subscapularis externus; si, M. supracoracoideus intermedius; svc, M. subscapularis ventralis cranialis; svt, M. serratus ventralis thoracis; t, M. trapezius; tbclps, M. triceps brachii; caput longus pars scapularis; tll, M. triceps longus lateralis; tm, M. terres major. Figure (B) and terminology adapted from Meers (2003); figure (E) and terminology adapted from Yasuda (2002). Although the terminology is retained for “dorsal" versus "ventral" muscles (e.g., m.c.b. ventralis), the more vertical position of the humerus in Tyrannosaurus indicates a need for modified terminology. " figureDoi="http://doi.org/10.5281/zenodo.3942839" httpUri="https://zenodo.org/record/3942839/files/figure.png" pageId="14" pageNumber="180" targetBox="[456,2335,199,1371]" targetPageId="13">Fig. 10.13</figureCitation>
|
||||
A and
|
||||
|
|
@ -1221,7 +1218,7 @@ D assumes the dorsal prominence just anterior to the scapular neck of the crocod
|
|||
<emphasis id="B9080A26362AFF6282FFF88FF7BAF89F" box="[2025,2051,1866,1904]" italics="true" pageId="14" pageNumber="180">is</emphasis>
|
||||
homologous to the posterodorsal corner of the acromion in
|
||||
<emphasis id="B9080A26362AFF6283F9F840F845F844" box="[1775,2044,1925,1963]" italics="true" pageId="14" pageNumber="180">
|
||||
<taxonomicName id="4C7CADB7362AFF6283F9F840F84FF844" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1775,2038,1925,1963]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362AFF6283F9F840F84FF844" authorityName="Osborn" authorityYear="1905" box="[1775,2038,1925,1963]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
,
|
||||
</emphasis>
|
||||
whereas
|
||||
|
|
@ -1230,7 +1227,7 @@ C does not. This is tested below.
|
|||
</paragraph>
|
||||
<paragraph id="8BC3D634362AFF628130F83EFB62F4B7" blockId="14.[976,2348,215,3024]" pageId="14" pageNumber="180">
|
||||
The muscle patterns of the deformed crocodile and avian scapulocoracoids were then compared against the few muscle scars on the scapulocoracoid of
|
||||
<taxonomicName id="4C7CADB7362AFF628175F7B5FADFF779" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1123,1382,2160,2198]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362AFF628175F7B5FADFF779" authorityName="Osborn" authorityYear="1905" box="[1123,1382,2160,2198]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362AFF628175F7B5FADFF779" box="[1123,1382,2160,2198]" italics="true" pageId="14" pageNumber="180">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
(
|
||||
|
|
@ -1239,13 +1236,13 @@ The muscle patterns of the deformed crocodile and avian scapulocoracoids were th
|
|||
<figureCitation id="1347CAB1362AFF628D1AF7B5F743F779" box="[2060,2298,2160,2198]" captionStart="Fig. 10.13" captionText="Figure 10.13. Deformation of a crocodilian and avian scapula and coracoid (SC) to approximate that of Tyrannosaurus. Wavy vertical lines show direction and degree of morphing. This method allows for the prediction of the position and shape of various muscles on the Tyrannosaurus SC (see text). (A) Tyrannosaurus scapula-coracoid used as the end point to morphing of the (B) crocodilian and (E) avian pectoral girdles. Note that 2 possible scenarios (C, D) occur in the morphing of the crocodilian SC depending on what portions of the crocodilian scapula is morphed into the acromion process of the Tyrannosaurus scapula. Location of actual muscle scars (G). Abbreviations: bb, M. biceps brachii; c-M. costocoracoideus; cbd, M. coracobrachialis brevis dorsalis; cbv, M. coracobrachialis brevis ventralis; ce, M. coracobrachialis externus; ch, M. coracohumeralis; dc, M. deltoideus clavicularis; ds, M. deltoideus scapularis; I, M. levator scapulae; rs, M. rhomboideus superficialis; sb, M. supracoracoideus brevis; sc, M. scapulohumeralis cranialis; scd, scapulohumeralis caudalis; se, M. subscapularis externus; si, M. supracoracoideus intermedius; svc, M. subscapularis ventralis cranialis; svt, M. serratus ventralis thoracis; t, M. trapezius; tbclps, M. triceps brachii; caput longus pars scapularis; tll, M. triceps longus lateralis; tm, M. terres major. Figure (B) and terminology adapted from Meers (2003); figure (E) and terminology adapted from Yasuda (2002). Although the terminology is retained for “dorsal" versus "ventral" muscles (e.g., m.c.b. ventralis), the more vertical position of the humerus in Tyrannosaurus indicates a need for modified terminology. " figureDoi="http://doi.org/10.5281/zenodo.3942839" httpUri="https://zenodo.org/record/3942839/files/figure.png" pageId="14" pageNumber="180" targetBox="[456,2335,199,1371]" targetPageId="13">Figure 10.13</figureCitation>
|
||||
<emphasis id="B9080A26362AFF628DECF7B5F6A3F779" box="[2298,2330,2160,2198]" italics="true" pageId="14" pageNumber="180">G</emphasis>
|
||||
, several muscle scars on the scapulocoracoid of
|
||||
<taxonomicName id="4C7CADB7362AFF628229F76EF7FDF73E" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1855,2116,2219,2257]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362AFF628229F76EF7FDF73E" authorityName="Osborn" authorityYear="1905" box="[1855,2116,2219,2257]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362AFF628229F76EF7FDF73E" box="[1855,2116,2219,2257]" italics="true" pageId="14" pageNumber="180">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
seem to be homologous with the origins
