1882 lines
262 KiB
XML
1882 lines
262 KiB
XML
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<mods:title id="666A00496E8FA362CF089190FB02A05D">Looking again at the forelimb of Tyrannosaurus rex</mods:title>
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<mods:namePart id="A1E3A2900825C484389B90DCED592545">Christine Lipkin</mods:namePart>
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<mods:namePart id="73F0D6FEC1245FAC2768DC635A5DEA39">Kenneth Carpenter</mods:namePart>
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<mods:dateIssued id="82EC9B3287D5F270EFD0CB5DB7B3479F">2008</mods:dateIssued>
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<mods:namePart id="345015DA44EEC4A6A501A2B6C0F82D54">Peter Larson</mods:namePart>
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<mods:namePart id="5E412BCA620BB6DA39E01E8362567F2D">Kenneth Carpenter</mods:namePart>
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<mods:title id="3D69E58962FEC61CD4551047B9724357">Tyrannosaurus rex, the tyrant king</mods:title>
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<treatment id="03D567223624FF7984DEFCDFF9A2F9D0" ID-DOI="http://doi.org/10.5281/zenodo.4710609" ID-GBIF-Taxon="165074824" ID-Zenodo-Dep="4710609" LSID="urn:lsid:plazi:treatment:03D567223624FF7984DEFCDFF9A2F9D0" httpUri="http://treatment.plazi.org/id/03D567223624FF7984DEFCDFF9A2F9D0" lastPageId="21" lastPageNumber="186" pageId="0" pageNumber="166">
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<subSubSection id="C36685BF3624FF6C84DEFCDFFAB4FC94" pageId="0" pageNumber="166" type="nomenclature">
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<paragraph id="8BC3D6343624FF6C84DEFCDFFAB4FC94" blockId="0.[450,1819,787,3011]" pageId="0" pageNumber="166">
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The large theropod
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<taxonomicName id="4C7CADB73624FF6C8639FCDFFBCCFCAF" baseAuthorityName="Osborn" baseAuthorityYear="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>
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is the archetype carnivorous dinosaur ever since it was named in 1905 (
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<bibRefCitation id="EFEDABC53624FF6C8104FC90FB45FC94" author="Osborn, H. F." box="[1042,1276,853,891]" 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>
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).
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</paragraph>
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</subSubSection>
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<subSubSection id="C36685BF3624FF6C800CFC90F941FAF9" pageId="0" pageNumber="166" type="discussion">
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<paragraph id="8BC3D6343624FF6C800CFC90F941FAF9" blockId="0.[450,1819,787,3011]" pageId="0" pageNumber="166">
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Even its name, “tyrant-lizard king,” invoked it as a top predator. Recent challenges to its title as the largest terrestrial carnivore (e.g.,
|
||
<bibRefCitation id="EFEDABC53624FF6C8687FC0EFB6BFC1E" author="Sereno, P. C. & Durheil. D. B. & Iarochene, M. & Larsson, H. C. E. & Lyon, C. H. & Mugwene, P. M. & Sidor, C. A. & Varricchio, D. J. & Wilson, J. A." box="[913,1234,971,1009]" etAl="et al." firstAuthor="Sereno" journalOrPublisher="Science" pageId="0" pageNumber="166" pagination="986 - 991" part="272" refId="ref10587" refString="Sereno, P. C., Durheil. D. B., Iarochene, M., Larsson, H. C. E., Lyon, C. H., Mugwene, P. M., Sidor, C. A., Varricchio, D. J., and Wilson, J. A. (1996). Predators dinosaurs from the Sahara and Late Cretaceous faunal differentiation. Science 272: 986 - 991." title="Predators dinosaurs from the Sahara and Late Cretaceous faunal differentiation" type="journal article" year="1996">Sereno et al. 1996</bibRefCitation>
|
||
;
|
||
<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>
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||
) 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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<emphasis id="B9080A263624FF6C8026FBC0F98CFBC4" box="[1328,1589,1029,1067]" italics="true" pageId="0" pageNumber="166">Tyrannosaurus</emphasis>
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||
</taxonomicName>
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||
was a predator on the basis of its teeth. Coprolitic material, some containing fossilized soft tissue, supports a carnivorous diet (e.g.,
|
||
<bibRefCitation id="EFEDABC53624FF6C81C9FBBEF9BDFB4E" author="Chin, K. & Eberth, D. A. & Schweitzer, M. H. & Rando, T. A. & Sloboda, W. J. & Horner, J. R." box="[1247,1540,1147,1185]" etAl="et al." firstAuthor="Chin" journalOrPublisher="Palaios" pageId="0" pageNumber="166" pagination="286 - 294" part="18" refId="ref9267" refString="Chin, K., Eberth, D. A., Schweitzer, M. H., Rando, T. A., Sloboda, W. J., and Horner, J. R. 2003. Remarkable preservation of undigested muscle tissue within a Late Cretaceous tyranosaurid coprolite from Alberta, Canada. Palaios 18: 286 - 294." title="Remarkable preservation of undigested muscle tissue within a Late Cretaceous tyranosaurid coprolite from Alberta, Canada" type="journal article" year="2003">Chin et al. 2003</bibRefCitation>
|
||
), as does toothgrooved bones of prey (
|
||
<bibRefCitation id="EFEDABC53624FF6C8675FB73FA95FB33" author="Erickson, G. M. & Olson, K. H." box="[867,1324,1206,1244]" firstAuthor="Erickson" journalOrPublisher="Journal ofVertebrate Paleontology" pageId="0" pageNumber="166" pagination="175 - 178" part="16" refId="ref9554" refString="Erickson, G. M., and Olson, K. H. 1996. Bite marks attributable to Tyrannosaurus rex: preliminary description and implications. Journal ofVertebrate Paleontology 16: 175 - 178." title="Bite marks attributable to Tyrannosaurus rex: preliminary description and implications" type="journal article" year="1996">Erickson and Olson 1996</bibRefCitation>
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||
) and evidence of a failed attack on a hadrosaur (
|
||
<bibRefCitation id="EFEDABC53624FF6C8628FB35FBE3FAF9" author="Carpenter, K." box="[830,1114,1264,1302]" editor="Perez-Moreno, B. R & Holtz, T. J. & Sanz, J. L. & Moratalla, J." firstAuthor="Carpenter" journalOrPublisher="Gaia: Revista de Geociencias, Museu National de Historia Natural, Lisbon" pageId="0" pageNumber="166" pagination="135 - 144" part="15" refId="ref9069" refString="Carpenter, K. 1998. Evidence of predatory behavior by carnivorous dinosaurs. P. 135 - 144 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="Evidence of predatory behavior by carnivorous dinosaurs" type="book" volumeTitle="Aspects of Theropod Paleobiology" year="1998">Carpenter 1998</bibRefCitation>
|
||
) and ceratopsian (Happ this volume).
|
||
</paragraph>
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||
</subSubSection>
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<subSubSection id="C36685BF3624FF6E870EFAE9F74AFE16" lastPageId="2" lastPageNumber="168" pageId="0" pageNumber="166" type="discussion">
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<paragraph id="8BC3D6343624FF6C870EFAE9F93EF7FB" blockId="0.[450,1819,787,3011]" pageId="0" pageNumber="166">
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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>
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||
, 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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<emphasis id="B9080A263624FF6C836EFAE9FD84FA62" italics="true" pageId="0" pageNumber="166">Tyrannosaurus</emphasis>
|
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</taxonomicName>
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||
a predator,
|
||
<bibRefCitation id="EFEDABC53624FF6C860BFAA2FBA4FA62" author="Lambe, L. M." box="[797,1053,1383,1421]" firstAuthor="Lambe" journalOrPublisher="Geological Survey of Canada Memoir" pageId="0" pageNumber="166" part="100" refId="ref9868" refString="Lambe, L. M. 1917. The Cretaceous Theropodous Dinosaur Gorgosaurus. Geological Survey of Canada Memoir 100." title="The Cretaceous Theropodous Dinosaur Gorgosaurus" type="journal volume" year="1917">Lambe (1917)</bibRefCitation>
|
||
dissented, considering tyrannosaurids to be scavengers instead, primarily because of the apparent absence of tooth wear. Tyrannosaurids as obligatory scavengers never gained popularity until
|
||
<emphasis id="B9080A263624FF6C84EFF9D2FD36F9D2" box="[505,655,1559,1597]" italics="true" pageId="0" pageNumber="166">recency</emphasis>
|
||
, 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>
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||
. 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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<emphasis id="B9080A263624FF6C86C9F997FB55F997" box="[991,1260,1618,1656]" italics="true" pageId="0" pageNumber="166">Tyrannosaurus</emphasis>
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</taxonomicName>
|
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has been considered an active predator, although the mode of attack remains controversial and includes flank bite and run (
|
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<bibRefCitation id="EFEDABC53624FF6C8623F90DFC4CF901" author="Paul, G. S." box="[821,1013,1736,1774]" editor="Currie, P. & Koster, E." firstAuthor="Paul" journalOrPublisher="Occasional Papers of the Tyrrell Museum of Palaeontology" pageId="0" pageNumber="166" pagination="173 - 178" part="3" refId="ref10285" refString="Paul, G. S. 1987. Predation in the meat eating dinosaurs. P. 173 - 178 in Currie, P., and Koster, E. (eds.). Fourth Symposium on Mesozoic Terrestrial Ecosystems, Short Papers. Occasional Papers of the Tyrrell Museum of Palaeontology 3." title="Predation in the meat eating dinosaurs" type="proceedings" volumeTitle="Fourth Symposium on Mesozoic Terrestrial Ecosystems, Short Papers" year="1987">Paul 1987</bibRefCitation>
|
||
), opportunistic (
|
||
<bibRefCitation id="EFEDABC53624FF6C8038F90DF9AEF901" author="Farlow, J. O." box="[1326,1559,1736,1774]" firstAuthor="Farlow" journalOrPublisher="Historical Biology" pageId="0" pageNumber="166" pagination="159 - 165" part="7" refId="ref9650" refString="Farlow, J. O. 1994. Speculations about the carrion-locating ability of tyrannosaurs. Historical Biology 7: 159 - 165." title="Speculations about the carrion-locating ability of tyrannosaurs" type="journal article" year="1994">Farlow 1994</bibRefCitation>
|
||
), and neck or snout crushing (
|
||
<bibRefCitation id="EFEDABC53624FF6C87E0F8C6FC55F8C6" author="Molnar, R. E." box="[758,1004,1795,1833]" editor="Perez-Moreno, B. P. & Holtz, T. J. & Sanz, J. L. & Moratalla, J." firstAuthor="Molnar" journalOrPublisher="Gaia: Revista de Geociencias, Museu Nacional de Historia Natural, Lisbon" pageId="0" pageNumber="166" pagination="193 - 218" part="15" refId="ref10091" refString="Molnar, R. E. 1998. Mechanical factors in the design of the skull of Tyrannosaurus rex (Osborn, 1905). P. 193 - 218 in Perez-Moreno, B. P., Holtz, T. J., Sanz, J. L., and Moratalla, J. (eds.). Aspects ofTheropod Paleobiology. Gaia: Revista de Geociencias, Museu Nacional de Historia Natural, Lisbon, 15." title="Mechanical factors in the design of the skull of Tyrannosaurus rex (Osborn, 1905)" type="book" volumeTitle="Aspects ofTheropod Paleobiology" year="1998">Molnar 1998</bibRefCitation>
|
||
). The method of killing is assumed to have been the jaws, and the bite force has been variously calculated or estimated to have been 13,400 N (
|
||
<bibRefCitation id="EFEDABC53624FF6C8694F8BDFB52F871" author="Rayfield, E. J. & Norman, D. B. & Horner, C. C. & Horner, J. R. & Smith, P. M. & Thomason, J. J. & Upchurch, P." box="[898,1259,1912,1950]" etAl="et al." firstAuthor="Rayfield" journalOrPublisher="Nature" pageId="0" pageNumber="166" pagination="1033 - 1037" part="409" refId="ref10358" refString="Rayfield, E. J., Norman, D. B., Horner, C. C., Horner, J. R., Smith, P. M., Thomason, J. J., and Upchurch, P. 2001. Cranial design and function in a large theropod dinosaur. Nature 409: 1033 - 1037." title="Cranial design and function in a large theropod dinosaur" type="journal article" year="2001">Rayfield et al. 2001</bibRefCitation>
|
||
), 6400—13,400 N (
|
||
<bibRefCitation id="EFEDABC53624FF6C8352F8BDFDE4F836" author="Erickson, G. M. & Olson, K. H." etAl="et al." firstAuthor="Erickson" journalOrPublisher="Journal ofVertebrate Paleontology" pageId="0" pageNumber="166" pagination="175 - 178" part="16" refId="ref9554" refString="Erickson, G. M., and Olson, K. H. 1996. Bite marks attributable to Tyrannosaurus rex: preliminary description and implications. Journal ofVertebrate Paleontology 16: 175 - 178." title="Bite marks attributable to Tyrannosaurus rex: preliminary description and implications" type="journal article" year="1996">Erickson et al. 1996</bibRefCitation>
|
||
), or even 183,000-235,000 N (
|
||
<bibRefCitation id="EFEDABC53624FF6C818BF876FAC5F836" author="Meers, M. B." box="[1181,1404,1971,2009]" firstAuthor="Meers" journalOrPublisher="Historical Biology" pageId="0" pageNumber="166" pagination="1 - 12" part="16" refId="ref10034" refString="Meers, M. B. 2002. Maximum bite force and prey size of Tyrannosaurus rex and their relationships to the inference of feeding behavior. Historical Biology 16: 1 - 12." title="Maximum bite force and prey size of Tyrannosaurus rex and their relationships to the inference of feeding behavior" type="journal article" year="2002">Meers 2002</bibRefCitation>
|
||
). As Meers has noted, the high forces are consistent with either scavenging or predation.
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343624FF6C870EF7ECF9D5F42F" blockId="0.[450,1819,787,3011]" pageId="0" pageNumber="166">
|
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In arguing against predation,
|
||
<bibRefCitation id="EFEDABC53624FF6C8125F7ECF9ABF7A0" author="Horner, J. R. & Lessem, D." box="[1075,1554,2089,2127]" 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>
|
||
cite the shortness of the forelimbs as being useless for holding prey. In support of this position,
|
||
<bibRefCitation id="EFEDABC53624FF6C8734F75BFC07F72B" author="Lingham-Soliar, T." box="[546,958,2206,2244]" firstAuthor="Lingham-Soliar" journalOrPublisher="Geology Today" pageId="0" pageNumber="166" pagination="16 - 20" part="14" refId="ref9932" refString="Lingham-Soliar, T. 1998. Guess who's coming to dinner: a portrait of Tyrannosaurus as a predator. Geology Today 14: 16 - 20." title="Guess who's coming to dinner: a portrait of Tyrannosaurus as a predator" type="journal article" year="1998">Lingham-Soliar (1998)</bibRefCitation>
|
||
considered head shaking as a means of flesh removal from small prey and direct ripping of flesh from large prey. Carpenter (2002) has shown that without exception, no theropod could extend its forelimb beyond the snout, thus limiting its usefulness as a
|
||
<emphasis id="B9080A263624FF6C80A6F68AF91DF69A" box="[1456,1700,2383,2421]" italics="true" pageId="0" pageNumber="166">prey-grasping</emphasis>
|
||
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">
|
||
<emphasis id="B9080A263624FF6C87B3F603FC15F606" box="[677,940,2502,2537]" italics="true" pageId="0" pageNumber="166">Tyrannosaurus</emphasis>
|
||
</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,
|
||
<bibRefCitation id="EFEDABC53624FF6C86B6F4E1FA1EF4A5" author="Carpenter, K. & Smith, M." box="[928,1447,2852,2890]" editor="Tanke, D. & Carpenter, K." firstAuthor="Carpenter" journalOrPublisher="Indiana University Press, Bloomington" pageId="0" pageNumber="166" 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>
|
||
concluded that the forelimb was powerful, an interpretation previously made by
|
||
<bibRefCitation id="EFEDABC53624FF6C8314F49AF958F46A" author="Brown, B." box="[1538,1761,2911,2949]" firstAuthor="Brown" journalOrPublisher="Museum Journal" pageId="0" pageNumber="166" pagination="271 - 279" part="15" refId="ref8906" refString="Brown, B. 1915. Tyrannosaurus, the largest flesh-eating animal that ever lived. Museum Journal 15: 271 - 279." title="Tyrannosaurus, the largest flesh-eating animal that ever lived" type="journal article" year="1915">Brown (1915</bibRefCitation>
|
||
, p. 271): “front limbs exceedingly small but set for a powerful clutch.”
|
||
</paragraph>
|
||
<caption id="DF0386BC3624FF6C827FF691F72CF454" ID-DOI="http://doi.org/10.5281/zenodo.3942815" ID-Zenodo-Dep="3942815" httpUri="https://zenodo.org/record/3942815/files/figure.png" pageId="0" pageNumber="166" targetBox="[465,2349,252,2501]" targetPageId="1">
|
||
<paragraph id="8BC3D6343624FF6C827FF691F72CF454" blockId="0.[1892,2346,2383,3003]" pageId="0" pageNumber="166">
|
||
Figure 10.1.