|
||||
<emphasis id="B9080A26362AFF6280CDF723F9B7F6E3" bold="true" box="[1499,1550,2278,2316]" pageId="14" pageNumber="180">for</emphasis>
|
||||
the M. costocoracoideus, M. triceps longus lateralis, and M. supracoracoideus intermedius on the deformed crocodilian scapulocoracoid, than with any muscle origin on the scapulocoracoid of the bird. We may infer, then, that the other muscles, for which scars are not evident on the scapulocoracoid, were more homologous with those ofthe crocodile than ofthe bird. The large depression or fossa on the acromion of
|
||||
<taxonomicName id="4C7CADB7362AFF628DB8F5CEFBCBF583" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362AFF628DB8F5CEFBCBF583" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362AFF628DB8F5CEFBCBF583" italics="true" pageId="14" pageNumber="180">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
seems to better match the pattern for the M. supracoracoideus intermedius in
|
||||
|
|
@ -1254,13 +1251,13 @@ C than for the multiple muscles in this region, as seen in
|
|||
<figureCitation id="1347CAB1362AFF628185F57EFAC2F50E" box="[1171,1403,2747,2785]" captionStart="Fig. 10.13" captionText="Figure 10.13. Deformation of a crocodilian and avian scapula and coracoid (SC) to approximate that of Tyrannosaurus. Wavy vertical lines show direction and degree of morphing. This method allows for the prediction of the position and shape of various muscles on the Tyrannosaurus SC (see text). (A) Tyrannosaurus scapula-coracoid used as the end point to morphing of the (B) crocodilian and (E) avian pectoral girdles. Note that 2 possible scenarios (C, D) occur in the morphing of the crocodilian SC depending on what portions of the crocodilian scapula is morphed into the acromion process of the Tyrannosaurus scapula. Location of actual muscle scars (G). Abbreviations: bb, M. biceps brachii; c-M. costocoracoideus; cbd, M. coracobrachialis brevis dorsalis; cbv, M. coracobrachialis brevis ventralis; ce, M. coracobrachialis externus; ch, M. coracohumeralis; dc, M. deltoideus clavicularis; ds, M. deltoideus scapularis; I, M. levator scapulae; rs, M. rhomboideus superficialis; sb, M. supracoracoideus brevis; sc, M. scapulohumeralis cranialis; scd, scapulohumeralis caudalis; se, M. subscapularis externus; si, M. supracoracoideus intermedius; svc, M. subscapularis ventralis cranialis; svt, M. serratus ventralis thoracis; t, M. trapezius; tbclps, M. triceps brachii; caput longus pars scapularis; tll, M. triceps longus lateralis; tm, M. terres major. Figure (B) and terminology adapted from Meers (2003); figure (E) and terminology adapted from Yasuda (2002). Although the terminology is retained for “dorsal" versus "ventral" muscles (e.g., m.c.b. ventralis), the more vertical position of the humerus in Tyrannosaurus indicates a need for modified terminology. " figureDoi="http://doi.org/10.5281/zenodo.3942839" httpUri="https://zenodo.org/record/3942839/files/figure.png" pageId="14" pageNumber="180" targetBox="[456,2335,199,1371]" targetPageId="13">Figure 10.13</figureCitation>
|
||||
D. This suggests that the dorsal prominence of the crocodile is not homologous to the posterodorsal corner of the acromion in
|
||||
<emphasis id="B9080A26362AFF6286C6F4F7FB62F4B7" box="[976,1243,2866,2904]" italics="true" pageId="14" pageNumber="180">
|
||||
<taxonomicName id="4C7CADB7362AFF6286C6F4F7FB6CF4B7" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[976,1237,2866,2904]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362AFF6286C6F4F7FB6CF4B7" authorityName="Osborn" authorityYear="1905" box="[976,1237,2866,2904]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
.
|
||||
</emphasis>
|
||||
</paragraph>
|
||||
<paragraph id="8BC3D634362AFF638133F4AEFA55F80F" blockId="14.[976,2348,215,3024]" lastBlockId="15.[443,1810,1351,2956]" lastPageId="15" lastPageNumber="181" pageId="14" pageNumber="180">
|
||||
Some independent support for the shoulder of
|
||||
<taxonomicName id="4C7CADB7362AFF628298F4AEF72BF47E" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1934,2194,2923,2961]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362AFF628298F4AEF72BF47E" authorityName="Osborn" authorityYear="1905" box="[1934,2194,2923,2961]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362AFF628298F4AEF72BF47E" box="[1934,2194,2923,2961]" italics="true" pageId="14" pageNumber="180">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
having the crocodilian muscle pattern rather than the avian pattern is seen in the scapular blade. In a random sample of various bird skeletons (
|
||||
|
|
@ -1303,7 +1300,7 @@ margin of the
|
|||
<emphasis id="B9080A26362BFF63875FF8BAFD12F84A" box="[585,683,1919,1957]" italics="true" pageId="15" pageNumber="181">terres</emphasis>
|
||||
major and M. deltoideus scapularis. The scapula of
|
||||
<emphasis id="B9080A26362BFF638366F8BAFD75F80F" italics="true" pageId="15" pageNumber="181">
|
||||
<taxonomicName id="4C7CADB7362BFF638366F8BAFD8AF80F" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362BFF638366F8BAFD8AF80F" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
also has
|
||||
</emphasis>
|
||||
<emphasis id="B9080A26362BFF6387CFF87FFD55F80F" bold="true" box="[729,748,1978,2016]" pageId="15" pageNumber="181">a</emphasis>
|
||||
|
|
@ -1319,7 +1316,7 @@ Figure 10.14.