|
||
<emphasis id="B9080A263624FF6C8D28F691F6B7F698" box="[2110,2318,2388,2423]" italics="true" pageId="0" pageNumber="166">Comparison</emphasis>
|
||
of
|
||
<emphasis id="B9080A263624FF6C8280F64CF7FBF643" box="[1942,2114,2441,2476]" italics="true" pageId="0" pageNumber="166">maximum</emphasis>
|
||
forelimb motion
|
||
<emphasis id="B9080A263624FF6C82E6F678F729F5FA" italics="true" pageId="0" pageNumber="166">in 3 well-known theropods. None</emphasis>
|
||
of the dinosaurs can reach its manus to its mouth as a result of constraints in
|
||
<emphasis id="B9080A263624FF6C827EF500F826F507" box="[1896,1951,2757,2792]" italics="true" pageId="0" pageNumber="166">the</emphasis>
|
||
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>
|
||
has the greatest range or retraction.
|
||
<emphasis id="B9080A263624FF6C82D7F45DF7BBF454" box="[1985,2050,2968,3003]" italics="true" pageId="0" pageNumber="166">Not</emphasis>
|
||
to scale.
|
||
</paragraph>
|
||
</caption>
|
||
<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>
|
||
,
|
||
</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>
|
||
must have had a rough, active life.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C36685BF3626FF6E812FFDCBF7B9FCF0" pageId="2" pageNumber="168" type="materials_examined">
|
||
<paragraph id="8BC3D6343626FF6E812FFDCBF7B9FCF0" blockId="2.[997,2358,226,799]" pageId="2" pageNumber="168">
|
||
The materials used in this study are as follows: scapula of
|
||
<materialsCitation id="3B14DC693626FF6E8D6AFDCBF697FDDB" ID-GBIF-Occurrence="2813095319" box="[2172,2350,526,564]" collectionCode="BHI" pageId="2" pageNumber="168" specimenCode="BHI 3033">BHI 3033</materialsCitation>
|
||
,
|
||
<materialsCitation id="3B14DC693626FF6E86FFFD8FFB58FD9F" ID-GBIF-Occurrence="2813095314" box="[1001,1249,586,624]" collectionCode="DMNH" pageId="2" pageNumber="168" specimenCode="DMNH 2827">DMNH 2827</materialsCitation>
|
||
, and
|
||
<materialsCitation id="3B14DC693626FF6E8050FD8FFA46FD9F" ID-GBIF-Occurrence="2813095346" box="[1350,1535,586,624]" collectionCode="MOR" pageId="2" pageNumber="168" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
; furcula of
|
||
<materialsCitation id="3B14DC693626FF6E83CEFD8FF7BEFD9F" ID-GBIF-Occurrence="2813095331" box="[1752,2055,586,624]" collectionCode="FMNH" pageId="2" pageNumber="168" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
,
|
||
<materialsCitation id="3B14DC693626FF6E8D0BFD8FF765FD9F" ID-GBIF-Occurrence="2813095313" box="[2077,2268,586,624]" collectionCode="MOR" pageId="2" pageNumber="168" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
, and
|
||
<materialsCitation id="3B14DC693626FF6E86F3FD41FAB4FD45" ID-GBIF-Occurrence="2813095347" box="[997,1293,644,682]" collectionCode="TCM" pageId="2" pageNumber="168" specimenCode="TCM 2001.90.1">TCM 2001.90.1</materialsCitation>
|
||
; humerus of
|
||
<materialsCitation id="3B14DC693626FF6E831EFD41F906FD45" ID-GBIF-Occurrence="2813095332" box="[1544,1727,644,682]" collectionCode="BHI" pageId="2" pageNumber="168" specimenCode="BHI 6230">BHI 6230</materialsCitation>
|
||
,
|
||
<materialsCitation id="3B14DC693626FF6E83CEFD41F7B1FD45" ID-GBIF-Occurrence="2813095322" box="[1752,2056,644,682]" collectionCode="FMNH" pageId="2" pageNumber="168" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
, and
|
||
<materialsCitation id="3B14DC693626FF6E8D65FD41F694FD45" ID-GBIF-Occurrence="2813095342" box="[2163,2349,644,682]" collectionCode="MOR" pageId="2" pageNumber="168" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
; ulna and radius of
|
||
<materialsCitation id="3B14DC693626FF6E802DFD7AF9DFFD0A" ID-GBIF-Occurrence="2813095341" box="[1339,1638,703,741]" collectionCode="FMNH" pageId="2" pageNumber="168" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
,
|
||
<materialsCitation id="3B14DC693626FF6E836CFD7AF88BFD0A" ID-GBIF-Occurrence="2813095338" box="[1658,1842,703,741]" collectionCode="MOR" pageId="2" pageNumber="168" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
, and
|
||
<materialsCitation id="3B14DC693626FF6E8280FD7AF7EDFD0A" ID-GBIF-Occurrence="2813095311" box="[1942,2132,703,741]" collectionCode="MOR" pageId="2" pageNumber="168" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
; and manus of
|
||
<materialsCitation id="3B14DC693626FF6E8100FD3CFAF9FCF0" ID-GBIF-Occurrence="2813095312" box="[1046,1344,761,799]" collectionCode="FMNH" pageId="2" pageNumber="168" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
,
|
||
<materialsCitation id="3B14DC693626FF6E8042FD3CF9B5FCF0" ID-GBIF-Occurrence="2813095324" box="[1364,1548,761,799]" collectionCode="MOR" pageId="2" pageNumber="168" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
,
|
||
<materialsCitation id="3B14DC693626FF6E8336FD3CF959FCF0" ID-GBIF-Occurrence="2813095343" box="[1568,1760,761,799]" collectionCode="MOR" pageId="2" pageNumber="168" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
, and
|
||
<materialsCitation id="3B14DC693626FF6E8250FD3CF842FCF0" ID-GBIF-Occurrence="2813095344" box="[1862,2043,761,799]" collectionCode="BHI" pageId="2" pageNumber="168" specimenCode="BHI 6230">BHI 6230</materialsCitation>
|
||
.
|
||
</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="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">
|
||
<emphasis id="B9080A263626FF6E8366FC74F8C4FC38" box="[1648,1917,945,983]" italics="true" pageId="2" pageNumber="168">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
have been described by
|
||
<bibRefCitation id="EFEDABC53626FF6E86F3FC29FA58FBFD" author="Carpenter, K. & Smith, M." box="[997,1505,1004,1042]" editor="Tanke, D. & Carpenter, K." firstAuthor="Carpenter" journalOrPublisher="Indiana University Press, Bloomington" pageId="2" pageNumber="168" 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 by
|
||
<bibRefCitation id="EFEDABC53626FF6E8366FC29F8C2FBFD" author="Brochu, C. A." box="[1648,1915,1004,1042]" 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>
|
||
. Since then, the furcula has been described (
|
||
<bibRefCitation id="EFEDABC53626FF6E807DFBE2F896FBA2" author="Larson, P & Rigby, J. K." box="[1387,1839,1063,1101]" 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>
|
||
), the third metacarpal and semilunate carpal have been found (described below), and new information is available for the scapula and coracoid. To date, no ossified sternal plates are known, but these are predicted to resemble those of
|
||
<taxonomicName id="4C7CADB73626FF6E8D5DFB12F688FB12" box="[2123,2353,1239,1277]" class="Reptilia" family="Tyrannosauridae" genus="Gorgosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="168" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9080A263626FF6E8D5DFB12F688FB12" box="[2123,2353,1239,1277]" italics="true" pageId="2" pageNumber="168">Gorgosaurus</emphasis>
|
||
</taxonomicName>
|
||
as described by
|
||
<bibRefCitation id="EFEDABC53626FF6E8015FAD7F9BAFAD7" author="Lambe, L. M." box="[1283,1539,1298,1336]" firstAuthor="Lambe" journalOrPublisher="Geological Survey of Canada Memoir" pageId="2" pageNumber="168" part="100" refId="ref9868" refString="Lambe, L. M. 1917. The Cretaceous Theropodous Dinosaur Gorgosaurus. Geological Survey of Canada Memoir 100." title="The Cretaceous Theropodous Dinosaur Gorgosaurus" type="journal volume" year="1917">Lambe (1917)</bibRefCitation>
|
||
.
|
||
<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">
|
||
<emphasis id="B9080A263626FF6E80C7FA88F967FA9C" box="[1489,1758,1357,1395]" italics="true" pageId="2" pageNumber="168">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
but came to no conclusion.
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343626FF6E86F7FA19FBCEF9EE" blockId="2.[993,1143,1491,1537]" box="[993,1143,1500,1537]" pageId="2" pageNumber="168">
|
||
<heading id="D08B61583626FF6E86F7FA19FBCEF9EE" bold="true" box="[993,1143,1500,1537]" fontSize="12" level="9" pageId="2" pageNumber="168" reason="2">
|
||
<emphasis id="B9080A263626FF6E86F7FA19FBCEF9EE" bold="true" box="[993,1143,1500,1537]" pageId="2" pageNumber="168">Furcula</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<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>
|
||
,
|
||
</emphasis>
|
||
the presence of which was predicted by
|
||
<bibRefCitation id="EFEDABC53626FF6E86F7F9ACFA52F960" author="Carpenter, K. & Smith, M." box="[993,1515,1641,1679]" editor="Tanke, D. & Carpenter, K." firstAuthor="Carpenter" journalOrPublisher="Indiana University Press, Bloomington" pageId="2" pageNumber="168" 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>
|
||
, is now known for several specimens (
|
||
<bibRefCitation id="EFEDABC53626FF6E8DB8F9ACFABEF925" 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>
|
||
). It is broadly U or boomerang shaped (
|
||
<figureCitation id="1347CAB13626FF6E82E3F961F72EF925" box="[2037,2199,1700,1738]" captionStart="Fig. 10.2" captionText="Figure 10.2. Furcula of Tyrannosaurus rex include several with pathologies, including fractures (black arrows) and localized exostosis of stress fractures (white arrows). FMNH PP2081 in posterior (A) and anterior (B) views; MOR 980 in posterior (C) and anterior (D) views; TCM 2001.90.1 in posterior (E), anterior (F), and lateral (G) views. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942817" httpUri="https://zenodo.org/record/3942817/files/figure.png" pageId="2" pageNumber="168" targetBox="[428,2309,188,1283]" targetPageId="3">Fig. 10.2</figureCitation>
|
||
), with a roughened sutural scar (acrominal facet) on the epicleidium for a ligamentous attachment to the acromion of the scapula (
|
||
<figureCitation id="1347CAB13626FF6E8248F8DFF7B9F8AF" box="[1886,2048,1818,1856]" captionStart="Fig. 10.3" captionText="Figure 10.3. Close-up of the epicleidial facet (arrow) on the dorsal edge of the scapula DMNH 2827 (A) and lateral view with the furcula articulated (B)." figureDoi="http://doi.org/10.5281/zenodo.3942819" httpUri="https://zenodo.org/record/3942819/files/figure.png" pageId="2" pageNumber="168" targetBox="[437,1808,1361,2928]" targetPageId="3">Fig. 10.3</figureCitation>
|
||
; see also below). In birds, the rami of the furcula have a nearly circular or laterally compressed cross-sectional geometry that allows them to act as a spring, with laterally directed tension added on the downstroke and medial directed recoil on the upstroke (
|
||
<bibRefCitation id="EFEDABC53626FF6E8090F7C0F962F7C4" author="Jenkins, F. A. & Dial, K. P. & Goslow, G. E." box="[1414,1755,2053,2091]" etAl="et al." firstAuthor="Jenkins" journalOrPublisher="Science" pageId="2" pageNumber="168" pagination="1495 - 1498" part="241" refId="ref9761" refString="Jenkins, F. A., Dial, K. P., and Goslow, G. E. 1988. A cineradiographic analysis of bird flight: the wishbone in starlings is a spring. Science 241: 1495 - 1498." title="A cineradiographic analysis of bird flight: the wishbone in starlings is a spring" type="journal article" year="1988">Jenkins et al. 1988</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFEDABC53626FF6E83F8F7C0F79FF7C4" author="Boggs, D. F. & Jenkins, F. A. & Dial, K. P." box="[1774,2086,2053,2091]" etAl="et al." firstAuthor="Boggs" journalOrPublisher="Journal of Experimental Biology" pageId="2" pageNumber="168" pagination="1403 - 1412" part="200" refId="ref8799" refString="Boggs, D. F., Jenkins, F. A., and Dial, K. P. 1997. The effects of the wingbeat cycle on respiration in black-billed magpies (Pica pica). Journal of Experimental Biology 200: 1403 - 1412." title="The effects of the wingbeat cycle on respiration in black-billed magpies (Pica pica)" type="journal article" year="1997">Boggs et al. 1997</bibRefCitation>
|
||
;
|
||
<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>
|
||
,
|
||
</emphasis>
|
||
however, the furcula is clearly designed to resist extreme lateral forces: (1) the ramfare anteroposteriorly flattened (cf.
|
||
<figureCitation id="1347CAB13626FF6E8D3FF7BFF773F74F" box="[2089,2250,2170,2208]" captionStart="Fig. 10.2" captionText="Figure 10.2. Furcula of Tyrannosaurus rex include several with pathologies, including fractures (black arrows) and localized exostosis of stress fractures (white arrows). FMNH PP2081 in posterior (A) and anterior (B) views; MOR 980 in posterior (C) and anterior (D) views; TCM 2001.90.1 in posterior (E), anterior (F), and lateral (G) views. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942817" httpUri="https://zenodo.org/record/3942817/files/figure.png" pageId="2" pageNumber="168" targetBox="[428,2309,188,1283]" targetPageId="3">Fig. 10.2</figureCitation>
|
||
F, G), thus prohibiting lateromedial springlike action; (2) the rami diverge, thus directing lateral stresses down the shaft (
|
||
<figureCitation id="1347CAB13626FF6E8215F735F814F6F9" box="[1795,1965,2288,2326]" captionStart="Fig. 10.4" captionText="Figure 10.4. Reconstruction of the pectoral girdle and forelimb of Tyrannosaurus showing (A) the distribution of force from the scapula (a arrows), through the furcula (b arrows), which results in cumulative force (c arrow) at the middle of the furcula. Resisting force c explains why the furcula is deepest at the midline, which is unlike any other theropod furcula. The range of motion for the forelimb segments (B), angle represented by arc a = 40°, arc b = 60°, arc c = 67°. " figureDoi="http://doi.org/10.5281/zenodo.3942821" httpUri="https://zenodo.org/record/3942821/files/figure.png" pageId="2" pageNumber="168" targetBox="[437,1808,1361,2928]" targetPageId="5">Fig. 10.4</figureCitation>
|
||
A; and (3) the rami deepen distally from the epicleidium and the furcula is deepest (thickest vertically in the anatomical position) near the midline to counter the stresses directed down the rami. In most theropods, the furcula
|
||
<emphasis id="B9080A263626FF6E8D84F665F715F629" box="[2194,2220,2464,2502]" italics="true" pageId="2" pageNumber="168">is</emphasis>
|
||
nearly uniform throughout its length (see
|
||
<bibRefCitation id="EFEDABC53626FF6E8377F61EF788F5EE" author="Chure, D. J. & J. H. Madsen, Jr." box="[1633,2097,2523,2561]" firstAuthor="Chure" journalOrPublisher="Journal ofVertebrate Paleontology" pageId="2" pageNumber="168" pagination="573 - 577" part="16" refId="ref9333" refString="Chure, D. J., and J. H. Madsen, Jr. 1996. On the presence of furculae in some nonmaniraptoran theropods. Journal ofVertebrate Paleontology 16: 573 - 577." title="On the presence of furculae in some nonmaniraptoran theropods" type="journal article" year="1996">Chure and Madsen 1996</bibRefCitation>
|
||
, fig. 2, 3;
|
||
<bibRefCitation id="EFEDABC53626FF6E8DF4F61EFA14F5D3" author="Makovicky, P & Currie, P. J." firstAuthor="Makovicky" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="2" pageNumber="168" pagination="143 - 149" part="18" refId="ref9995" refString="Makovicky, P, and Currie, P. J. 1998. The presence of a furcula in tyrannosaurid theropods, and its phylogenetic and functional implications. Journal of Vertebrate Paleontology 18: 143 - 149." title="The presence of a furcula in tyrannosaurid theropods, and its phylogenetic and functional implications" type="journal article" year="1998">Makovicky and Currie 1998</bibRefCitation>
|
||
, fig. 1;
|
||
<bibRefCitation id="EFEDABC53626FF6E832BF5D3F829F5D3" author="Currie, P. J. & Trexler, D. & Koppelhus, E. B. & Wicks, K. & Murphy, N." box="[1597,1936,2582,2620]" editor="Carpenter, K." etAl="et al." firstAuthor="Currie" journalOrPublisher="Indiana University Press, Bloomington" pageId="2" pageNumber="168" pagination="313 - 324" refId="ref9392" refString="Currie, P. J., Trexler, D., Koppelhus, E. B., Wicks, K., and Murphy, N. 2005. An unusual multi-individual bonebed in the Two Medicine Formation (Late Cretaceous, Campanian) of Montana (USA). P. 313 - 324, in Carpenter, K. (ed.). The Carnivorous Dinosaurs. Indiana University Press, Bloomington." title="An unusual multi-individual bonebed in the Two Medicine Formation (Late Cretaceous, Campanian) of Montana (USA)" type="book" volumeTitle="The Carnivorous Dinosaurs" year="2005">Currie et al. 2005</bibRefCitation>
|
||
, 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">
|
||
<emphasis id="B9080A263626FF6E82C2F594F758F598" box="[2004,2273,2641,2679]" italics="true" pageId="2" pageNumber="168">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
furcula
|
||
<emphasis id="B9080A263626FF6E8121F549FBE9F55D" box="[1079,1104,2700,2738]" italics="true" pageId="2" pageNumber="168">is</emphasis>
|
||
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">
|
||
<emphasis id="B9080A263626FF6E80FEF502F94CF502" box="[1512,1781,2759,2797]" italics="true" pageId="2" pageNumber="168">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
furculae are pathologic (
|
||
<figureCitation id="1347CAB13626FF6E8DF2F502FB9EF4C8" captionStart="Fig. 10.2" captionText="Figure 10.2. Furcula of Tyrannosaurus rex include several with pathologies, including fractures (black arrows) and localized exostosis of stress fractures (white arrows). FMNH PP2081 in posterior (A) and anterior (B) views; MOR 980 in posterior (C) and anterior (D) views; TCM 2001.90.1 in posterior (E), anterior (F), and lateral (G) views. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942817" httpUri="https://zenodo.org/record/3942817/files/figure.png" pageId="2" pageNumber="168" targetBox="[428,2309,188,1283]" targetPageId="3">Fig. 10.2</figureCitation>
|
||
) and were examined by computed tomographic scanning and X-ray. Two of them are missing portions of a ramus (
|
||
<materialsCitation id="3B14DC693626FF6E820BF4F9F7FDF48D" ID-GBIF-Occurrence="2813095315" box="[1821,2116,2876,2914]" collectionCode="FMNH" pageId="2" pageNumber="168" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
[
|
||
<figureCitation id="1347CAB13626FF6E8D77F4F9F6BDF48D" box="[2145,2308,2876,2914]" captionStart="Fig. 10.2" captionText="Figure 10.2. Furcula of Tyrannosaurus rex include several with pathologies, including fractures (black arrows) and localized exostosis of stress fractures (white arrows). FMNH PP2081 in posterior (A) and anterior (B) views; MOR 980 in posterior (C) and anterior (D) views; TCM 2001.90.1 in posterior (E), anterior (F), and lateral (G) views. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942817" httpUri="https://zenodo.org/record/3942817/files/figure.png" pageId="2" pageNumber="168" targetBox="[428,2309,188,1283]" targetPageId="3">Fig. 10.2</figureCitation>
|
||
A, B] and
|
||
<materialsCitation id="3B14DC693626FF6E8177F4B3FA3BF473" ID-GBIF-Occurrence="2813095328" box="[1121,1410,2934,2972]" collectionCode="TCM" pageId="2" pageNumber="168" specimenCode="TCM 2001.90.1">TCM 2001.90.1</materialsCitation>
|
||
[
|
||
<figureCitation id="1347CAB13626FF6E80B4F4B3F9F3F473" box="[1442,1610,2934,2972]" captionStart="Fig. 10.2" captionText="Figure 10.2. Furcula of Tyrannosaurus rex include several with pathologies, including fractures (black arrows) and localized exostosis of stress fractures (white arrows). FMNH PP2081 in posterior (A) and anterior (B) views; MOR 980 in posterior (C) and anterior (D) views; TCM 2001.90.1 in posterior (E), anterior (F), and lateral (G) views. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942817" httpUri="https://zenodo.org/record/3942817/files/figure.png" pageId="2" pageNumber="168" targetBox="[428,2309,188,1283]" targetPageId="3">Fig. 10.2</figureCitation>
|
||
E, F]), and 2 show localized swelling of the cortical bone (
|
||
<materialsCitation id="3B14DC693626FF6E8037F474FA6EF438" ID-GBIF-Occurrence="2813095303" box="[1313,1495,2993,3031]" collectionCode="MOR" pageId="2" pageNumber="168" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
[
|
||
<figureCitation id="1347CAB13626FF6E80E6F474F92BF438" box="[1520,1682,2993,3031]" captionStart="Fig. 10.2" captionText="Figure 10.2. Furcula of Tyrannosaurus rex include several with pathologies, including fractures (black arrows) and localized exostosis of stress fractures (white arrows). FMNH PP2081 in posterior (A) and anterior (B) views; MOR 980 in posterior (C) and anterior (D) views; TCM 2001.90.1 in posterior (E), anterior (F), and lateral (G) views. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942817" httpUri="https://zenodo.org/record/3942817/files/figure.png" pageId="2" pageNumber="168" targetBox="[428,2309,188,1283]" targetPageId="3">Fig. 10.2</figureCitation>
|
||
D] and
|
||
<materialsCitation id="3B14DC693626FF6E8202F474F788F438" ID-GBIF-Occurrence="2813095302" box="[1812,2097,2993,3031]" collectionCode="TCM" pageId="2" pageNumber="168" specimenCode="TCM 2001.90.1">TCM 2001.90.1</materialsCitation>
|
||
[
|
||
<figureCitation id="1347CAB13626FF6E8D5DF474F748F438" box="[2123,2289,2993,3031]" captionStart="Fig. 10.2" captionText="Figure 10.2. Furcula of Tyrannosaurus rex include several with pathologies, including fractures (black arrows) and localized exostosis of stress fractures (white arrows). FMNH PP2081 in posterior (A) and anterior (B) views; MOR 980 in posterior (C) and anterior (D) views; TCM 2001.90.1 in posterior (E), anterior (F), and lateral (G) views. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942817" httpUri="https://zenodo.org/record/3942817/files/figure.png" pageId="2" pageNumber="168" targetBox="[428,2309,188,1283]" targetPageId="3">Fig. 10.2</figureCitation>
|
||
F]).
|
||
</paragraph>
|
||
<caption id="DF0386BC3627FF6F825DFA91F7A1F77C" ID-DOI="http://doi.org/10.5281/zenodo.3942817" ID-Zenodo-Dep="3942817" httpUri="https://zenodo.org/record/3942817/files/figure.png" pageId="3" pageNumber="169" startId="3.[1867,1971,1364,1399]" targetBox="[428,2309,188,1283]" targetPageId="3">
|
||
<paragraph id="8BC3D6343627FF6F825DFA91F7A1F77C" blockId="3.[1862,2308,1358,2195]" pageId="3" pageNumber="169">
|
||
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>
|
||
<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
|
||
<emphasis id="B9080A263627FF6F8D9DFA7BF718F9F9" italics="true" pageId="3" pageNumber="169">pathologies, including</emphasis>
|
||
<emphasis id="B9080A263627FF6F8DBBFA36F819F9A4" bold="true" pageId="3" pageNumber="169">fractures</emphasis>
|
||
<emphasis id="B9080A263627FF6F82BAF9EDF801F9A4" box="[1964,1976,1576,1611]" italics="true" pageId="3" pageNumber="169">(</emphasis>
|
||
black arrows
|
||
<emphasis id="B9080A263627FF6F8D82F9EDF719F9A4" box="[2196,2208,1576,1611]" italics="true" pageId="3" pageNumber="169">)</emphasis>
|
||
and localized exostosis of stress fractures
|
||
<emphasis id="B9080A263627FF6F8D4DF957F7D1F95A" box="[2139,2152,1682,1717]" italics="true" pageId="3" pageNumber="169">(</emphasis>
|
||
white arrows
|
||
<emphasis id="B9080A263627FF6F82A8F902F876F905" box="[1982,1999,1735,1770]" italics="true" pageId="3" pageNumber="169">).</emphasis>
|
||
FMNH
|
||
<emphasis id="B9080A263627FF6F8D4FF902F8DFF8F0" italics="true" pageId="3" pageNumber="169">PP2081 in</emphasis>
|
||
posterior
|
||
<emphasis id="B9080A263627FF6F8D0AF939F791F8F0" box="[2076,2088,1788,1823]" italics="true" pageId="3" pageNumber="169">(</emphasis>
|
||
A
|
||
<emphasis id="B9080A263627FF6F8D54F939F7F7F8F0" box="[2114,2126,1788,1823]" italics="true" pageId="3" pageNumber="169">)</emphasis>
|
||
and
|
||
<emphasis id="B9080A263627FF6F8DB0F939F805F8BB" italics="true" pageId="3" pageNumber="169">anterior (B)</emphasis>
|
||
views;
|
||
<materialsCitation id="3B14DC693627FF6F8D28F8F4F753F8BB" ID-GBIF-Occurrence="2813095337" box="[2110,2282,1841,1876]" collectionCode="MOR" pageId="3" pageNumber="169" specimenCode="MOR 980">
|
||
MOR
|
||
<emphasis id="B9080A263627FF6F8DB2F8F4F753F8BB" box="[2212,2282,1841,1876]" italics="true" pageId="3" pageNumber="169">980</emphasis>
|
||
</materialsCitation>
|
||
<emphasis id="B9080A263627FF6F8251F8A3F79EF866" box="[1863,2087,1894,1929]" italics="true" pageId="3" pageNumber="169">in posterior (</emphasis>
|
||
C
|
||
<emphasis id="B9080A263627FF6F8D29F8A3F7F3F866" box="[2111,2122,1894,1929]" italics="true" pageId="3" pageNumber="169">)</emphasis>
|
||
and
|
||
<emphasis id="B9080A263627FF6F8DB5F8A3F879F851" italics="true" pageId="3" pageNumber="169">anterior (D)</emphasis>
|
||
views;
|
||
<materialsCitation id="3B14DC693627FF6F8D52F85EF84BF81C" ID-GBIF-Occurrence="2813095307" collectionCode="TCM" pageId="3" pageNumber="169" specimenCode="TCM 2001.90.1">
|
||
TCM
|
||
<emphasis id="B9080A263627FF6F8250F815F84BF81C" box="[1862,2034,2000,2035]" italics="true" pageId="3" pageNumber="169">2001.90.1</emphasis>
|
||
</materialsCitation>
|
||
in posterior
|
||
<emphasis id="B9080A263627FF6F825EF7C0F8CEF7C7" box="[1864,1911,2053,2088]" italics="true" pageId="3" pageNumber="169">(E),</emphasis>
|
||
anterior
|
||
<emphasis id="B9080A263627FF6F8D08F7C0F7F2F7C7" box="[2078,2123,2053,2088]" italics="true" pageId="3" pageNumber="169">(F),</emphasis>
|
||
and lateral
|
||
<emphasis id="B9080A263627FF6F8285F7FEF827F7B1" box="[1939,1950,2107,2142]" italics="true" pageId="3" pageNumber="169">(</emphasis>
|
||
G
|
||
<emphasis id="B9080A263627FF6F82AFF7FEF781F7B1" box="[1977,2104,2107,2142]" italics="true" pageId="3" pageNumber="169">) views.</emphasis>
|
||
Scale
|
||
<emphasis id="B9080A263627FF6F8DBBF7FEF772F7B1" box="[2221,2251,2107,2142]" italics="true" pageId="3" pageNumber="169">in</emphasis>
|
||
centimeters.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF0386BC3627FF6F8250F5D1F86FF49A" ID-DOI="http://doi.org/10.5281/zenodo.3942819" ID-Zenodo-Dep="3942819" httpUri="https://zenodo.org/record/3942819/files/figure.png" pageId="3" pageNumber="169" startId="3.[1862,1966,2580,2615]" targetBox="[437,1808,1361,2928]" targetPageId="3">
|
||
<paragraph id="8BC3D6343627FF6F8250F5D1F86FF49A" blockId="3.[1859,2300,2574,2933]" pageId="3" pageNumber="169">
|
||
Figure 10.3.
|
||
<emphasis id="B9080A263627FF6F8D34F5D1F702F5D8" box="[2082,2235,2580,2615]" italics="true" pageId="3" pageNumber="169">Close-up</emphasis>
|
||
of
|
||
<emphasis id="B9080A263627FF6F8250F58CF78FF583" box="[1862,2102,2633,2668]" italics="true" pageId="3" pageNumber="169">the epicleidial</emphasis>
|
||
facet
|
||
<emphasis id="B9080A263627FF6F8DB2F58CF82BF54F" italics="true" pageId="3" pageNumber="169">(arrow)</emphasis>
|
||
on
|
||
<emphasis id="B9080A263627FF6F82CCF5B8F7ABF54F" box="[2010,2066,2685,2720]" italics="true" pageId="3" pageNumber="169">the</emphasis>
|
||
dorsal edge of
|
||
<emphasis id="B9080A263627FF6F8263F576F812F539" box="[1909,1963,2739,2774]" italics="true" pageId="3" pageNumber="169">the</emphasis>
|
||
scapula
|
||
<materialsCitation id="3B14DC693627FF6F8D53F576F824F4E4" ID-GBIF-Occurrence="2813095326" collectionCode="DMNH" pageId="3" pageNumber="169" specimenCode="DMNH 2827">
|
||
DMNH
|
||
<emphasis id="B9080A263627FF6F8255F52DF824F4E4" box="[1859,1949,2792,2827]" italics="true" pageId="3" pageNumber="169">2827</emphasis>
|
||
</materialsCitation>
|
||
<emphasis id="B9080A263627FF6F82BCF52DF863F4E4" box="[1962,2010,2792,2827]" italics="true" pageId="3" pageNumber="169">(A)</emphasis>
|
||
and lateral
|
||
<emphasis id="B9080A263627FF6F8DBAF52DF745F4E4" box="[2220,2300,2792,2827]" italics="true" pageId="3" pageNumber="169">view</emphasis>
|
||
with
|
||
<emphasis id="B9080A263627FF6F8288F4D9F868F4D0" box="[1950,2001,2844,2879]" italics="true" pageId="3" pageNumber="169">the</emphasis>
|
||
furcula
|
||
<emphasis id="B9080A263627FF6F8D73F4D9F821F49A" italics="true" pageId="3" pageNumber="169">articulated</emphasis>
|
||
(B).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BC3D6343620FF6886F5FF36F804FDEC" blockId="4.[991,2355,236,930]" pageId="4" pageNumber="170">The missing sections of the rami were broken in life, with subsequent remodeling of the fracture surface. Surprisingly, none show a pseudoarthrosis joint at the site of the break, thus indicating significant displacement of the broken portion, possibly due to the contraction of the M. supracoracoideus brevis, which probably inserted along the ramus.</paragraph>
|
||
<paragraph id="8BC3D6343620FF688125FDDDF711FC71" blockId="4.[991,2355,236,930]" pageId="4" pageNumber="170">
|
||
The amount of force needed to break a ramus was calculated from the largest furcula,
|
||
<materialsCitation id="3B14DC693620FF6881EDFD96F999FD96" ID-GBIF-Occurrence="2813095349" box="[1275,1568,595,633]" collectionCode="TCM" pageId="4" pageNumber="170" specimenCode="TCM 2001.90.1">TCM 2001.90.1</materialsCitation>
|
||
. The fracture is 2.5 cm wide; however, the bone is remodeled on the anterior and posterior sides so as to exaggerate the original thickness of the bone. Therefore, the anterior-posterior thickness is approximated from the undamaged ramus, where it is has the same width, which gives a thickness of 1.35 cm. The ramus can be modeled as an ellipse; therefore the area of the break is given by the following:
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343620FF688359FC28F693FBFC" blockId="4.[1615,1699,1001,1050]" box="[1615,2346,1005,1043]" lastBlockId="4.[2303,2347,998,1043]" pageId="4" pageNumber="170">AB, (1)</paragraph>
|
||
<paragraph id="8BC3D6343620FF6886C8FBA1F69EFA5F" blockId="4.[980,2350,1117,2458]" pageId="4" pageNumber="170">
|
||
or 2.6 cm2, where A
|
||
<emphasis id="B9080A263620FF688053FBA1FAE7FB65" box="[1349,1374,1124,1162]" italics="true" pageId="4" pageNumber="170">is</emphasis>
|
||
the radius ofthe ramus width and B
|
||
<emphasis id="B9080A263620FF6882FAFBA1F7BCFB65" box="[2028,2053,1124,1162]" italics="true" pageId="4" pageNumber="170">is</emphasis>
|
||
the radius of the thickness. Given that the shear strength of living cortical bone
|
||
<emphasis id="B9080A263620FF688D4CFB5BF7CAFB2B" box="[2138,2163,1182,1220]" italics="true" pageId="4" pageNumber="170">is</emphasis>
|
||
conservatively ~102 N/cm2 (
|
||
<bibRefCitation id="EFEDABC53620FF688009FB1CFA47FB10" author="Currey, J. D." box="[1311,1534,1241,1279]" firstAuthor="Currey" journalOrPublisher="Princeton University Press, Princeton, NJ" pageId="4" pageNumber="170" refId="ref9370" refString="Currey, J. D. 2002. Bones: Structure and Mechanics. Princeton University Press, Princeton, NJ." title="Bones: Structure and Mechanics" type="book" year="2002">Currey 2002</bibRefCitation>
|
||
), about 26,000 N (i.e., 2652 kg offorce) was required to break the furcula. As seen by computed tomography, trabecular bone occupies a small portion of the normal ramus, so it was not considered in the calculations because it would have reduced the fracturing force only slightly.