|
|||
muscle map for the
|
||||
<emphasis id="B9080A26362BFF638DD1FF39F86BFEBB" italics="true" pageId="15" pageNumber="181">humerus</emphasis>
|
||||
of
|
||||
<taxonomicName id="4C7CADB7362BFF638D07FEF4F69FFEBB" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[2065,2342,305,340]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362BFF638D07FEF4F69FFEBB" authorityName="Osborn" authorityYear="1905" box="[2065,2342,305,340]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
based on actual muscle scars and those inferred from
|
||||
<emphasis id="B9080A26362BFF6382D1FE0BF71AFE1E" box="[1991,2211,462,497]" italics="true" pageId="15" pageNumber="181">Figure 10.13.</emphasis>
|
||||
Some differences between
|
||||
|
|
@ -1410,7 +1407,7 @@ A and C, F and H), there are also some positional differences. For example, the
|
|||
for the M. terres major and M. latissimus dorsi is lower on
|
||||
<emphasis id="B9080A26362BFF638383F690F969F694" box="[1685,1744,2389,2427]" italics="true" pageId="15" pageNumber="181">the</emphasis>
|
||||
diaphysis and more centrally located in
|
||||
<taxonomicName id="4C7CADB7362BFF638166F655FAC0F659" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1136,1401,2448,2486]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362BFF638166F655FAC0F659" authorityName="Osborn" authorityYear="1905" box="[1136,1401,2448,2486]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362BFF638166F655FAC0F659" box="[1136,1401,2448,2486]" italics="true" pageId="15" pageNumber="181">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
(cf.
|
||||
|
|
@ -1418,7 +1415,7 @@ diaphysis and more centrally located in
|
|||
F, G, and H). Furthermore, there
|
||||
<emphasis id="B9080A26362BFF638691F60EFBDAF61E" box="[903,1123,2507,2545]" italics="true" pageId="15" pageNumber="181">seems to be</emphasis>
|
||||
a distinct scar on all
|
||||
<taxonomicName id="4C7CADB7362BFF638314F60EF8B2F61E" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1538,1803,2507,2545]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB7362BFF638314F60EF8B2F61E" authorityName="Osborn" authorityYear="1905" box="[1538,1803,2507,2545]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A26362BFF638314F60EF8B2F61E" box="[1538,1803,2507,2545]" italics="true" pageId="15" pageNumber="181">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
humeri, suggesting that the M. supracoracoideus longus had a separate
|
||||
|
|
@ -1435,7 +1432,7 @@ the deltopectoral crest than
|
|||
matches the muscle pattern
|
||||
<emphasis id="B9080A26362BFF638055F535FAD0F4F9" box="[1347,1385,2800,2838]" italics="true" pageId="15" pageNumber="181">of</emphasis>
|
||||
the crocodile than the bird. With this information, it is possible to reconstruct the muscles of the pectoral girdle and arm of
|
||||
<taxonomicName id="4C7CADB7362BFF6386A6F4A3FB01F463" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[944,1208,2918,2956]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB7362BFF6386A6F4A3FB01F463" authorityName="Osborn" authorityYear="1905" box="[944,1208,2918,2956]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
(
|
||||
<figureCitation id="1347CAB1362BFF6381C2F4A3FA31F463" box="[1236,1416,2918,2956]" captionStart="Fig. 10.15" captionText="Figure 10.15. Reconstructed of forelimb and pectoral girdle musculature in Tyrannosaurus based on results of Figures 10.13 and 10.14. Deep muscles in lateral (A) and anterior (B) views; intermediate muscles in lateral (C) and anterior (D) views; surficial muscles in lateral (E) and anterior (F) views. Scale in centimeters. Abbreviations: b, M. brachialis; bb, M. biceps brachii; cb, M. coracobrachialis brevis; cbd, M. coracobrachialis brevis dorsalis; dc, M. deltoideus davicularis; ds, M. deltoideus scapularis; hr, M. humeroradialis; Id, tendon for M. latissimus dorsi; p, M. pectoralis; sb, M. supracoracoideus brevis; sc, M. scapulohumeralis cranialis; sed, M. scapulohumeralis caudalis; sci, M. supracoracoideus intermedius; si, M. supracoracoideus longus; tbi, M. triceps brevis intermedius; tll, M. triceps longus lateralis; tm, M. terres major. Terminology adapted from Meers (2003). " figureDoi="http://doi.org/10.5281/zenodo.3942843" httpUri="https://zenodo.org/record/3942843/files/figure.png" pageId="15" pageNumber="181" targetBox="[993,2179,246,3040]" targetPageId="16">Fig. 10.15</figureCitation>
|
||||
).
|
||||
|
|
@ -1448,7 +1445,7 @@ the crocodile than the bird. With this information, it is possible to reconstruc
|
|||
of forelimb and
|
||||
<emphasis id="B9080A263634FF7C8623FEE4FD33FE96" italics="true" pageId="16" pageNumber="182">pectoral girdle</emphasis>
|
||||
musculature in
|
||||
<taxonomicName id="4C7CADB73634FF7C84EFFE4EFCB7FE41" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[505,782,395,430]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="182" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73634FF7C84EFFE4EFCB7FE41" authorityName="Osborn" authorityYear="1905" box="[505,782,395,430]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="16" pageNumber="182" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
based on results of Figures
|
||||
<emphasis id="B9080A263634FF7C84D9FE30FD9FFDF7" box="[463,550,501,536]" italics="true" pageId="16" pageNumber="182">10.13</emphasis>
|
||||
and
|
||||
|
|
@ -1521,7 +1518,7 @@ from Meers (2003).
|
|||
<paragraph id="8BC3D6343635FF7D8263FF05F6B3FE57" blockId="17.[1905,2314,192,440]" pageId="17" pageNumber="183">
|
||||
<heading id="D08B61583635FF7D8263FF05F6B3FE57" bold="true" fontSize="18" level="3" pageId="17" pageNumber="183" reason="0">
|
||||
Biomechanical Analysis of the Forelimb in
|
||||
<taxonomicName id="4C7CADB73635FF7D8262FE48F6B3FE57" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1908,2314,397,440]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="17" pageNumber="183" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7CADB73635FF7D8262FE48F6B3FE57" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1908,2314,397,440]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="17" pageNumber="183" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B9080A263635FF7D8262FE48F70DFE57" box="[1908,2228,397,440]" italics="true" pageId="17" pageNumber="183">Tyrannosaurus</emphasis>
|
||||
<emphasis id="B9080A263635FF7D8DD5FE48F6B3FE57" bold="true" box="[2243,2314,397,440]" pageId="17" pageNumber="183">rex</emphasis>
|
||||
</taxonomicName>
|
||||
|
|
@ -1531,7 +1528,7 @@ Biomechanical Analysis of the Forelimb in
|
|||
In this section, we do a reanalysis of
|
||||
<bibRefCitation id="EFEDABC53635FF7D8195FF00F936FF04" author="Carpenter, K. & Smith, M." box="[1155,1679,197,235]" editor="Tanke, D. & Carpenter, K." firstAuthor="Carpenter" journalOrPublisher="Indiana University Press, Bloomington" pageId="17" pageNumber="183" pagination="90 - 116" refId="ref9216" refString="Carpenter, K., and Smith, M. 2001. Forelimb osteology and biomechanics of Tyrannosaurus rex. P. 90 - 116 in Tanke, D., and Carpenter, K. (eds.). Mesozoic Vertebrate Life. Indiana University Press, Bloomington." title="Forelimb osteology and biomechanics of Tyrannosaurus rex" type="book chapter" volumeTitle="Mesozoic Vertebrate Life" year="2001">Carpenter and Smith (2001)</bibRefCitation>
|
||||
and an extension of the biomechanical properties of the forelimb in
|
||||
<taxonomicName id="4C7CADB73635FF7D8329FF3AFDF0FEB0" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="17" pageNumber="183" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7CADB73635FF7D8329FF3AFDF0FEB0" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="17" pageNumber="183" phylum="Chordata" rank="species" species="rex">
|
||||
<emphasis id="B9080A263635FF7D8329FF3AFDF0FEB0" italics="true" pageId="17" pageNumber="183">Tyrannosaurus rex</emphasis>
|
||||
</taxonomicName>
|
||||
based mostly on
|
||||
|
|
@ -1559,7 +1556,7 @@ obtain the MFA of the radius, we measured from the radial head to the midscar of
|
|||
</paragraph>
|
||||
<paragraph id="8BC3D6343635FF7D870FFAE5FCF8F9DD" blockId="17.[452,1825,190,2293]" pageId="17" pageNumber="183">
|
||||
A tendon tensile strength of 100 MPa, the global mean across all species, was used to estimate the tendon tensile strength in
|
||||
<taxonomicName id="4C7CADB73635FF7D8043FA9EF9E2FA6E" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1365,1627,1371,1409]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="17" pageNumber="183" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73635FF7D8043FA9EF9E2FA6E" authorityName="Osborn" authorityYear="1905" box="[1365,1627,1371,1409]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="17" pageNumber="183" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
(
|
||||
<bibRefCitation id="EFEDABC53635FF7D8364FA9EFD03FA53" author="Nigg, B. N. & Herzog, W." firstAuthor="Nigg" journalOrPublisher="J. Wiley, New York" pageId="17" pageNumber="183" refId="ref10173" refString="Nigg, B. N., and Herzog, W. 1999. Biomechanics of the Musculo-Skeletal System. 2 nd ed. J. Wiley, New York." title="Biomechanics of the Musculo-Skeletal System. 2 nd ed." type="book" year="1999">Nigg and Herzog 1999</bibRefCitation>
|
||||
). The safety factors in the values of bird tendons range from 1.19 to 4.10 (
|
||||
|
|
@ -1624,7 +1621,7 @@ surface area of the scar for insertion of the M. biceps = estimated tendon tensi
|
|||
<paragraph id="8BC3D6343635FF7D84D4F55CF965F550" blockId="17.[450,1756,2706,2751]" box="[450,1756,2713,2751]" pageId="17" pageNumber="183">where MWR is 12,211 N/3 = 4070 N, and NWR is 4070 N/3 = 1357 N</paragraph>
|
||||
<paragraph id="8BC3D6343635FF7D84D7F4C8FB28F4DC" blockId="17.[449,1169,2824,2871]" box="[449,1169,2829,2867]" pageId="17" pageNumber="183">Tendon tensile strength for the ulna is:</paragraph>
|
||||
<paragraph id="8BC3D6343635FF7D8602F4B5FA01F478" blockId="17.[788,1464,2923,2967]" box="[788,1464,2928,2967]" pageId="17" pageNumber="183">100 N/mm2 X 192 mm2 = 19,200 N</paragraph>
|
||||
<caption id="DF0386BC3636FF7E84DDFF2CFCB5FDEA" ID-Table-UUID="DF0386BC3636FF7E84DDFF2CFCB5FDEA" httpUri="http://table.plazi.org/id/DF0386BC3636FF7E84DDFF2CFCB5FDEA" pageId="18" pageNumber="184" subCaptionStartIDs="18.[467,708,387,420]" subCaptionStarts="Abbr" targetBox="[1010,2246,256,606]" targetIsTable="true" targetPageId="18">
|
||||
<caption id="DF0386BC3636FF7E84DDFF2CFCB5FDEA" ID-DOI="http://doi.org/10.5281/zenodo.13845839" ID-Table-UUID="DF0386BC3636FF7E84DDFF2CFCB5FDEA" ID-Zenodo-Dep="13845839" httpUri="http://table.plazi.org/id/DF0386BC3636FF7E84DDFF2CFCB5FDEA" pageId="18" pageNumber="184" subCaptionStartIDs="18.[467,708,387,420]" subCaptionStarts="Abbr" targetBox="[1010,2246,256,606]" targetIsTable="true" targetPageId="18">
|
||||
<paragraph id="8BC3D6343636FF7E84DDFF2CFCB5FDEA" blockId="18.[459,897,227,320]" lastBlockId="18.[463,852,383,531]" pageId="18" pageNumber="184">
|
||||
Table 10.1.