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343620FF68813DFA00F9D4F678" blockId="4.[980,2350,1117,2458]" pageId="4" pageNumber="170">
|
||
Two of the pathologic furculae also show a characteristic bony callus from a healed stress fracture (
|
||
<bibRefCitation id="EFEDABC53620FF688305F9C5F8FFF9C9" author="Rothschild, B. M." box="[1555,1862,1536,1574]" firstAuthor="Rothschild" journalOrPublisher="Journal of Paleontology" pageId="4" pageNumber="170" pagination="302 - 304" part="62" refId="ref10445" refString="Rothschild, B. M. 1988. Stress fracture in a ceratopsian phalanx. Journal of Paleontology 62: 302 - 304." title="Stress fracture in a ceratopsian phalanx" type="journal article" year="1988">Rothschild 1988</bibRefCitation>
|
||
) located on the posterior side (
|
||
<figureCitation id="1347CAB13620FF68812CF9FFFB64F98F" box="[1082,1245,1594,1632]" captionStart="Fig. 10.2" captionText="Figure 10.2. Furcula of Tyrannosaurus rex include several with pathologies, including fractures (black arrows) and localized exostosis of stress fractures (white arrows). FMNH PP2081 in posterior (A) and anterior (B) views; MOR 980 in posterior (C) and anterior (D) views; TCM 2001.90.1 in posterior (E), anterior (F), and lateral (G) views. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942817" httpUri="https://zenodo.org/record/3942817/files/figure.png" pageId="4" pageNumber="170" targetBox="[428,2309,188,1283]" targetPageId="3">Fig. 10.2</figureCitation>
|
||
D, F;
|
||
<figureCitation id="1347CAB13620FF688029F9FFFA67F98F" box="[1343,1502,1594,1632]" captionStart="Fig. 10.5" captionText="Figure 10.5. Evidence for stress fracture in the furcula of TCM 2001.90.1 is the prominent callus (A), seen clearly in dorsal view (B) and in close-up showing periosteal reactive bone (C). The region is X-ray opaque because of the greater deposit of bone (D, between arrows). Scale for A in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942823" httpUri="https://zenodo.org/record/3942823/files/figure.png" pageId="4" pageNumber="170" targetBox="[429,1868,1369,3017]" targetPageId="5">Fig. 10.5</figureCitation>
|
||
). Stress fractures occur as
|
||
<emphasis id="B9080A263620FF6882A9F9FFF86BF98F" bold="true" box="[1983,2002,1594,1632]" pageId="4" pageNumber="170">a</emphasis>
|
||
result of repetitive loading on bone, which leads to mechanical failure and microfracturing (
|
||
<bibRefCitation id="EFEDABC53620FF6886F0F969FB59F93D" author="Resnick, D." box="[998,1248,1708,1746]" firstAuthor="Resnick" journalOrPublisher="W. B. Saunders, Philadelphia" pageId="4" pageNumber="170" refId="ref10424" refString="Resnick, D. 2002. Diagnosis of Bone and Joint Disorders. W. B. Saunders, Philadelphia." title="Diagnosis of Bone and Joint Disorders" type="book" year="2002">Resnick 2002</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFEDABC53620FF6881E7F969F944F93D" author="Rothschild, B. M. & Martin, L. D." box="[1265,1789,1708,1746]" firstAuthor="Rothschild" journalOrPublisher="CRC Press, Boca Raton, FL" pageId="4" pageNumber="170" 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>
|
||
;
|
||
<bibRefCitation id="EFEDABC53620FF688219F969F7F9F93D" author="Rothschild, B. M." box="[1807,2112,1708,1746]" firstAuthor="Rothschild" journalOrPublisher="Journal of Paleontology" pageId="4" pageNumber="170" pagination="302 - 304" part="62" refId="ref10445" refString="Rothschild, B. M. 1988. Stress fracture in a ceratopsian phalanx. Journal of Paleontology 62: 302 - 304." title="Stress fracture in a ceratopsian phalanx" type="journal article" year="1988">Rothschild 1988</bibRefCitation>
|
||
). R&thschild and Tanke (2005) and Rothschild and Molnar (this volume) note a high incidence of stress fractures
|
||
<emphasis id="B9080A263620FF6880EEF8E3F9A4F8A3" bold="true" box="[1528,1565,1830,1868]" pageId="4" pageNumber="170">in</emphasis>
|
||
the manual elements of tyrannosaurids. They also note, “Active resistance of prey is required to overstress the manus,” which would also essentially include the rest of the forelimb and pectoral girdle. Because a stress fracture is a partial fracture, the amount of force must be less than the maximum required to completely break the bone.
|
||
<emphasis id="B9080A263620FF688153F789FB0FF79D" bold="true" box="[1093,1206,2124,2162]" pageId="4" pageNumber="170">Again</emphasis>
|
||
, the furcula of
|
||
<materialsCitation id="3B14DC693620FF6880D3F789F95DF79D" ID-GBIF-Occurrence="2813095327" box="[1477,1764,2124,2162]" collectionCode="TCM" pageId="4" pageNumber="170" specimenCode="TCM 2001.90.1">TCM 2001.90.1</materialsCitation>
|
||
was used to calculate the force. The normal portion of the furcula corresponding to the pathological portion measures 4.5 cm dorsoventrally and 1.7 cm anteroposteriorly. Assuming the regions of the stress fracture were approximately the same dimensions, then the force must have been less than 60,080 N (6128 kg of pressure), resulting from equation 1.
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343620FF6886C0F63AFBD7F5CB" blockId="4.[982,1134,2552,2599]" box="[982,1134,2559,2596]" pageId="4" pageNumber="170">
|
||
<heading id="D08B61583620FF6886C0F63AFBD7F5CB" bold="true" box="[982,1134,2559,2596]" fontSize="9" level="11" pageId="4" pageNumber="170" reason="2">Scapula</heading>
|
||
</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">
|
||
T.
|
||
<emphasis id="B9080A263620FF688001F596FAF4F596" box="[1303,1357,2643,2681]" italics="true" pageId="4" pageNumber="170">rex</emphasis>
|
||
</taxonomicName>
|
||
was mostly described by
|
||
<bibRefCitation id="EFEDABC53620FF688203F596F6A5F596" author="Carpenter, K. & Smith, M." box="[1813,2332,2643,2681]" editor="Tanke, D. & Carpenter, K." firstAuthor="Carpenter" journalOrPublisher="Indiana University Press, Bloomington" pageId="4" pageNumber="170" 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 information is available based on
|
||
<materialsCitation id="3B14DC693620FF6883A3F548F80BF55C" ID-GBIF-Occurrence="2813095350" box="[1717,1970,2701,2739]" collectionCode="DMNH" pageId="4" pageNumber="170" specimenCode="DMNH 2827">DMNH 2827</materialsCitation>
|
||
. The acromion
|
||
<emphasis id="B9080A263620FF688DF1F548F6B8F55C" box="[2279,2305,2701,2739]" italics="true" pageId="4" pageNumber="170">is</emphasis>
|
||
a thin plate that is often damaged or lost in other specimens (e.g.,
|
||
<materialsCitation id="3B14DC693620FF688D41F50DF6A9F501" ID-GBIF-Occurrence="2813095323" box="[2135,2320,2760,2798]" collectionCode="MOR" pageId="4" pageNumber="170" specimenCode="MOR 555">MOR 555</materialsCitation>
|
||
). Fortunately, this region
|
||
<emphasis id="B9080A263620FF68808CF4C6FA0AF4C6" box="[1434,1459,2819,2857]" italics="true" pageId="4" pageNumber="170">is</emphasis>
|
||
preserved in
|
||
<materialsCitation id="3B14DC693620FF6883AAF4C6F803F4C6" ID-GBIF-Occurrence="2813095310" box="[1724,1978,2819,2857]" collectionCode="DMNH" pageId="4" pageNumber="170" specimenCode="DMNH 2827">DMNH 2827</materialsCitation>
|
||
and shows a small facet for the epicleidium near the scapulocoracoid suture (
|
||
<figureCitation id="1347CAB13620FF688D1AF4F8F716F48C" box="[2060,2223,2877,2915]" captionStart="Fig. 10.3" captionText="Figure 10.3. Close-up of the epicleidial facet (arrow) on the dorsal edge of the scapula DMNH 2827 (A) and lateral view with the furcula articulated (B)." figureDoi="http://doi.org/10.5281/zenodo.3942819" httpUri="https://zenodo.org/record/3942819/files/figure.png" pageId="4" pageNumber="170" targetBox="[437,1808,1361,2928]" targetPageId="3">Fig. 10.3</figureCitation>
|
||
). This epicleidial facet measures 4.8 cm by 1.1 cm. Placement of the furcula connecting the epicleidial facets of the left and right scapula show how close together the coracoids really were in life (
|
||
<figureCitation id="1347CAB13623FF6B81CDFF1EFAC0FEEE" box="[1243,1401,219,257]" captionStart="Fig. 10.4" captionText="Figure 10.4. Reconstruction of the pectoral girdle and forelimb of Tyrannosaurus showing (A) the distribution of force from the scapula (a arrows), through the furcula (b arrows), which results in cumulative force (c arrow) at the middle of the furcula. Resisting force c explains why the furcula is deepest at the midline, which is unlike any other theropod furcula. The range of motion for the forelimb segments (B), angle represented by arc a = 40°, arc b = 60°, arc c = 67°. " figureDoi="http://doi.org/10.5281/zenodo.3942821" httpUri="https://zenodo.org/record/3942821/files/figure.png" pageId="7" pageNumber="173" targetBox="[437,1808,1361,2928]" targetPageId="5">Fig. 10.4</figureCitation>
|
||
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">
|
||
<emphasis id="B9080A263623FF6B812AFED0FAFCFED4" box="[1084,1349,277,315]" italics="true" pageId="7" pageNumber="173">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
chest region found in situ that
|
||
<emphasis id="B9080A263623FF6B8726FE95FDF0FE99" box="[560,585,336,374]" italics="true" pageId="7" pageNumber="173">is</emphasis>
|
||
currently under study (
|
||
<bibRefCitation id="EFEDABC53623FF6B8100FE95FA50FE99" author="Lipkin, C. & Sereno, P. C." box="[1046,1513,336,374]" firstAuthor="Lipkin" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="7" pageNumber="173" pagination="83 A" part="24 (Suppl. to 3)" refId="ref9960" refString="Lipkin, C., and Sereno, P. C. 2004. The furcula in Tyrannosaurus rex. Journal of Vertebrate Paleontology 24 (Suppl. to 3): 83 A." title="The furcula in Tyrannosaurus rex" type="journal article" year="2004">Lipkin and Sereno 2004</bibRefCitation>
|
||
). A similar close placement is also known in hadrosaurs (e.g., Osborn 1912), suggesting that coracoids were closely placed in all dinosaurs (including sauropods), as has been discussed elsewhere (Carpenter 2002;
|
||
<bibRefCitation id="EFEDABC53623FF6B8014FDC5F935FDC9" author="Carpenter, K. & Madsen, J. & Lewis, A." box="[1282,1676,512,550]" editor="Leiggi, P. & May, P." etAl="et al." firstAuthor="Carpenter" journalOrPublisher="Cambridge University Press, New York" pageId="7" pageNumber="173" pagination="285 - 322" refId="ref9161" refString="Carpenter, K., Madsen, J., and Lewis, A. 1994. Mounting of fossil vertebrate skeletons. P. 285 - 322 in Leiggi, P., and May, P. (eds.). Vertebrate Paleontological Techniques. Cambridge University Press, New York." title="Mounting of fossil vertebrate skeletons" type="book" volumeTitle="Vertebrate Paleontological Techniques" year="1994">Carpenter et al. 1994</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="DF0386BC3621FF698264FB45F6BAF77C" ID-DOI="http://doi.org/10.5281/zenodo.3942821" ID-Zenodo-Dep="3942821" httpUri="https://zenodo.org/record/3942821/files/figure.png" pageId="5" pageNumber="171" targetBox="[462,2057,178,1135]" targetPageId="5">
|
||
<paragraph id="8BC3D6343621FF698264FB45F6BAF77C" blockId="5.[1898,2354,1152,2195]" pageId="5" pageNumber="171">
|
||
Figure 10.4.
|
||
<emphasis id="B9080A263621FF698D5AFB45F80AFB37" italics="true" pageId="5" pageNumber="171">Reconstruction</emphasis>
|
||
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>
|
||
showing (A)
|
||
</emphasis>
|
||
the distribution of force from the
|
||
<emphasis id="B9080A263621FF698D04FA4CF715FA43" box="[2066,2220,1417,1452]" italics="true" pageId="5" pageNumber="171">scapula (</emphasis>
|
||
a
|
||
<emphasis id="B9080A263621FF698DDDFA4CF86DFA0E" italics="true" pageId="5" pageNumber="171">arrows),</emphasis>
|
||
through the furcula
|
||
<emphasis id="B9080A263621FF6982D7FA36F874F9F9" box="[1985,1997,1523,1558]" italics="true" pageId="5" pageNumber="171">(</emphasis>
|
||
b arrows
|
||
<emphasis id="B9080A263621FF698D72FA36F7CFF9F9" box="[2148,2166,1523,1558]" italics="true" pageId="5" pageNumber="171">),</emphasis>
|
||
which results
|
||
<emphasis id="B9080A263621FF6982FDF9ECF7B3F9A3" box="[2027,2058,1577,1612]" italics="true" pageId="5" pageNumber="171">in</emphasis>
|
||
cumulative force
|
||
<emphasis id="B9080A263621FF6982C2F99BF859F96E" box="[2004,2016,1630,1665]" italics="true" pageId="5" pageNumber="171">(</emphasis>
|
||
c arrow
|
||
<emphasis id="B9080A263621FF698D74F99BF7D7F96E" box="[2146,2158,1630,1665]" italics="true" pageId="5" pageNumber="171">)</emphasis>
|
||
at the middle of the furcula.
|
||
<emphasis id="B9080A263621FF698278F90DF7B1F904" box="[1902,2056,1736,1771]" italics="true" pageId="5" pageNumber="171">Resisting</emphasis>
|
||
force
|
||
<emphasis id="B9080A263621FF698D61F90DF699F904" box="[2167,2336,1736,1771]" italics="true" pageId="5" pageNumber="171">
|
||
<emphasis id="B9080A263621FF698D61F90DF732F904" bold="true" box="[2167,2187,1736,1771]" italics="true" pageId="5" pageNumber="171">c</emphasis>
|
||
explains
|
||
</emphasis>
|
||
why the furcula
|
||
<emphasis id="B9080A263621FF698D9CF938F81BF8B9" italics="true" pageId="5" pageNumber="171">is deepest</emphasis>
|
||
at the
|
||
<emphasis id="B9080A263621FF698D0BF8F6F71AF8B9" box="[2077,2211,1843,1878]" italics="true" pageId="5" pageNumber="171">midline,</emphasis>
|
||
which
|
||
<emphasis id="B9080A263621FF69827DF8A2F7FCF865" box="[1899,2117,1895,1930]" italics="true" pageId="5" pageNumber="171">is unlike any</emphasis>
|
||
other
|
||
<emphasis id="B9080A263621FF698DA0F8A2F80AF850" italics="true" pageId="5" pageNumber="171">theropod</emphasis>
|
||
furcula. The range of motion for the forelimb segments
|
||
<emphasis id="B9080A263621FF698D34F7C3F782F7B0" italics="true" pageId="5" pageNumber="171">(B), angle represented by</emphasis>
|
||
arc a
|
||
<emphasis id="B9080A263621FF698DB2F7F9F81BF77C" italics="true" pageId="5" pageNumber="171">= 40°, arc</emphasis>
|
||
b = 60°, arc c = 67°.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF0386BC3621FF69827AF6D3F783F45B" ID-DOI="http://doi.org/10.5281/zenodo.3942823" ID-Zenodo-Dep="3942823" httpUri="https://zenodo.org/record/3942823/files/figure.png" pageId="5" pageNumber="171" targetBox="[429,1868,1369,3017]" targetPageId="5">
|
||
<paragraph id="8BC3D6343621FF69827AF6D3F783F45B" blockId="5.[1896,2333,2326,2996]" pageId="5" pageNumber="171">
|
||
Figure 10.5.