|
||||
<emphasis id="B9080A263636FF7E878DFF23FCB0FEE4" box="[667,777,230,267]" italics="true" pageId="18" pageNumber="184">Power</emphasis>
|
||||
|
|
@ -1663,9 +1660,9 @@ AnalysisMeasurements Abbreviations.—MFA, motive force arm; RFA, restive force
|
|||
<paragraph id="8BC3D6343636FF7E86F1FD7BF6B6FD0B" blockId="18.[997,2360,695,1034]" box="[999,2319,702,740]" pageId="18" pageNumber="184">where MWR is 19,200 N/3 = 6400 N, and NWR is 6400 N/3 = 2133 N.</paragraph>
|
||||
<paragraph id="8BC3D6343636FF7E8121FD3DF9C7FBE6" blockId="18.[997,2360,695,1034]" pageId="18" pageNumber="184">
|
||||
The values for the MWR and NWR represent the estimated strength of the tendon at the insertion of the M. biceps and are used as the MF in the analysis of the power of the
|
||||
<taxonomicName id="4C7CADB73636FF7E8350FCABF8E8FC7B" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1606,1873,878,916]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73636FF7E8350FCABF8E8FC7B" authorityName="Osborn" authorityYear="1905" box="[1606,1873,878,916]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
forelimbs. The following equations are used to estimate the amount of force the arm of
|
||||
<taxonomicName id="4C7CADB73636FF7E8D45FC6DFBA2FBE6" baseAuthorityName="Osborn" baseAuthorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB73636FF7E8D45FC6DFBA2FBE6" authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A263636FF7E8D45FC6DFBA2FBE6" italics="true" pageId="18" pageNumber="184">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
can resist (resistive force, or RF):
|
||||
|
|
@ -1679,7 +1676,7 @@ Measurement of the manus (177.6 mm) was taken from a cast of
|
|||
</paragraph>
|
||||
<paragraph id="8BC3D6343636FF7E8123FA30F88FF9F4" blockId="18.[996,2358,1341,1563]" box="[1077,1846,1525,1563]" pageId="18" pageNumber="184">
|
||||
MWR for the radius of
|
||||
<taxonomicName id="4C7CADB73636FF7E80F0FA30F9F2F9F4" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1510,1611,1525,1563]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7CADB73636FF7E80F0FA30F9F2F9F4" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1510,1611,1525,1563]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="species" species="rex">
|
||||
T.
|
||||
<emphasis id="B9080A263636FF7E8300FA30F9F2F9F4" box="[1558,1611,1525,1563]" italics="true" pageId="18" pageNumber="184">rex</emphasis>
|
||||
</taxonomicName>
|
||||
|
|
@ -1694,7 +1691,7 @@ is as follows:
|
|||
<paragraph id="8BC3D6343636FF7E8097F925F8C7F8E9" blockId="18.[1340,1988,1637,1801]" box="[1409,1918,1760,1798]" pageId="18" pageNumber="184">RF = 179.93 N (or 18.36 kg)</paragraph>
|
||||
<paragraph id="8BC3D6343636FF7E86F5F893F963F893" blockId="18.[995,1754,1871,1916]" box="[995,1754,1878,1916]" pageId="18" pageNumber="184">
|
||||
NWR for the radius of
|
||||
<taxonomicName id="4C7CADB73636FF7E809DF893FA56F893" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1419,1519,1878,1916]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7CADB73636FF7E809DF893FA56F893" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1419,1519,1878,1916]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="species" species="rex">
|
||||
T.
|
||||
<emphasis id="B9080A263636FF7E80ACF893FA56F893" box="[1466,1519,1878,1916]" italics="true" pageId="18" pageNumber="184">rex</emphasis>
|
||||
</taxonomicName>
|
||||
|
|
@ -1713,7 +1710,7 @@ RF
|
|||
<paragraph id="8BC3D6343636FF7E8086F784F8C8F788" blockId="18.[1350,1981,1991,2151]" box="[1424,1905,2113,2151]" pageId="18" pageNumber="184">RF = 60.01 N (or 6.12 kg)</paragraph>
|
||||
<paragraph id="8BC3D6343636FF7E86F7F772F97CF732" blockId="18.[993,1733,2224,2269]" box="[993,1733,2231,2269]" pageId="18" pageNumber="184">
|
||||
MWR for the ulna of
|
||||
<taxonomicName id="4C7CADB73636FF7E8060F772FA63F732" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1398,1498,2231,2269]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7CADB73636FF7E8060F772FA63F732" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1398,1498,2231,2269]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="species" species="rex">
|
||||
T.
|
||||
<emphasis id="B9080A263636FF7E80B3F772FA63F732" box="[1445,1498,2231,2269]" italics="true" pageId="18" pageNumber="184">rex</emphasis>
|
||||
</taxonomicName>
|
||||
|
|
@ -1728,7 +1725,7 @@ is as follows:
|
|||
<paragraph id="8BC3D6343636FF7E8096F664F8C7F628" blockId="18.[1330,1995,2344,2504]" box="[1408,1918,2465,2503]" pageId="18" pageNumber="184">F = 804.83 N (or 82.13 kg)</paragraph>
|
||||
<paragraph id="8BC3D6343636FF7E86F7F5D2F905F5D2" blockId="18.[993,1724,2577,2621]" box="[993,1724,2583,2621]" pageId="18" pageNumber="184">
|
||||
NWR for the ulna of
|
||||
<taxonomicName id="4C7CADB73636FF7E807BF5D2FA69F5D2" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1389,1488,2583,2621]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="species" species="rex">
|
||||
<taxonomicName id="4C7CADB73636FF7E807BF5D2FA69F5D2" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1389,1488,2583,2621]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="18" pageNumber="184" phylum="Chordata" rank="species" species="rex">
|
||||
T.
|
||||
<emphasis id="B9080A263636FF7E808BF5D2FA69F5D2" box="[1437,1488,2583,2621]" italics="true" pageId="18" pageNumber="184">rex</emphasis>
|
||||
</taxonomicName>
|
||||
|
|
@ -1752,11 +1749,11 @@ Adding the resistive forces of the radius and ulna results in 984.76 N (100.49 k
|
|||
</paragraph>
|
||||
<paragraph id="8BC3D6343637FF7F8738FEA9F94EFBF7" blockId="19.[474,1840,181,1051]" pageId="19" pageNumber="185">
|
||||
An average strength of 5 kg/cm2 per cross-sectional area of muscle was used to determine the cross-sectional area of the M. biceps in
|
||||
<taxonomicName id="4C7CADB73637FF7F833DFE63F895FE23" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1579,1836,422,460]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="19" pageNumber="185" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73637FF7F833DFE63F895FE23" authorityName="Osborn" authorityYear="1905" box="[1579,1836,422,460]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="19" pageNumber="185" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
(
|
||||
<bibRefCitation id="EFEDABC53637FF7F84FDFE24FC71FDE8" author="Carpenter, K. & Smith, M." box="[491,968,481,519]" editor="Tanke, D. & Carpenter, K." firstAuthor="Carpenter" journalOrPublisher="Indiana University Press, Bloomington" pageId="19" pageNumber="185" pagination="90 - 116" refId="ref9216" refString="Carpenter, K., and Smith, M. 2001. Forelimb osteology and biomechanics of Tyrannosaurus rex. P. 90 - 116 in Tanke, D., and Carpenter, K. (eds.). Mesozoic Vertebrate Life. Indiana University Press, Bloomington." title="Forelimb osteology and biomechanics of Tyrannosaurus rex" type="book chapter" volumeTitle="Mesozoic Vertebrate Life" year="2001">Carpenter and Smith 2001</bibRefCitation>
|
||||
). The NWR ofthe tendon tensile strength for the radius and ulna were added together to get the MF: 1357 N + 2133 N = 4490 N (356.12 kg). The formula used to determine the cross-sectional area of muscle is MF (kg)/strength (kg x cm-2) = cross-sectional area (cm2). Thus, the estimated cross section of the
|
||||
<taxonomicName id="4C7CADB73637FF7F812DFD09FA87FD1D" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1083,1342,716,754]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="19" pageNumber="185" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73637FF7F812DFD09FA87FD1D" authorityName="Osborn" authorityYear="1905" box="[1083,1342,716,754]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="19" pageNumber="185" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
M. biceps is 356.12 kg/5 kg x cm2 = 71.224 cm2. This translates into a diameter of9.52 cm. Of
|
||||
<emphasis id="B9080A263637FF7F8371FCC3F8A5FCC3" box="[1639,1820,774,812]" italics="true" pageId="19" pageNumber="185">course the</emphasis>
|
||||
M. biceps is not the only arm protractor. In fact, by using half the estimated cross-sectional area of the upper arm (based on a diameter of 25 cm), the amount offorce generated
|
||||
|
|
@ -1768,7 +1765,7 @@ estimated to have been around 1150 kg, or 11,270 N. Of this, the biceps generate
|
|||
Mechanical
|
||||
<emphasis id="B9080A263637FF7F8D93FB6DF68FFB3C" box="[2181,2358,1192,1235]" italics="true" pageId="19" pageNumber="185">Analysis</emphasis>
|
||||
of the Forelimb in
|
||||
<taxonomicName id="4C7CADB73637FF7F8269FAEBF6AAFAB6" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1919,2323,1326,1369]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="19" pageNumber="185" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73637FF7F8269FAEBF6AAFAB6" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1919,2323,1326,1369]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="19" pageNumber="185" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
|
||||
</heading>
|
||||
</paragraph>
|
||||
<paragraph id="8BC3D6343637FF7F84C0FB6CFA26FA1A" blockId="19.[469,1836,1186,2410]" pageId="19" pageNumber="185">
|
||||
|
|
@ -1783,15 +1780,15 @@ The mechanical advantage is the amount of force a given effort can produce. It c
|
|||
</paragraph>
|
||||
<paragraph id="8BC3D6343637FF7F873FF933FBE4F7E8" blockId="19.[469,1836,1186,2410]" pageId="19" pageNumber="185">
|
||||
The
|
||||
<taxonomicName id="4C7CADB73637FF7F8794F933FC36F8F3" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[642,911,1782,1820]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="19" pageNumber="185" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73637FF7F8794F933FC36F8F3" authorityName="Osborn" authorityYear="1905" box="[642,911,1782,1820]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="19" pageNumber="185" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
forelimb is found to have a mechanical advantage of the 0.09 (RFA measurement including the hand) and 0.18 (RFA measurement excluding the hand). The mechanical advantage of a human forearm is 0.07 (RFA measurement including the hand) and 0.13 (RFA measurement excluding the hand).