|
||
<emphasis id="B9080A263621FF698D50F6D3F767F6D6" box="[2118,2270,2326,2361]" italics="true" pageId="5" pageNumber="171">Evidence</emphasis>
|
||
for stress fracture
|
||
<emphasis id="B9080A263621FF698D7BF68EF768F681" box="[2157,2257,2379,2414]" italics="true" pageId="5" pageNumber="171">in the</emphasis>
|
||
furcula of
|
||
<materialsCitation id="3B14DC693621FF6982F9F645F74CF64C" ID-GBIF-Occurrence="2813095340" box="[2031,2293,2432,2467]" collectionCode="TCM" pageId="5" pageNumber="171" specimenCode="TCM 2001.90.1">
|
||
TCM
|
||
<emphasis id="B9080A263621FF698D51F645F74CF64C" box="[2119,2293,2432,2467]" italics="true" pageId="5" pageNumber="171">2001.90.1</emphasis>
|
||
</materialsCitation>
|
||
<emphasis id="B9080A263621FF698C12F645F6A4F64C" box="[2308,2333,2432,2467]" italics="true" pageId="5" pageNumber="171">is</emphasis>
|
||
the prominent callus
|
||
<emphasis id="B9080A263621FF698DC8F670F750F637" box="[2270,2281,2485,2520]" italics="true" pageId="5" pageNumber="171">(</emphasis>
|
||
A
|
||
<emphasis id="B9080A263621FF698C15F670F800F5E2" italics="true" pageId="5" pageNumber="171">), seen</emphasis>
|
||
clearly
|
||
<emphasis id="B9080A263621FF698D57F62FF7E7F5E2" box="[2113,2142,2538,2573]" italics="true" pageId="5" pageNumber="171">in</emphasis>
|
||
dorsal
|
||
<emphasis id="B9080A263621FF69827DF5DAF84DF5AD" box="[1899,2036,2591,2626]" italics="true" pageId="5" pageNumber="171">view (B)</emphasis>
|
||
and
|
||
<emphasis id="B9080A263621FF698D5AF5DAF6B0F5AD" box="[2124,2313,2591,2626]" italics="true" pageId="5" pageNumber="171">in close-up</emphasis>
|
||
showing
|
||
<emphasis id="B9080A263621FF698D1EF591F81FF543" italics="true" pageId="5" pageNumber="171">periosteal reactive</emphasis>
|
||
bone
|
||
<emphasis id="B9080A263621FF698D0CF54CF79CF543" box="[2074,2085,2697,2732]" italics="true" pageId="5" pageNumber="171">(</emphasis>
|
||
C
|
||
<emphasis id="B9080A263621FF698D28F54CF7F4F543" box="[2110,2125,2697,2732]" italics="true" pageId="5" pageNumber="171">).</emphasis>
|
||
The region
|
||
<emphasis id="B9080A263621FF69827FF57BF852F50E" box="[1897,2027,2750,2785]" italics="true" pageId="5" pageNumber="171">is X-ray</emphasis>
|
||
opaque because of the greater
|
||
<emphasis id="B9080A263621FF698D71F535F751F4FA" box="[2151,2280,2800,2837]" italics="true" pageId="5" pageNumber="171">deposit</emphasis>
|
||
of bone
|
||
<emphasis id="B9080A263621FF6982D9F4E2F843F4A5" box="[1999,2042,2855,2890]" italics="true" pageId="5" pageNumber="171">(D,</emphasis>
|
||
between
|
||
<emphasis id="B9080A263621FF698DA4F4E2F781F490" italics="true" pageId="5" pageNumber="171">arrows). Scale</emphasis>
|
||
for A in centimeters.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF0386BC3623FF6B826CFF1EF778FDD5" ID-DOI="http://doi.org/10.5281/zenodo.3942825" ID-Zenodo-Dep="3942825" httpUri="https://zenodo.org/record/3942825/files/figure.png" pageId="7" pageNumber="173" targetBox="[234,2497,25,3292]" targetPageId="6">
|
||
<paragraph id="8BC3D6343623FF6B826CFF1EF778FDD5" blockId="7.[1910,2361,212,570]" pageId="7" pageNumber="173">
|
||
<emphasis id="B9080A263623FF6B826CFF1EF85AFF11" box="[1914,2019,219,254]" italics="true" pageId="7" pageNumber="173">Figure</emphasis>
|
||
10.6.
|
||
<emphasis id="B9080A263623FF6B8D45FF1EF765FF11" box="[2131,2268,219,254]" italics="true" pageId="7" pageNumber="173">Position</emphasis>
|
||
of the furcula
|
||
<emphasis id="B9080A263623FF6B8D57FECAF707FEDD" box="[2113,2238,271,306]" italics="true" pageId="7" pageNumber="173">relative</emphasis>
|
||
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">
|
||
<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>
|
||
cast of
|
||
<emphasis id="B9080A263623FF6B8261FE27F793FDEA" box="[1911,2090,482,517]" italics="true" pageId="7" pageNumber="173">
|
||
<materialsCitation id="3B14DC693623FF6B8261FE27F7A1FDEA" ID-GBIF-Occurrence="2813095336" box="[1911,2072,482,517]" collectionCode="BHI" pageId="7" pageNumber="173" specimenCode="BHI 3033">BHI3033</materialsCitation>
|
||
).
|
||
</emphasis>
|
||
Human
|
||
<emphasis id="B9080A263623FF6B8DD0FE27F7A4FDD5" italics="true" pageId="7" pageNumber="173">(Neal L. Larson)</emphasis>
|
||
for scale.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BC3D6343623FF648731FDFEF7F4FD22" blockId="7.[462,1829,212,1200]" lastBlockId="8.[985,2346,262,721]" lastPageId="8" lastPageNumber="174" pageId="7" pageNumber="173">
|
||
<materialsCitation id="3B14DC693623FF6B8731FDFEFCA7FD8E" ID-GBIF-Occurrence="2813095345" box="[551,798,571,609]" collectionCode="DMNH" pageId="7" pageNumber="173" specimenCode="DMNH 2827">DMNH 2827</materialsCitation>
|
||
also shows an unusual pathology of the glenoid, which is partially collapsed as a result of ventroposterior rotation of the coracoid. Although the glenoid was partially damaged during preparation (the bone in the region was crumbly), it is clear that the 2 bones were initially damaged before co-ossification because the 2 bones are now firmly fused by remodeled bone (
|
||
<figureCitation id="1347CAB13623FF6B878AFCA5FC83FC69" box="[668,826,864,902]" captionStart="Fig. 10.7" captionText="Figure 10.7. Partial collapse of the glenoid in DMNH 2827 as seen in lateral view (A), with close-up (B); in medial view (C), with close-up (D); and ventral view showing the telescoping that occurred between the arrows. Sclerotic bone overhangs the lateral surface. The amount of deformation decreases dorsally to about the level of the coracoid foremen and indicates a posteroventral rotation of the coracoid due to great stress. " figureDoi="http://doi.org/10.5281/zenodo.3942827" httpUri="https://zenodo.org/record/3942827/files/figure.png" pageId="7" pageNumber="173" targetBox="[464,1829,1266,3030]" targetPageId="7">Fig. 10.7</figureCitation>
|
||
). 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>
|
||
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.
|
||
</paragraph>
|
||
<caption id="DF0386BC3623FF6B8262FAC1F86FF70E" ID-DOI="http://doi.org/10.5281/zenodo.3942827" ID-Zenodo-Dep="3942827" httpUri="https://zenodo.org/record/3942827/files/figure.png" pageId="7" pageNumber="173" startId="7.[1908,2014,1284,1319]" targetBox="[464,1829,1266,3030]" targetPageId="7">
|
||
<paragraph id="8BC3D6343623FF6B8262FAC1F86FF70E" blockId="7.[1901,2355,1278,2273]" pageId="7" pageNumber="173">
|
||
Figure 10.7. Partial
|
||
<emphasis id="B9080A263623FF6B8DD4FAC1F875FAB3" italics="true" pageId="7" pageNumber="173">collapse</emphasis>
|
||
of the
|
||
<emphasis id="B9080A263623FF6B8D5FFAFCF74EFAB3" box="[2121,2295,1337,1372]" italics="true" pageId="7" pageNumber="173">glenoid in</emphasis>
|
||
<materialsCitation id="3B14DC693623FF6B8265FAABF7E9FA7E" ID-GBIF-Occurrence="2813095334" box="[1907,2128,1390,1425]" collectionCode="DMNH" pageId="7" pageNumber="173" specimenCode="DMNH 2827">
|
||
DMNH
|
||
<emphasis id="B9080A263623FF6B82E1FAABF7E9FA7E" box="[2039,2128,1390,1425]" italics="true" pageId="7" pageNumber="173">2827</emphasis>
|
||
</materialsCitation>
|
||
as
|
||
<emphasis id="B9080A263623FF6B8D9FFAABF6BCFA7E" box="[2185,2309,1390,1425]" italics="true" pageId="7" pageNumber="173">seen in</emphasis>
|
||
lateral
|
||
<emphasis id="B9080A263623FF6B82FAFA66F7EBFA29" box="[2028,2130,1443,1478]" italics="true" pageId="7" pageNumber="173">view (</emphasis>
|
||
A
|
||
<emphasis id="B9080A263623FF6B8D7DFA66F7C4FA29" box="[2155,2173,1443,1478]" italics="true" pageId="7" pageNumber="173">),</emphasis>
|
||
with
|
||
<emphasis id="B9080A263623FF6B8264FA1DF7CAFA14" box="[1906,2163,1496,1531]" italics="true" pageId="7" pageNumber="173">close-up (B); in</emphasis>
|
||
medial view
|
||
<emphasis id="B9080A263623FF6B82D9F9C8F863F9DF" box="[1999,2010,1549,1584]" italics="true" pageId="7" pageNumber="173">(</emphasis>
|
||
C
|
||
<emphasis id="B9080A263623FF6B82E5F9C8F7BDF9DF" box="[2035,2052,1549,1584]" italics="true" pageId="7" pageNumber="173">),</emphasis>
|
||
with
|
||
<emphasis id="B9080A263623FF6B8D7AF9C8F814F98A" italics="true" pageId="7" pageNumber="173">close-up (D);</emphasis>
|
||
and ventral
|
||
<emphasis id="B9080A263623FF6B8D9FF987F7BFF975" italics="true" pageId="7" pageNumber="173">view showing</emphasis>
|
||
the
|
||
<emphasis id="B9080A263623FF6B8D43F9B2F6A5F975" box="[2133,2332,1655,1690]" italics="true" pageId="7" pageNumber="173">telescoping</emphasis>
|
||
that occurred between the arrows. Sclerotic bone overhangs the lateral surface.
|
||
<emphasis id="B9080A263623FF6B8D45F889F728F880" box="[2131,2193,1868,1903]" italics="true" pageId="7" pageNumber="173">The</emphasis>
|
||
amount of deformation decreases
|
||
<emphasis id="B9080A263623FF6B8266F873F843F836" box="[1904,2042,1974,2009]" italics="true" pageId="7" pageNumber="173">dorsally</emphasis>
|
||
to about the level of the coracoid foremen and indicates a
|
||
<emphasis id="B9080A263623FF6B8DCEF7E5F796F797" italics="true" pageId="7" pageNumber="173">posteroventral</emphasis>
|
||
rotation of the coracoid due to great stress.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF0386BC362CFF6484D0FE55FCA6FB77" ID-DOI="http://doi.org/10.5281/zenodo.3942829" ID-Zenodo-Dep="3942829" httpUri="https://zenodo.org/record/3942829/files/figure.png" pageId="8" pageNumber="174" startId="8.[454,559,400,433]" targetBox="[443,2332,1275,3078]" targetPageId="8">
|
||
<paragraph id="8BC3D634362CFF6484D0FE55FCA6FB77" blockId="8.[448,900,394,1176]" pageId="8" pageNumber="174">
|
||
<emphasis id="B9080A26362CFF6484D0FE55FD96FE5E" box="[454,559,400,433]" italics="true" pageId="8" pageNumber="174">Figure</emphasis>
|
||
10.8.
|
||
<emphasis id="B9080A26362CFF6487B7FE4AFCD5FE5D" box="[673,876,399,434]" italics="true" pageId="8" pageNumber="174">Comparison</emphasis>
|
||
of normal right humerus of MOR
|
||
<emphasis id="B9080A26362CFF64874FFE3CFD25FDF3" box="[601,668,505,540]" italics="true" pageId="8" pageNumber="174">690</emphasis>
|
||
and its
|
||
<emphasis id="B9080A26362CFF648633FE3CFDF5FDBE" italics="true" pageId="8" pageNumber="174">pathological</emphasis>
|
||
left
|
||
<emphasis id="B9080A26362CFF64878DFDEBFD01FDBE" box="[667,696,558,593]" italics="true" pageId="8" pageNumber="174">in</emphasis>
|
||
anterior
|
||
<emphasis id="B9080A26362CFF6484D3FDA6FD89FD69" box="[453,560,611,646]" italics="true" pageId="8" pageNumber="174">(A, D),</emphasis>
|
||
lateral
|
||
<emphasis id="B9080A26362CFF6487A3FDA6FCA8FD69" box="[693,785,611,646]" italics="true" pageId="8" pageNumber="174">(B, E),</emphasis>
|
||
and
|
||
<emphasis id="B9080A26362CFF6484D7FD5DFDCFFD54" box="[449,630,664,699]" italics="true" pageId="8" pageNumber="174">posterior (</emphasis>
|
||
C,
|
||
<emphasis id="B9080A26362CFF6487B2FD5DFDAFFD1F" italics="true" pageId="8" pageNumber="174">F) views. Note</emphasis>
|
||
spur at dart
|
||
<emphasis id="B9080A26362CFF6487E0FD08FD44FCCA" italics="true" pageId="8" pageNumber="174">in (E), perisoteal reactive</emphasis>
|
||
bone opposite arrow
|
||
<emphasis id="B9080A26362CFF6487C3FCF2FC97FCB5" box="[725,814,823,858]" italics="true" pageId="8" pageNumber="174">in (F),</emphasis>
|
||
with
|
||
<emphasis id="B9080A26362CFF64870DFCA9FD49FC60" box="[539,752,876,911]" italics="true" pageId="8" pageNumber="174">close-up in (</emphasis>
|
||
G
|
||
<emphasis id="B9080A26362CFF64861AFCA9FDB4FC2B" italics="true" pageId="8" pageNumber="174">). Region</emphasis>
|
||
between darts
|
||
<emphasis id="B9080A26362CFF648635FC64FE41FC17" italics="true" pageId="8" pageNumber="174">
|
||
in (F)
|
||
<emphasis id="B9080A26362CFF6484D6FC10FE41FC17" bold="true" box="[448,504,981,1016]" italics="true" pageId="8" pageNumber="174">are</emphasis>
|
||
</emphasis>
|
||
shown
|
||
<emphasis id="B9080A26362CFF648794FC10FD19FC17" box="[642,672,981,1016]" italics="true" pageId="8" pageNumber="174">in</emphasis>
|
||
lateral
|
||
<emphasis id="B9080A26362CFF648635FC10FDA7FBC1" italics="true" pageId="8" pageNumber="174">view in (H)</emphasis>
|
||
and
|
||
<emphasis id="B9080A26362CFF648760FBCEFD8CFB8C" italics="true" pageId="8" pageNumber="174">in close-up in (I). See</emphasis>
|
||
<emphasis id="B9080A26362CFF648757FB85FD3FFB8C" bold="true" box="[577,646,1088,1123]" pageId="8" pageNumber="174">text</emphasis>
|
||
for
|
||
<emphasis id="B9080A26362CFF6487DDFB85FCC7FB8C" box="[715,894,1088,1123]" italics="true" pageId="8" pageNumber="174">discussion.</emphasis>
|
||
<emphasis id="B9080A26362CFF6484D6FBB0FDA0FB77" bold="true" box="[448,537,1141,1176]" pageId="8" pageNumber="174">Scale</emphasis>
|
||
<emphasis id="B9080A26362CFF648733FBB0FDFAFB77" box="[549,579,1141,1176]" italics="true" pageId="8" pageNumber="174">in</emphasis>
|
||
centimeters.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BC3D634362CFF6486CCFCF2FB35FCB3" blockId="8.[986,1164,817,860]" box="[986,1164,823,860]" pageId="8" pageNumber="174">
|
||
<heading id="D08B6158362CFF6486CCFCF2FB35FCB3" bold="true" box="[986,1164,823,860]" fontSize="13" level="8" pageId="8" pageNumber="174" reason="0">Humerus</heading>
|
||
</paragraph>
|
||
<paragraph id="8BC3D634362CFF6586CCFC4DFB4AF960" blockId="8.[983,2343,897,1181]" lastBlockId="9.[465,1828,1560,3032]" lastPageId="9" lastPageNumber="175" pageId="8" pageNumber="174">
|
||
Several additional humeri are now known, including more with pathologies that reflect behavior. Overall, these specimens resemble those described by
|
||
<bibRefCitation id="EFEDABC5362CFF64818FFC3BF920FBCB" author="Carpenter, K. & Smith, M." box="[1177,1689,1022,1060]" editor="Tanke, D. & Carpenter, K." firstAuthor="Carpenter" journalOrPublisher="Indiana University Press, Bloomington" pageId="8" pageNumber="174" 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>
|
||
, differing only in minor detail. Two of the new specimens are of gracile morphs (e.g.,
|
||
<materialsCitation id="3B14DC69362CFF64825BFBFCF7BFFBB0" ID-GBIF-Occurrence="2813095306" box="[1869,2054,1081,1119]" collectionCode="MOR" pageId="8" pageNumber="174" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
and
|
||
<materialsCitation id="3B14DC69362CFF648D48FBFCF6A8FBB0" ID-GBIF-Occurrence="2813095309" box="[2142,2321,1081,1119]" collectionCode="BHI" pageId="8" pageNumber="174" specimenCode="BHI 6230">BHI6230</materialsCitation>
|
||
), which are probably male (P. Larson this volume). The robust morph (e.g.,
|
||
<materialsCitation id="3B14DC69362DFF6584C3F9EDFD4EF9A1" ID-GBIF-Occurrence="2813095339" box="[469,759,1576,1614]" collectionCode="FMNH" pageId="9" pageNumber="175" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
) is broader proximally than the gracile morph (cf.
|
||
<bibRefCitation id="EFEDABC5362DFF658391F9EDFC84F960" author="Carpenter, K. & Smith, M." editor="Tanke, D. & Carpenter, K." firstAuthor="Carpenter" journalOrPublisher="Indiana University Press, Bloomington" pageId="9" pageNumber="175" 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>
|
||
, fig. 9.4, with
|
||
<figureCitation id="1347CAB1362DFF658156F9ACFB5BF960" box="[1088,1250,1641,1679]" captionStart="Fig. 10.8" captionText="Figure 10.8. Comparison of normal right humerus of MOR 690 and its pathological left in anterior (A, D), lateral (B, E), and posterior (C, F) views. Note spur at dart in (E), perisoteal reactive bone opposite arrow in (F), with close-up in (G). Region between darts in (F) are shown in lateral view in (H) and in close-up in (I). See text for discussion. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942829" httpUri="https://zenodo.org/record/3942829/files/figure.png" pageId="9" pageNumber="175" targetBox="[443,2332,1275,3078]" targetPageId="8">Fig. 10.8</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="DF0386BC362DFF658269FF17F7FEF965" ID-DOI="http://doi.org/10.5281/zenodo.3942831" ID-Zenodo-Dep="3942831" httpUri="https://zenodo.org/record/3942831/files/figure.png" pageId="9" pageNumber="175" startId="9.[1919,2024,210,245]" subCaptionStartIDs="9.[2215,2322,1533,1568]" subCaptionStarts="Figure 10" targetBox="[480,1829,206,1482]" targetPageId="9">
|
||
<paragraph id="8BC3D634362DFF658269FF17F7FEF965" blockId="9.[1910,2368,205,1674]" pageId="9" pageNumber="175">
|
||
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">
|
||
<emphasis id="B9080A26362DFF658D9BFEC2F841FEB1" bold="true" pageId="9" pageNumber="175">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C7CADB7362DFF658D1BFEFEF70CFEB1" box="[2061,2229,315,350]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Alligator</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
and
|
||
<taxonomicName id="4C7CADB7362DFF658268FEB5F84DFE7C" box="[1918,2036,368,403]" class="Aves" family="Phasianidae" genus="Gallus" kingdom="Animalia" order="Galliformes" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9080A26362DFF658268FEB5F84DFE7C" bold="true" box="[1918,2036,368,403]" pageId="9" pageNumber="175">Gallus</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
<emphasis id="B9080A26362DFF658D1DFEB5F807FE28" italics="true" pageId="9" pageNumber="175">Top row is anterior,</emphasis>
|
||
bottom row
|
||
<emphasis id="B9080A26362DFF658DBDFE61F807FE13" italics="true" pageId="9" pageNumber="175">is posterior.</emphasis>
|
||
Muscle map based on
|
||
<emphasis id="B9080A26362DFF6582A5FDCBF79DFDDE" box="[1971,2084,526,561]" italics="true" pageId="9" pageNumber="175">scars (</emphasis>
|
||
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>
|
||
.
|
||
</emphasis>
|
||
Map for
|
||
<emphasis id="B9080A26362DFF658DB3FD86F855FD74" italics="true" pageId="9" pageNumber="175">
|
||
<taxonomicName id="4C7CADB7362DFF658DB3FD86F801FD74" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">Alligator</taxonomicName>
|
||
(B,
|
||
</emphasis>
|
||
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>
|
||
<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>
|
||
based on deformation of
|
||
<taxonomicName id="4C7CADB7362DFF65826EFCD3F791FCD6" box="[1912,2088,790,825]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9080A26362DFF65826EFCD3F791FCD6" bold="true" box="[1912,2088,790,825]" pageId="9" pageNumber="175">Alligator</emphasis>
|
||
</taxonomicName>
|
||
humerus. Map for
|
||
<taxonomicName id="4C7CADB7362DFF6582ADFC8EF797FC81" box="[1979,2094,843,878]" class="Aves" family="Phasianidae" genus="Gallus" kingdom="Animalia" order="Galliformes" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9080A26362DFF6582ADFC8EF797FC81" bold="true" box="[1979,2094,843,878]" pageId="9" pageNumber="175">Gallus</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B9080A26362DFF658D2DFC8EF733FC81" box="[2107,2186,843,878]" italics="true" pageId="9" pageNumber="175">(D, I)</emphasis>
|
||
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>
|
||
(E, J)
|
||
</emphasis>
|
||
based on deformation of
|
||
<taxonomicName id="4C7CADB7362DFF658D25FC2FF71CFBE2" box="[2099,2213,1002,1037]" class="Aves" family="Phasianidae" genus="Gallus" kingdom="Animalia" order="Galliformes" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9080A26362DFF658D25FC2FF71CFBE2" bold="true" box="[2099,2213,1002,1037]" pageId="9" pageNumber="175">Gallus</emphasis>
|
||
</taxonomicName>
|
||
humerus.
|
||
<emphasis id="B9080A26362DFF658D3DFBDAF7C7FBAD" box="[2091,2174,1055,1090]" italics="true" pageId="9" pageNumber="175">Note</emphasis>
|
||
that
|
||
<emphasis id="B9080A26362DFF658DCFFBDAF7FEFB98" italics="true" pageId="9" pageNumber="175">predicted scars</emphasis>
|
||
for deformed
|
||
<emphasis id="B9080A26362DFF658261FB4CF785FB43" box="[1911,2108,1161,1196]" italics="true" pageId="9" pageNumber="175">
|
||
<taxonomicName id="4C7CADB7362DFF658261FB4CF79EFB43" box="[1911,2087,1161,1196]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="9" pageNumber="175" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9080A26362DFF658261FB4CF79EFB43" bold="true" box="[1911,2087,1161,1196]" italics="true" pageId="9" pageNumber="175">Alligator</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
</emphasis>
|
||
C,
|
||
<emphasis id="B9080A26362DFF658D7CFB4CF76AFB43" box="[2154,2259,1161,1196]" italics="true" pageId="9" pageNumber="175">H) are</emphasis>
|
||
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>
|
||
<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>
|
||
humerus
|
||
<emphasis id="B9080A26362DFF6582E7FA38F86CF9BA" italics="true" pageId="9" pageNumber="175">(K, L). See Figure 10.13</emphasis>
|
||
and text for further explanation.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BC3D634362DFF658733F966FAE3F4C9" blockId="9.[465,1828,1560,3032]" pageId="9" pageNumber="175">
|
||
As noted by
|
||
<bibRefCitation id="EFEDABC5362DFF658635F966FA86F926" author="Carpenter, K. & Smith, M." box="[803,1343,1699,1737]" editor="Tanke, D. & Carpenter, K." firstAuthor="Carpenter" journalOrPublisher="Indiana University Press, Bloomington" pageId="9" pageNumber="175" 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 humerus of
|
||
<materialsCitation id="3B14DC69362DFF65838EF966FDD9F8EA" ID-GBIF-Occurrence="2813095330" collectionCode="FMNH" pageId="9" pageNumber="175" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
shows several pathologies, as does the new left humerus of
|
||
<materialsCitation id="3B14DC69362DFF6583ADF91AFDAEF8D0" ID-GBIF-Occurrence="2813095318" collectionCode="MOR" pageId="9" pageNumber="175" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
(e.g.,
|
||
<figureCitation id="1347CAB1362DFF65879BF8DCFC88F8D0" box="[653,817,1817,1855]" captionStart="Fig. 10.8" captionText="Figure 10.8. Comparison of normal right humerus of MOR 690 and its pathological left in anterior (A, D), lateral (B, E), and posterior (C, F) views. Note spur at dart in (E), perisoteal reactive bone opposite arrow in (F), with close-up in (G). Region between darts in (F) are shown in lateral view in (H) and in close-up in (I). See text for discussion. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942829" httpUri="https://zenodo.org/record/3942829/files/figure.png" pageId="9" pageNumber="175" targetBox="[443,2332,1275,3078]" targetPageId="8">Fig. 10.8</figureCitation>
|
||
). This latter humerus shows an extensive juxtacortical pathology located proximally on the anterolateral, lateral, and posterolateral sides of the diaphysis, which has all but obliterated the deltopectoral crest (
|
||
<figureCitation id="1347CAB1362DFF6584F4F80CFD3AF800" box="[482,643,1993,2031]" captionStart="Fig. 10.8" captionText="Figure 10.8. Comparison of normal right humerus of MOR 690 and its pathological left in anterior (A, D), lateral (B, E), and posterior (C, F) views. Note spur at dart in (E), perisoteal reactive bone opposite arrow in (F), with close-up in (G). Region between darts in (F) are shown in lateral view in (H) and in close-up in (I). See text for discussion. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942829" httpUri="https://zenodo.org/record/3942829/files/figure.png" pageId="9" pageNumber="175" targetBox="[443,2332,1275,3078]" targetPageId="8">Fig. 10.8</figureCitation>
|
||
D, E). The pathology shows extreme disruption ofthe cortical bone and irregular patches of sclerotic bone that suggest uneven loss and regrowth of the periosteum. The resulting periostitis implies extensive inflammation of the region. The region of the pathology corresponds with the crocodile humerus to the insertions ofthe M. pectoralis, the common insertion for the M. supracoracoideus complex, M. deltoideus clavicularis, common insertion for the M. teres major and M. latisimus dorsi, and origins of the M. brachialis, M. humeroradialis, and part of the triceps brevis cranialis (cf.
|
||
<figureCitation id="1347CAB1362DFF658349F6A0F8BFF664" box="[1631,1798,2405,2443]" captionStart="Fig. 10.9" captionText="Figure 10.9. Muscle maps for humerus in Tyrannosaurus, Alligator, and Gallus. Top row is anterior, bottom row is posterior. Muscle map based on scars (A, F) Tyrannosaurus. Map for Alligator (B, G) and predicted for Tyrannosaurus (C, D) based on deformation of Alligator humerus. Map for Gallus (D, I) and predicted for Tyrannosaurus (E, J) based on deformation of Gallus humerus. Note that predicted scars for deformed Alligator (C, H) are a better match for the scars of Tyrannosaurus (A, F). This prediction is also supported by the pattern ofavulsion seen in a Tyrannosaurus humerus (K, L). See Figure 10.13 and text for further explanation. " figureDoi="http://doi.org/10.5281/zenodo.3942831" httpUri="https://zenodo.org/record/3942831/files/figure.png" pageId="9" pageNumber="175" targetBox="[480,1829,206,1482]" targetPageId="9">Figs. 10.9</figureCitation>
|
||
B with
|
||
<figureCitation id="1347CAB1362DFF658721F665FDC6F629" box="[567,639,2464,2502]" captionStart="Fig. 10.9" captionText="Figure 10.9. Muscle maps for humerus in Tyrannosaurus, Alligator, and Gallus. Top row is anterior, bottom row is posterior. Muscle map based on scars (A, F) Tyrannosaurus. Map for Alligator (B, G) and predicted for Tyrannosaurus (C, D) based on deformation of Alligator humerus. Map for Gallus (D, I) and predicted for Tyrannosaurus (E, J) based on deformation of Gallus humerus. Note that predicted scars for deformed Alligator (C, H) are a better match for the scars of Tyrannosaurus (A, F). This prediction is also supported by the pattern ofavulsion seen in a Tyrannosaurus humerus (K, L). See Figure 10.13 and text for further explanation. " figureDoi="http://doi.org/10.5281/zenodo.3942831" httpUri="https://zenodo.org/record/3942831/files/figure.png" pageId="9" pageNumber="175" targetBox="[480,1829,206,1482]" targetPageId="9">10.9</figureCitation>
|
||
K, G with L). There is also a juxtacortical lesion on the posterior side that, in the crocodile, corresponds with the origin for the proximal part of the M. triceps brevis intermedius (
|
||
<figureCitation id="1347CAB1362DFF65819AF5D0FA94F5D4" box="[1164,1325,2581,2619]" captionStart="Fig. 10.8" captionText="Figure 10.8. Comparison of normal right humerus of MOR 690 and its pathological left in anterior (A, D), lateral (B, E), and posterior (C, F) views. Note spur at dart in (E), perisoteal reactive bone opposite arrow in (F), with close-up in (G). Region between darts in (F) are shown in lateral view in (H) and in close-up in (I). See text for discussion. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942829" httpUri="https://zenodo.org/record/3942829/files/figure.png" pageId="9" pageNumber="175" targetBox="[443,2332,1275,3078]" targetPageId="8">Fig. 10.8</figureCitation>
|
||
F, G; compare 9G with L). More distally, there is a small spur on the posterior side as well (
|
||
<figureCitation id="1347CAB1362DFF65834DF595F945F599" box="[1627,1788,2640,2678]" captionStart="Fig. 10.8" captionText="Figure 10.8. Comparison of normal right humerus of MOR 690 and its pathological left in anterior (A, D), lateral (B, E), and posterior (C, F) views. Note spur at dart in (E), perisoteal reactive bone opposite arrow in (F), with close-up in (G). Region between darts in (F) are shown in lateral view in (H) and in close-up in (I). See text for discussion. Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942829" httpUri="https://zenodo.org/record/3942829/files/figure.png" pageId="9" pageNumber="175" targetBox="[443,2332,1275,3078]" targetPageId="8">Fig. 10.8</figureCitation>
|
||
E, F, H, I). These 2 lesions blend with the diaphysis proximally but have sharp, overhanging margins distally that may be due to osteoblastic response to the extensor motion of the overlying M. triceps group.