|
||||
</paragraph>
|
||||
<paragraph id="8BC3D6343637FF7F873FF7D9F8A5F688" blockId="19.[469,1836,1186,2410]" pageId="19" pageNumber="185">
|
||||
Next we evaluate the force at the elbow joint. The sum of the MFs (NWR + MWR) at the radius (138.3 kg) and the ulna (217.5 kg) minus the RF at the manus (33.5 kg) must equal the force at the elbow for a static configuration. Therefore,
|
||||
<taxonomicName id="4C7CADB73637FF7F8667F709FBCCF71D" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[881,1141,2252,2290]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="19" pageNumber="185" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73637FF7F8667F709FBCCF71D" authorityName="Osborn" authorityYear="1905" box="[881,1141,2252,2290]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="19" pageNumber="185" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
has a force of 138.3 + 217.5 - 33.5 = 322.3 kg at the elbow joint for the NWR. This compares with a force of about 128.25 kg at the elbow joint of an average adult male human for the NWR.
|
||||
</paragraph>
|
||||
<caption id="DF0386BC3637FF7F8262F58FF860F45E" ID-Table-UUID="DF0386BC3637FF7F8262F58FF860F45E" httpUri="http://table.plazi.org/id/DF0386BC3637FF7F8262F58FF860F45E" pageId="19" pageNumber="185" subCaptionStartIDs="19.[1908,2147,2766,2799]" subCaptionStarts="Abbr" targetBox="[478,1752,2662,2945]" targetIsTable="true" targetPageId="19">
|
||||
<caption id="DF0386BC3637FF7F8262F58FF860F45E" ID-DOI="http://doi.org/10.5281/zenodo.13845841" ID-Table-UUID="DF0386BC3637FF7F8262F58FF860F45E" ID-Zenodo-Dep="13845841" httpUri="http://table.plazi.org/id/DF0386BC3637FF7F8262F58FF860F45E" pageId="19" pageNumber="185" subCaptionStartIDs="19.[1908,2147,2766,2799]" subCaptionStarts="Abbr" targetBox="[478,1752,2662,2945]" targetIsTable="true" targetPageId="19">
|
||||
<paragraph id="8BC3D6343637FF7F8262F58FF860F45E" blockId="19.[1908,2344,2628,2716]" lastBlockId="19.[1908,2340,2763,2995]" pageId="19" pageNumber="185">
|
||||
Table 10.2.
|
||||
<emphasis id="B9080A263637FF7F8D56F58FF824F574" italics="true" pageId="19" pageNumber="185">Power Analysis</emphasis>
|
||||
|
|
@ -1806,25 +1803,25 @@ working range.
|
|||
</caption>
|
||||
<paragraph id="8BC3D6343637FF7F81FEF5A3F9C0F46F" pageId="19" pageNumber="185">
|
||||
<table id="F97C24943637009384C8F5A3F961F46E" box="[478,1752,2662,2945]" gridcols="4" gridrows="4" pageId="19" pageNumber="185">
|
||||
<tr id="354CD4763637009384C8F5A3F961F562" box="[478,1752,2662,2701]" gridrow="0" pageId="19" pageNumber="185" rowspan-0="1" rowspan-1="1" rowspan-3="1">
|
||||
<th id="769DBD0A3637009381B8F5A3FAD8F562" box="[1198,1377,2662,2701]" gridcol="2" gridrow="0" pageId="19" pageNumber="185">RF (kg)</th>
|
||||
<tr id="354CD4763637009384C8F5A3F961F562" box="[478,1752,2662,2701]" gridrow="0" pageId="19" pageNumber="185" rowspan-0="1">
|
||||
<th id="769DBD0A363700938624F5A3F961F562" box="[818,1752,2662,2701]" colspan="3" colspanRight="2" gridcol="1" gridrow="0" pageId="19" pageNumber="185">RF (kg)</th>
|
||||
</tr>
|
||||
<tr id="354CD4763637009384C8F57FF961F50D" box="[478,1752,2746,2786]" gridrow="1" pageId="19" pageNumber="185">
|
||||
<th id="769DBD0A3637009384C8F57FFDEBF50D" box="[478,594,2746,2786]" gridcol="0" gridrow="1" pageId="19" pageNumber="185">Range</th>
|
||||
<td id="769DBD0A363700938624F57FFC12F50D" box="[818,939,2746,2786]" gridcol="1" gridrow="1" pageId="19" pageNumber="185">Radius</td>
|
||||
<td id="769DBD0A3637009381B8F57FFAD8F50D" box="[1198,1377,2746,2786]" gridcol="2" gridrow="1" pageId="19" pageNumber="185">Ulna</td>
|
||||
<td id="769DBD0A3637009381B8F57FFABEF50D" box="[1198,1287,2746,2786]" gridcol="2" gridrow="1" pageId="19" pageNumber="185">Ulna</td>
|
||||
<td id="769DBD0A36370093830AF57FF961F50D" box="[1564,1752,2746,2786]" gridcol="3" gridrow="1" pageId="19" pageNumber="185">Combined</td>
|
||||
</tr>
|
||||
<tr id="354CD4763637009384C8F4D6F961F4DA" box="[478,1752,2835,2869]" gridrow="2" pageId="19" pageNumber="185">
|
||||
<th id="769DBD0A3637009384C8F4D6FDEBF4DA" box="[478,594,2835,2869]" gridcol="0" gridrow="2" pageId="19" pageNumber="185">MWR</th>
|
||||
<td id="769DBD0A363700938624F4D6FC12F4DA" box="[818,939,2835,2869]" gridcol="1" gridrow="2" pageId="19" pageNumber="185">18.36</td>
|
||||
<td id="769DBD0A3637009381B8F4D6FAD8F4DA" box="[1198,1377,2835,2869]" gridcol="2" gridrow="2" pageId="19" pageNumber="185">82.13</td>
|
||||
<td id="769DBD0A3637009381B8F4D6FABEF4DA" box="[1198,1287,2835,2869]" gridcol="2" gridrow="2" pageId="19" pageNumber="185">82.13</td>
|
||||
<td id="769DBD0A36370093830AF4D6F961F4DA" box="[1564,1752,2835,2869]" gridcol="3" gridrow="2" pageId="19" pageNumber="185">100.49</td>
|
||||
</tr>
|
||||
<tr id="354CD4763637009384C8F499F961F46E" box="[478,1752,2908,2945]" gridrow="3" pageId="19" pageNumber="185">
|
||||
<th id="769DBD0A3637009384C8F499FDEBF46E" box="[478,594,2908,2945]" gridcol="0" gridrow="3" pageId="19" pageNumber="185">NWR</th>
|
||||