|
||
</paragraph>
|
||
<paragraph id="8BC3D634362DFF668732F4FFF956FA0D" blockId="9.[465,1828,1560,3032]" lastBlockId="10.[978,2345,227,1509]" lastPageId="10" lastPageNumber="176" pageId="9" pageNumber="175">
|
||
The large pathology on the proximal end is irregular or knobby. Superfacially, it suggests a malignant neoplasm or extensive osteomyelitis, but
|
||
<bibRefCitation id="EFEDABC5362DFF6583E8F4B0FCE6F439" author="Donnelly, L. F. & Helms, C. A. & Bisset, G. S." etAl="et al." firstAuthor="as Donnelly" journalOrPublisher="Radiology" pageId="9" pageNumber="175" pagination="233" part="211" refId="ref9470" refString="Donnelly, L. F., Helms, C. A., and Bisset, G. S. 1999. Chronic avulsive injury of the deltoid insertion in adolescents: imaging findings in three cases. Radiology 211: 233." title="Chronic avulsive injury of the deltoid insertion in adolescents: imaging findings in three cases" type="journal article" year="1999">as Donnelly et al. (1999)</bibRefCitation>
|
||
have noted, site location is important for differential diagnosis. The location and extent of the pathology is most probably due to avulsion of muscles in the vicinity of the deltopectoral crest, and the posterior pathologies due to periostitis from trauma to the periosteum associated with the event that caused the avulsion. The aggressive appearance of the proximal pathology, caused by bone resorption resulting in a lytic appearance of bone, is characteristic of a healing avulsion and can mimic the appearance of osteomyelitis or skeletal Ewing sarcoma (
|
||
<bibRefCitation id="EFEDABC5362EFF66829EFD8FF76BFD9F" author="Stevens, AL A. & El-Khoury, G. Y. & Kathol, M. H. & Brandser, E. A. & Chow, S." box="[1928,2258,586,624]" etAl="et al." firstAuthor="Stevens" journalOrPublisher="Radiographics" pageId="10" pageNumber="176" pagination="655 - 672" part="19" refId="ref10672" refString="Stevens, AL A., El-Khoury, G. Y., Kathol, M. H., Brandser, E. A., and Chow, S. 1999. Imaging features of avulsion injuries. Radiographics 19: 655 - 672." title="Imaging features of avulsion injuries" type="journal article" year="1999">Stevens et al. 1999</bibRefCitation>
|
||
). An avulsion is a failure of bone at a tendinous or aponeurotic insertion of muscle (
|
||
<bibRefCitation id="EFEDABC5362EFF6686F3FD05FA81FD09" author="Tehranzadeh, J." box="[997,1336,704,742]" firstAuthor="Tehranzadeh" journalOrPublisher="Radiographics" pageId="10" pageNumber="176" pagination="945 - 974" part="7" refId="ref10719" refString="Tehranzadeh, J. 1987. The spectrum of avulsion and avulsion-like injuries of the musculoskeletal system. Radiographics 7: 945 - 974." title="The spectrum of avulsion and avulsion-like injuries of the musculoskeletal system" type="journal article" year="1987">Tehranzadeh 1987</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFEDABC5362EFF668058FD05F959FD09" author="El-Khoury, G. Y. & Daniel, W. W. & Kathol, M. H." box="[1358,1760,704,742]" etAl="et al." firstAuthor="El-Khoury" journalOrPublisher="Radiological Clines of North America" pageId="10" pageNumber="176" pagination="747 - 766" part="35" refId="ref9514" refString="El-Khoury, G. Y., Daniel, W. W., and Kathol, M. H. 1997. Acute and chronic avulsive injuries. Radiological Clines of North America 35: 747 - 766." title="Acute and chronic avulsive injuries" type="journal article" year="1997">El-Khoury et al. 1997</bibRefCitation>
|
||
). Avulsions can either be acute (the result of extreme, abnormal muscle contractions) or chronic (the result of repeated microtrauma or overuse) (
|
||
<bibRefCitation id="EFEDABC5362EFF668386FCF0F854FCB4" author="Stevens, AL A. & El-Khoury, G. Y. & Kathol, M. H. & Brandser, E. A. & Chow, S." box="[1680,2029,821,859]" etAl="et al." firstAuthor="Stevens" journalOrPublisher="Radiographics" pageId="10" pageNumber="176" pagination="655 - 672" part="19" refId="ref10672" refString="Stevens, AL A., El-Khoury, G. Y., Kathol, M. H., Brandser, E. A., and Chow, S. 1999. Imaging features of avulsion injuries. Radiographics 19: 655 - 672." title="Imaging features of avulsion injuries" type="journal article" year="1999">Stevens et al. 1999</bibRefCitation>
|
||
), where reoccurrence of the injury is more frequent than the ability of the tissue to repair itself (
|
||
<bibRefCitation id="EFEDABC5362EFF66813DFC6EFA0BFC3E" author="El-Khoury, G. Y. & Daniel, W. W. & Kathol, M. H." box="[1067,1458,939,977]" etAl="et al." firstAuthor="El-Khoury" journalOrPublisher="Radiological Clines of North America" pageId="10" pageNumber="176" pagination="747 - 766" part="35" refId="ref9514" refString="El-Khoury, G. Y., Daniel, W. W., and Kathol, M. H. 1997. Acute and chronic avulsive injuries. Radiological Clines of North America 35: 747 - 766." title="Acute and chronic avulsive injuries" type="journal article" year="1997">El-Khoury et al. 1997</bibRefCitation>
|
||
). The presence of a stress fracture in the furcula associated with this specimen suggests that repetitive overuse of the forelimb may have been a major factor leading to the avulsion. A secondary factor may have been a violent stress on the arm because so many different muscles were apparently affected. Such stresses would be generated in the sudden pull of the arm toward the chest at the same time prey was struggling in the opposite direction. It may not be incidental that
|
||
<materialsCitation id="3B14DC69362EFF66838AFAC9F8EDFADD" ID-GBIF-Occurrence="2813095333" box="[1692,1876,1292,1330]" collectionCode="MOR" pageId="10" pageNumber="176" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
is a young adult because
|
||
<bibRefCitation id="EFEDABC5362EFF6686C4FA82FA8EFA82" author="Tehranzadeh, J." box="[978,1335,1351,1389]" firstAuthor="Tehranzadeh" journalOrPublisher="Radiographics" pageId="10" pageNumber="176" pagination="945 - 974" part="7" refId="ref10719" refString="Tehranzadeh, J. 1987. The spectrum of avulsion and avulsion-like injuries of the musculoskeletal system. Radiographics 7: 945 - 974." title="The spectrum of avulsion and avulsion-like injuries of the musculoskeletal system" type="journal article" year="1987">Tehranzadeh (1987)</bibRefCitation>
|
||
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>
|
||
.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph id="8BC3D634362EFF6686C7F98EFBECF99F" blockId="10.[977,1109,1604,1648]" box="[977,1109,1611,1648]" pageId="10" pageNumber="176">
|
||
<heading id="D08B6158362EFF6686C7F98EFBECF99F" bold="true" box="[977,1109,1611,1648]" fontSize="13" level="8" pageId="10" pageNumber="176" reason="0">
|
||
<emphasis id="B9080A26362EFF6686C7F98EFBECF99F" bold="true" box="[977,1109,1611,1648]" pageId="10" pageNumber="176">Manus</emphasis>
|
||
</heading>
|
||
</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>
|
||
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,
|
||
<materialsCitation id="3B14DC69362EFF6680E9F91CF90CF910" ID-GBIF-Occurrence="2813095348" box="[1535,1717,1753,1791]" collectionCode="MOR" pageId="10" pageNumber="176" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
and
|
||
<materialsCitation id="3B14DC69362EFF668206F91CF87BF910" ID-GBIF-Occurrence="2813095335" box="[1808,1986,1753,1791]" collectionCode="BHI" pageId="10" pageNumber="176" specimenCode="BHI 6230">BHI 6230</materialsCitation>
|
||
, provide new information, including a carpal and metacarpal III (
|
||
<figureCitation id="1347CAB1362EFF66821BF8D1F870F8D5" box="[1805,1993,1812,1850]" captionStart="Fig. 10.10" captionText="Figure 10.10. Distal carpal (BHI6230) of Tyrannosaurus in multiple views: proximal or dorsal (A); distal or ventral (B); anterior (C); posterior (D); extensor side (E); palmar side (F). Metacarpal III of BHI 6230 in lateral (G) and extensor side (H). Metacarpal III of MOR 690 in lateral (I) and extensor side (J). Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942833" httpUri="https://zenodo.org/record/3942833/files/figure.png" pageId="10" pageNumber="176" targetBox="[457,1819,194,2046]" targetPageId="11">Figs. 10.10</figureCitation>
|
||
,
|
||
<figureCitation id="1347CAB1362EFF6682F1F8D1F782F8D5" box="[2023,2107,1812,1850]" captionStart="Fig. 10.11" captionText="Figure 10.11. Distal carpal and metacarpals in articulation (BHI 6230). Proximal (A), digit I side (B), extensor side (C), digit III side (D). Scale in centimeters." figureDoi="http://doi.org/10.5281/zenodo.3942835" httpUri="https://zenodo.org/record/3942835/files/figure.png" pageId="10" pageNumber="176" targetBox="[976,2381,209,1786]" targetPageId="11">10.11</figureCitation>
|
||
). The carpal (
|
||
<figureCitation id="1347CAB1362EFF6686C8F88BFB20F89B" box="[990,1177,1870,1908]" captionStart="Fig. 10.10" captionText="Figure 10.10. Distal carpal (BHI6230) of Tyrannosaurus in multiple views: proximal or dorsal (A); distal or ventral (B); anterior (C); posterior (D); extensor side (E); palmar side (F). Metacarpal III of BHI 6230 in lateral (G) and extensor side (H). Metacarpal III of MOR 690 in lateral (I) and extensor side (J). Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942833" httpUri="https://zenodo.org/record/3942833/files/figure.png" pageId="10" pageNumber="176" targetBox="[457,1819,194,2046]" targetPageId="11">Fig. 10.10</figureCitation>
|
||
A-F) is distal carpal I (“semilunate”) and shows
|
||
<emphasis id="B9080A26362EFF668D0AF88BF789F89B" bold="true" box="[2076,2096,1870,1908]" pageId="10" pageNumber="176">a</emphasis>
|
||
facet on the proximal surface for the
|
||
<emphasis id="B9080A26362EFF66806FF84CF9BBF840" box="[1401,1538,1929,1967]" italics="true" pageId="10" pageNumber="176">missing</emphasis>
|
||
radiale. This carpal shows that the damaged, incomplete one described by
|
||
<bibRefCitation id="EFEDABC5362EFF6680F2F801F85CF805" author="Carpenter, K. & Smith, M." box="[1508,2021,1988,2026]" 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>
|
||
is in fact a distal carpal I. Ventrally, the new carpal is faceted and fits snugly between the proximal ends of metacarpals I and II (
|
||
<figureCitation id="1347CAB1362EFF668394F7FCF88FF7B0" box="[1666,1846,2105,2143]" captionStart="Fig. 10.11" captionText="Figure 10.11. Distal carpal and metacarpals in articulation (BHI 6230). Proximal (A), digit I side (B), extensor side (C), digit III side (D). Scale in centimeters." figureDoi="http://doi.org/10.5281/zenodo.3942835" httpUri="https://zenodo.org/record/3942835/files/figure.png" pageId="10" pageNumber="176" targetBox="[976,2381,209,1786]" targetPageId="11">Fig. 10.11</figureCitation>
|
||
<emphasis id="B9080A26362EFF668220F7FCF8EBF7B0" box="[1846,1874,2105,2143]" italics="true" pageId="10" pageNumber="176">C</emphasis>
|
||
), rather than across them as in Deinonychus (Carpenter 2002). Metacarpal III is slender, posteriorly curving, and tapering, and lacks a distal condyle, so it thus had no phalanges.
|
||
<materialsCitation id="3B14DC69362EFF6686D8F72FFB3BF6FF" ID-GBIF-Occurrence="2813095304" box="[974,1154,2282,2320]" collectionCode="MOR" pageId="10" pageNumber="176" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
is from a larger individual than
|
||
<materialsCitation id="3B14DC69362EFF6683ABF72FF8D7F6FF" ID-GBIF-Occurrence="2813095305" box="[1725,1902,2282,2320]" collectionCode="BHI" pageId="10" pageNumber="176" specimenCode="BHI 6230">BHI 6230</materialsCitation>
|
||
, and the differences between them are probably ontogenetic. These metacarpals suggest that as the individual grew, the metacarpal became more robust and the proximal end formed a broader attachment to metacarpal II.
|
||
</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">
|
||
<emphasis id="B9080A26362EFF6683C8F5CAF852F5DA" box="[1758,2027,2575,2613]" italics="true" pageId="10" pageNumber="176">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="1347CAB1362EFF668D10F5CAF702F5DA" box="[2054,2235,2575,2613]" captionStart="Fig. 10.12" captionText="Figure 10.12. Articulated pectoral girdle and forelimb of Tyrannosaurus in lateral (A) and anterior (B) views" figureDoi="http://doi.org/10.5281/zenodo.3942837" httpUri="https://zenodo.org/record/3942837/files/figure.png" pageId="10" pageNumber="176" targetBox="[970,2327,1946,2984]" targetPageId="12">Fig. 10.12</figureCitation>
|
||
) and was used in following biomechanical study.
|
||
</paragraph>
|
||
<paragraph id="8BC3D634362EFF6684A2F537FD7CF442" blockId="10.[436,766,2802,2989]" box="[436,765,2802,2989]" pageId="10" pageNumber="176">
|
||
<heading id="D08B6158362EFF6684A2F537FD44F4C7" bold="true" box="[436,765,2802,2856]" centered="true" fontSize="17" level="4" pageId="10" pageNumber="176" reason="0">
|
||
<emphasis id="B9080A26362EFF6684A2F537FD44F4C7" bold="true" box="[436,765,2802,2856]" pageId="10" pageNumber="176">Reconstructing</emphasis>
|
||
</heading>
|
||
<heading id="D08B6158362EFF6684A2F4F1FD7CF442" bold="true" box="[436,709,2868,2989]" fontSize="17" level="5" pageId="10" pageNumber="176" reason="0">Forelimb Musculature</heading>
|
||
</paragraph>
|
||
<paragraph id="8BC3D634362EFF6786DCF4C4FAD7F696" blockId="10.[969,2329,2810,3035]" lastBlockId="11.[456,1820,2144,3013]" lastPageId="11" lastPageNumber="177" pageId="10" pageNumber="176">
|
||
Mathematical models for scavengers not withstanding (e.g.,
|
||
<bibRefCitation id="EFEDABC5362EFF668D2CF4C4FB76F48E" author="Ruxton. G. D. & Houston. D. C." firstAuthor="Ruxton" journalOrPublisher="Proceedings: Biological Sciences" pageId="10" pageNumber="176" pagination="731 - 733" part="270" refId="ref10545" refString="Ruxton. G. D., and Houston. D. C. 2003. Could Tyrannosaurus rex have been a scavenger rather than a predator? An energetics approach. Proceedings: Biological Sciences 270: 731 - 733." title="Could Tyrannosaurus rex have been a scavenger rather than a predator? An energetics approach" type="journal article" year="2003">Ruxton and Houston 2003</bibRefCitation>
|
||
), many of the same criteria and assumptions for scavenging theropods also hold for predatory theropods as well (see Holtz this volume). In reality, there are too many unknown variables (e.g., population densities, energentics, locomotion capabilities of both predator and prey) to ever have testable results. With so many assumptions built on assumptions, a house of cards results. Our approach is to minimize assumptions (including minimizing untested “common sense,” referred to by
|
||
<bibRefCitation id="EFEDABC5362FFF67809EF6DDFD9DF695" author="Ruxton. G. D. & Houston. D. C." firstAuthor="Ruxton" journalOrPublisher="Proceedings: Biological Sciences" pageId="11" pageNumber="177" pagination="731 - 733" part="270" refId="ref10545" refString="Ruxton. G. D., and Houston. D. C. 2003. Could Tyrannosaurus rex have been a scavenger rather than a predator? An energetics approach. Proceedings: Biological Sciences 270: 731 - 733." title="Could Tyrannosaurus rex have been a scavenger rather than a predator? An energetics approach" type="journal article" year="2003">Ruxton and Houston 2003</bibRefCitation>
|
||
) and to rely on evidence and testable models.
|
||
</paragraph>
|
||
<caption id="DF0386BC362FFF678263FF0EF7FBFC74" ID-DOI="http://doi.org/10.5281/zenodo.3942833" ID-Zenodo-Dep="3942833" httpUri="https://zenodo.org/record/3942833/files/figure.png" pageId="11" pageNumber="177" startId="11.[1909,2013,203,238]" targetBox="[457,1819,194,2046]" targetPageId="11">
|
||
<paragraph id="8BC3D634362FFF678263FF0EF7FBFC74" blockId="11.[1904,2359,196,923]" pageId="11" pageNumber="177">
|
||
<emphasis id="B9080A26362FFF678263FF0EF864FF01" box="[1909,2013,203,238]" italics="true" pageId="11" pageNumber="177">Figure</emphasis>
|
||
10.10.
|
||
<emphasis id="B9080A26362FFF678D76FF0EF794FECD" italics="true" pageId="11" pageNumber="177">
|
||
Distal
|
||
<emphasis id="B9080A26362FFF678DDAFF0EF68EFF01" bold="true" box="[2252,2359,203,238]" italics="true" pageId="11" pageNumber="177">carpal</emphasis>
|
||
(
|
||
<materialsCitation id="3B14DC69362FFF678297FF3AF798FECD" ID-GBIF-Occurrence="2813095308" box="[1921,2081,255,290]" collectionCode="BHI" pageId="11" pageNumber="177" specimenCode="BHI 6230">BHI6230</materialsCitation>
|
||
)
|
||
</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>
|
||
in multiple views: proximal
|
||
</emphasis>
|
||
or dorsal
|
||
<emphasis id="B9080A26362FFF678DADFEACF77EFE63" box="[2235,2247,361,396]" italics="true" pageId="11" pageNumber="177">(</emphasis>
|
||
A
|
||
<emphasis id="B9080A26362FFF678DF7FEACF74FFE63" box="[2273,2294,361,396]" italics="true" pageId="11" pageNumber="177">);</emphasis>
|
||
distal
|
||
<emphasis id="B9080A26362FFF6782CDFE58F7BBFE2F" box="[2011,2050,413,448]" italics="true" pageId="11" pageNumber="177">or</emphasis>
|
||
ventral
|
||
<emphasis id="B9080A26362FFF678D9BFE58F87FFE1B" italics="true" pageId="11" pageNumber="177">(B); anterior (</emphasis>
|
||
C
|
||
<emphasis id="B9080A26362FFF6782C9FE14F84AFE1B" box="[2015,2035,465,500]" italics="true" pageId="11" pageNumber="177">);</emphasis>
|
||
posterior
|
||
<emphasis id="B9080A26362FFF678DBFFE14F75BFE1B" box="[2217,2274,465,500]" italics="true" pageId="11" pageNumber="177">(D);</emphasis>
|
||
extensor
|
||
<emphasis id="B9080A26362FFF678D05FDC2F848FDB1" italics="true" pageId="11" pageNumber="177">side (E); palmar side (F).</emphasis>
|
||
Metacarpal
|
||
<emphasis id="B9080A26362FFF678DC4FDFEF748FDB1" box="[2258,2289,571,606]" italics="true" pageId="11" pageNumber="177">III</emphasis>
|
||
of
|
||
<materialsCitation id="3B14DC69362FFF678265FDB5F7ACFD7C" ID-GBIF-Occurrence="2813095321" box="[1907,2069,624,659]" collectionCode="BHI" pageId="11" pageNumber="177" specimenCode="BHI 6230">
|
||
<emphasis id="B9080A26362FFF678265FDB5F7ACFD7C" box="[1907,2069,624,659]" italics="true" pageId="11" pageNumber="177">BHI 6230</emphasis>
|
||
</materialsCitation>
|
||
in lateral
|
||
<emphasis id="B9080A26362FFF678DD2FDB5F776FD7C" box="[2244,2255,624,659]" italics="true" pageId="11" pageNumber="177">(</emphasis>
|
||
G
|
||
<emphasis id="B9080A26362FFF678DFCFDB5F74CFD7C" box="[2282,2293,624,659]" italics="true" pageId="11" pageNumber="177">)</emphasis>
|
||
and extensor
|
||
<emphasis id="B9080A26362FFF678D4BFD61F75FFD28" box="[2141,2278,676,711]" italics="true" pageId="11" pageNumber="177">side (H).</emphasis>
|
||
Metacarpal
|
||
<emphasis id="B9080A26362FFF678D54FD1CF7D9FD13" box="[2114,2144,729,764]" italics="true" pageId="11" pageNumber="177">III</emphasis>
|
||
of
|
||
<emphasis id="B9080A26362FFF678D8CFD1CF859FCDE" italics="true" pageId="11" pageNumber="177">MOR 690 in</emphasis>
|
||
lateral
|
||
<emphasis id="B9080A26362FFF678D73FCCBF73AFCDE" box="[2149,2179,782,817]" italics="true" pageId="11" pageNumber="177">(I)</emphasis>
|
||
and extensor
|
||
<emphasis id="B9080A26362FFF6782FFFC86F7DAFC89" box="[2025,2147,835,870]" italics="true" pageId="11" pageNumber="177">side (J).</emphasis>
|
||
Scale
|
||
<emphasis id="B9080A26362FFF678DCCFC86F741FC89" box="[2266,2296,835,870]" italics="true" pageId="11" pageNumber="177">in</emphasis>
|
||
centimeters.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BC3D634362FFF61870AF64BFD17F925" blockId="11.[456,1820,2144,3013]" lastBlockId="13.[444,1809,1457,2975]" lastPageId="13" lastPageNumber="779" pageId="11" pageNumber="177">
|
||
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">
|
||
<emphasis id="B9080A263629FF618366F9ACFD8AF925" italics="true" pageId="13" pageNumber="779">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
below.