<td id="769DBD0A363700938624F499FC12F46E" box="[818,939,2908,2945]" gridcol="1" gridrow="3" pageId="19" pageNumber="185">6.12</td>
|
||||
<td id="769DBD0A3637009381B8F499FAD8F46E" box="[1198,1377,2908,2945]" gridcol="2" gridrow="3" pageId="19" pageNumber="185">
|
||||
<td id="769DBD0A3637009381B8F499FABEF46E" box="[1198,1287,2908,2945]" gridcol="2" gridrow="3" pageId="19" pageNumber="185">
|
||||
<emphasis id="B9080A263637FF7F81B8F499FABEF46E" box="[1198,1287,2908,2945]" italics="true" pageId="19" pageNumber="185">27.37</emphasis>
|
||||
</td>
|
||||
<td id="769DBD0A36370093830AF499F961F46E" box="[1564,1752,2908,2945]" gridcol="3" gridrow="3" pageId="19" pageNumber="185">33.49</td>
|
||||
|
|
@ -1838,7 +1835,7 @@ working range.
|
|||
From the torque ot the
|
||||
<emphasis id="B9080A263630FF788062FF33F9BAFEF3" box="[1396,1539,246,284]" italics="true" pageId="20" pageNumber="186">Forearm</emphasis>
|
||||
, the force that could be applied at the manus and the resultant Force at the elbow joint were determined. We now estimate the acceleration that could be generated at the claws using the moment of inertia. The fleshed-out version ot the arm of
|
||||
<taxonomicName id="4C7CADB73630FF788234FE63F792FE23" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1826,2091,422,460]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="20" pageNumber="186" phylum="Chordata" rank="genus">
|
||||
<taxonomicName id="4C7CADB73630FF788234FE63F792FE23" authorityName="Osborn" authorityYear="1905" box="[1826,2091,422,460]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="20" pageNumber="186" phylum="Chordata" rank="genus">
|
||||
<emphasis id="B9080A263630FF788234FE63F792FE23" box="[1826,2091,422,460]" italics="true" pageId="20" pageNumber="186">Tyrannosaurus</emphasis>
|
||||
</taxonomicName>
|
||||
(
|
||||
|
|
@ -1860,15 +1857,15 @@ elliptical cylinders. The cross sections of each elliptical cylinder were determ
|
|||
</paragraph>
|
||||
<paragraph id="8BC3D6343630FF7886D9FA9DF802F8F5" blockId="20.[974,2337,1362,2056]" pageId="20" pageNumber="186">
|
||||
As we have shown, the forearm of
|
||||
<taxonomicName id="4C7CADB73630FF788344FA9DF8E5FA91" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1618,1884,1368,1406]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="20" pageNumber="186" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73630FF788344FA9DF8E5FA91" authorityName="Osborn" authorityYear="1905" box="[1618,1884,1368,1406]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="20" pageNumber="186" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
was capable of resisting large forces and moving at high accelerations. These results strengthen the hypothesis that the forelimbs were used during predation. However, because of the small size of the forelimb relative to the body size, it is unlikely that the
|
||||
<taxonomicName id="4C7CADB73630FF788171F981FAC8F985" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1127,1393,1604,1642]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="20" pageNumber="186" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73630FF788171F981FAC8F985" authorityName="Osborn" authorityYear="1905" box="[1127,1393,1604,1642]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="20" pageNumber="186" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
would use the manus for striking prey, as discussed in Carpenter (2002). Rather, the forelimbs may have been used to cling to prey. Our results of finding large forces at the elbow joint and possible signs of injury at the furcula further support this hypothesis.
|
||||
</paragraph>
|
||||
<paragraph id="8BC3D6343630FF798134F8EAF9A2F9D0" blockId="20.[974,2337,1362,2056]" lastBlockId="21.[471,1836,1319,1605]" lastPageId="21" lastPageNumber="187" pageId="20" pageNumber="186">
|
||||
Finally, in contrast to the belief of Lockley et al. (this volume) that “no useful function is plausible” to explain the forelimb of
|
||||
<emphasis id="B9080A263630FF7882ADF8AFF777F87F" box="[1979,2254,1898,1936]" italics="true" pageId="20" pageNumber="186">
|
||||
<taxonomicName id="4C7CADB73630FF7882ADF8AFF771F87F" baseAuthorityName="Osborn" baseAuthorityYear="1905" box="[1979,2248,1898,1936]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="20" pageNumber="186" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
<taxonomicName id="4C7CADB73630FF7882ADF8AFF771F87F" authorityName="Osborn" authorityYear="1905" box="[1979,2248,1898,1936]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="20" pageNumber="186" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
||||
,
|
||||
</emphasis>
|
||||
our results support the previous assertion that the forelimb played a functional role in predation. By implication, the short forelimbs of other tyrannosaurids had a similar function. In support of this, we note that a progressive reduction in the forelimb does not occur in the
|
||||
|
|
|
|||
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