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343628FF608CAFFFCCF678FFC7" box="[2489,2497,9,40]" pageId="12" pageNumber="178">!</paragraph>
|
||
<paragraph id="8BC3D6343628FF608CACFFFBF678FFBF" box="[2490,2497,62,80]" pageId="12" pageNumber="178">s</paragraph>
|
||
<caption id="DF0386BC3628FF6084DFFF16FD21FDDC" ID-DOI="http://doi.org/10.5281/zenodo.3942835" ID-Zenodo-Dep="3942835" httpUri="https://zenodo.org/record/3942835/files/figure.png" pageId="12" pageNumber="178" targetBox="[976,2381,209,1786]" targetPageId="12">
|
||
<paragraph id="8BC3D6343628FF6084DFFF16FD21FDDC" blockId="12.[453,903,207,563]" pageId="12" pageNumber="178">
|
||
Figure 10.11. Distal carpal and metacarpals in
|
||
<emphasis id="B9080A263628FF608608FEC0FC8BFEB0" italics="true" pageId="12" pageNumber="178">
|
||
articulation (
|
||
<materialsCitation id="3B14DC693628FF608797FEFFFC98FEB0" ID-GBIF-Occurrence="2813095320" box="[641,801,314,351]" collectionCode="BHI" pageId="12" pageNumber="178" specimenCode="BHI 6230">BHI 6230</materialsCitation>
|
||
).
|
||
</emphasis>
|
||
Proximal
|
||
<emphasis id="B9080A263628FF60877FFEAAFE4EFE26" italics="true" pageId="12" pageNumber="178">(A), digit I side (B),</emphasis>
|
||
extensor
|
||
<emphasis id="B9080A263628FF6087BCFE61FCB1FE26" box="[682,776,420,457]" italics="true" pageId="12" pageNumber="178">side (</emphasis>
|
||
C
|
||
<emphasis id="B9080A263628FF608636FE61FC96FE26" box="[800,815,420,457]" italics="true" pageId="12" pageNumber="178">),</emphasis>
|
||
digit
|
||
<emphasis id="B9080A263628FF608736FE1CFD76FE11" box="[544,719,473,510]" italics="true" pageId="12" pageNumber="178">III side (D).</emphasis>
|
||
Scale in centimeters.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF0386BC3628FF6084A8F86CFDE5F770" ID-DOI="http://doi.org/10.5281/zenodo.3942837" ID-Zenodo-Dep="3942837" httpUri="https://zenodo.org/record/3942837/files/figure.png" pageId="12" pageNumber="178" startId="12.[446,551,1961,1996]" targetBox="[970,2327,1946,2984]" targetPageId="12">
|
||
<paragraph id="8BC3D6343628FF6084A8F86CFDE5F770" blockId="12.[442,885,1955,2207]" pageId="12" pageNumber="178">
|
||
Figure 10.12. Articulated pectoral
|
||
<emphasis id="B9080A263628FF60874EF818FD03F7EF" box="[600,698,2013,2048]" italics="true" pageId="12" pageNumber="178">girdle</emphasis>
|
||
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>
|
||
<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>
|
||
A
|
||
<emphasis id="B9080A263628FF608790F78DFD2BF784" box="[646,658,2120,2155]" italics="true" pageId="12" pageNumber="178">)</emphasis>
|
||
and anterior (B) views.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF0386BC3629FF628272FA72FD2CF6A8" ID-DOI="http://doi.org/10.5281/zenodo.3942839" ID-Zenodo-Dep="3942839" httpUri="https://zenodo.org/record/3942839/files/figure.png" lastPageId="14" lastPageNumber="180" pageId="13" pageNumber="779" startId="13.[1892,1995,1463,1498]" subCaptionStartIDs="14.[569,676,1651,1686] 14.[704,807,1757,1792]" subCaptionStarts="Figu & figu" targetBox="[456,2335,199,1371]" targetPageId="13">
|
||
<paragraph id="8BC3D6343629FF628272FA72FD2CF6A8" blockId="13.[1882,2333,1458,2981]" lastBlockId="14.[442,903,216,2376]" lastPageId="14" lastPageNumber="180" pageId="13" pageNumber="779">
|
||
Figure 10.13. Deformation of a crocodilian and
|
||
<emphasis id="B9080A263629FF618276F9E4F7FEF9AB" box="[1888,2119,1569,1604]" italics="true" pageId="13" pageNumber="779">avian scapula</emphasis>
|
||
and
|
||
<emphasis id="B9080A263629FF618D88F9E4F878F996" italics="true" pageId="13" pageNumber="779">coracoid (</emphasis>
|
||
SC
|
||
<emphasis id="B9080A263629FF6182F9F993F843F996" box="[2031,2042,1622,1657]" italics="true" pageId="13" pageNumber="779">)</emphasis>
|
||
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>
|
||
. Wavy vertical
|
||
<emphasis id="B9080A263629FF618D45F905F718F90C" box="[2131,2209,1728,1763]" italics="true" pageId="13" pageNumber="779">lines</emphasis>
|
||
show direction and degree of
|
||
<emphasis id="B9080A263629FF618276F8EFF7ABF8A2" box="[1888,2066,1834,1869]" italics="true" pageId="13" pageNumber="779">morphing.</emphasis>
|
||
This method allows for the
|
||
<emphasis id="B9080A263629FF618D4EF89AF6B5F86D" box="[2136,2316,1887,1922]" italics="true" pageId="13" pageNumber="779">prediction</emphasis>
|
||
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>
|
||
<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>
|
||
<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>
|
||
the end point to
|
||
<emphasis id="B9080A263629FF618D93F758F82DF71A" italics="true" pageId="13" pageNumber="779">morphing</emphasis>
|
||
of the
|
||
<emphasis id="B9080A263629FF618D03F717F786F71A" box="[2069,2111,2258,2293]" italics="true" pageId="13" pageNumber="779">(B)</emphasis>
|
||
crocodilian and
|
||
<emphasis id="B9080A263629FF6182BDF6C2F780F6C5" box="[1963,2105,2311,2346]" italics="true" pageId="13" pageNumber="779">(E) avian</emphasis>
|
||
pectoral
|
||
<emphasis id="B9080A263629FF61824AF6F9F783F6B0" box="[1884,2106,2364,2399]" italics="true" pageId="13" pageNumber="779">girdles. Note</emphasis>
|
||
that
|
||
<emphasis id="B9080A263629FF618D80F6F9F812F67B" italics="true" pageId="13" pageNumber="779">2 possible</emphasis>
|
||
scenarios
|
||
<emphasis id="B9080A263629FF618D74F6B4F7D5F67B" box="[2146,2156,2417,2452]" italics="true" pageId="13" pageNumber="779">(</emphasis>
|
||
C,
|
||
<emphasis id="B9080A263629FF618D8AF6B4F805F626" italics="true" pageId="13" pageNumber="779">D) occur in</emphasis>
|
||
the
|
||
<emphasis id="B9080A263629FF618D19F663F700F626" box="[2063,2233,2470,2505]" italics="true" pageId="13" pageNumber="779">morphing</emphasis>
|
||
of the crocodilian SC
|
||
<emphasis id="B9080A263629FF618DB4F61FF855F5DD" italics="true" pageId="13" pageNumber="779">depending</emphasis>
|
||
on what portions of the
|
||
<emphasis id="B9080A263629FF618D26F581F7BBF573" italics="true" pageId="13" pageNumber="779">crocodilian scapula is</emphasis>
|
||
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>
|
||
scapula. Location of
|
||
<emphasis id="B9080A263629FF618DD4F4D0F824F49F" italics="true" pageId="13" pageNumber="779">actual</emphasis>
|
||
muscle scars
|
||
<emphasis id="B9080A263629FF618D9BF48EF721F49F" box="[2189,2200,2891,2928]" italics="true" pageId="13" pageNumber="779">(</emphasis>
|
||
G
|
||
<emphasis id="B9080A263629FF618DA2F48EF77DF49F" box="[2228,2244,2891,2928]" italics="true" pageId="13" pageNumber="779">).</emphasis>
|
||
Abbreviations: bb, M.
|
||
<emphasis id="B9080A263629FF628DBBF445FCAFFF10" italics="true" lastPageId="14" lastPageNumber="180" pageId="13" pageNumber="779">biceps (continued) brachii;</emphasis>
|
||
c-M. costocoracoideus; cbd, M. coracobrachialis
|
||
<emphasis id="B9080A26362AFF628632FE86FDEAFE72" italics="true" pageId="14" pageNumber="180">brevis dorsalis;</emphasis>
|
||
cbv, M.
|
||
<emphasis id="B9080A26362AFF6287F8FEBDFD60FE3D" italics="true" pageId="14" pageNumber="180">coracobrachialis brevis</emphasis>
|
||
ventralis;
|
||
<emphasis id="B9080A26362AFF6284D3FE27FE48FDE8" box="[453,497,482,519]" italics="true" pageId="14" pageNumber="180">ce,</emphasis>
|
||
M. coracobrachialis externus; ch, M.
|
||
<emphasis id="B9080A26362AFF6287F9FDD2FDD6FD9C" italics="true" pageId="14" pageNumber="180">coracohumeralis</emphasis>
|
||
; dc, M. deltoideus clavicularis;
|
||
<emphasis id="B9080A26362AFF6287E4FD47FCA6FD48" box="[754,799,642,679]" italics="true" pageId="14" pageNumber="180">ds,</emphasis>
|
||
M. deltoideus scapularis;
|
||
<emphasis id="B9080A26362AFF628652FD73FCEBFD34" box="[836,850,694,731]" italics="true" pageId="14" pageNumber="180">I,</emphasis>
|
||
M. levator scapulae;
|
||
<emphasis id="B9080A26362AFF628638FD28FCEAFCFF" box="[814,851,749,784]" italics="true" pageId="14" pageNumber="180">rs,</emphasis>
|
||
M. rhomboideus
|
||
<emphasis id="B9080A26362AFF6287E4FCE6FDA6FC95" italics="true" pageId="14" pageNumber="180">superficialis;</emphasis>
|
||
sb, M. supracoracoideus
|
||
<emphasis id="B9080A26362AFF628745FC48FD7BFC5F" box="[595,706,909,944]" italics="true" pageId="14" pageNumber="180">brevis;</emphasis>
|
||
sc,
|
||
<emphasis id="B9080A26362AFF62861EFC48FC8AFC5F" box="[776,819,909,944]" italics="true" pageId="14" pageNumber="180">M.</emphasis>
|
||
scapulohumeralis
|
||
<emphasis id="B9080A26362AFF6287EBFC07FE5DFBF5" italics="true" pageId="14" pageNumber="180">cranialis</emphasis>
|
||
; scd, scapulohumeralis caudalis;
|
||
<emphasis id="B9080A26362AFF628775FBEEFD29FBA1" box="[611,656,1067,1102]" italics="true" pageId="14" pageNumber="180">se,</emphasis>
|
||
M.
|
||
<emphasis id="B9080A26362AFF6287CCFBEEFDA4FB6B" italics="true" pageId="14" pageNumber="180">subscapularis</emphasis>
|
||
externus;
|
||
<emphasis id="B9080A26362AFF6287C2FBA4FD4CFB6B" box="[724,757,1121,1156]" italics="true" pageId="14" pageNumber="180">si,</emphasis>
|
||
M.
|
||
<emphasis id="B9080A26362AFF6284A9FB53FD2BFB02" italics="true" pageId="14" pageNumber="180">supracoracoideus intermedius; svc,</emphasis>
|
||
M.
|
||
<emphasis id="B9080A26362AFF6287CAFB0FFDA5FACC" italics="true" pageId="14" pageNumber="180">subscapularis</emphasis>
|
||
ventralis cranialis;
|
||
<emphasis id="B9080A26362AFF6284A9FAF0FE40FAB7" box="[447,505,1333,1368]" italics="true" pageId="14" pageNumber="180">svt,</emphasis>
|
||
M. serratus ventralis thoracis;
|
||
<emphasis id="B9080A26362AFF628774FAAFFD96FA2E" italics="true" pageId="14" pageNumber="180">t, M. trapezius; tbclps,</emphasis>
|
||
M.
|
||
<emphasis id="B9080A26362AFF62876BFA5BFCCFFA2E" box="[637,886,1438,1473]" italics="true" pageId="14" pageNumber="180">triceps brachii;</emphasis>
|
||
caput
|
||
<emphasis id="B9080A26362AFF628738FA11FD1AFA18" box="[558,675,1492,1527]" italics="true" pageId="14" pageNumber="180">longus</emphasis>
|
||
pars
|
||
<emphasis id="B9080A26362AFF628617FA11FDF8F9C4" italics="true" pageId="14" pageNumber="180">scapularis; tll,</emphasis>
|
||
M.
|
||
<emphasis id="B9080A26362AFF628786F9CDFCB8F9C4" box="[656,769,1544,1579]" italics="true" pageId="14" pageNumber="180">triceps</emphasis>
|
||
longus
|
||
<emphasis id="B9080A26362AFF6284A9F9FBFDF6F98E" box="[447,591,1598,1633]" italics="true" pageId="14" pageNumber="180">lateralis;</emphasis>
|
||
tm,
|
||
<emphasis id="B9080A26362AFF6287B4F9FBFD67F979" italics="true" pageId="14" pageNumber="180">M. terres major. Figure (B)</emphasis>
|
||
and
|
||
<emphasis id="B9080A26362AFF62862EF9B6FDDDF924" italics="true" pageId="14" pageNumber="180">terminology</emphasis>
|
||
adapted from
|
||
<emphasis id="B9080A26362AFF6284ABF918FD08F8EF" box="[445,689,1757,1792]" italics="true" pageId="14" pageNumber="180">Meers (2003);</emphasis>
|
||
figure
|
||
<emphasis id="B9080A26362AFF628625F918FCE2F8EF" box="[819,859,1757,1792]" italics="true" pageId="14" pageNumber="180">(E)</emphasis>
|
||
and terminology adapted from
|
||
<emphasis id="B9080A26362AFF628734F882FD84F870" italics="true" pageId="14" pageNumber="180">
|
||
<bibRefCitation id="EFEDABC5362AFF628734F882FCA3F885" author="Yasuda, M." box="[546,794,1863,1898]" firstAuthor="Yasuda" journalOrPublisher="Tokyo, University of Tokyo Press" pageId="14" pageNumber="180" refId="ref10803" refString="Yasuda, M. 2002. The Anatomy of Gallus. Tokyo, University of Tokyo Press." title="The Anatomy of Gallus" type="book" year="2002">Yasuda (2002)</bibRefCitation>
|
||
. Although
|
||
</emphasis>
|
||
the
|
||
<emphasis id="B9080A26362AFF628787F8B9FE6FF83B" italics="true" pageId="14" pageNumber="180">terminology is</emphasis>
|
||
retained for
|
||
<emphasis id="B9080A26362AFF6287D6F874FD74F83B" box="[704,717,1969,2004]" italics="true" pageId="14" pageNumber="180">“</emphasis>
|
||
dorsal
|
||
<emphasis id="B9080A26362AFF62862AF874FDFEF7E6" italics="true" pageId="14" pageNumber="180">" versus "</emphasis>
|
||
ventral
|
||
<emphasis id="B9080A26362AFF6287D3F823FD6AF7E6" box="[709,723,2022,2057]" italics="true" pageId="14" pageNumber="180">"</emphasis>
|
||
muscles
|
||
<emphasis id="B9080A26362AFF6284A8F7DEFE74F7D1" box="[446,461,2075,2110]" italics="true" pageId="14" pageNumber="180">(</emphasis>
|
||
<emphasis id="B9080A26362AFF6284DBF7DEFE58F7D1" bold="true" box="[461,481,2075,2110]" pageId="14" pageNumber="180">e</emphasis>
|
||
.g., m.c.b. ventralis
|
||
<emphasis id="B9080A26362AFF628633F7DEFC8EF7D1" box="[805,823,2075,2110]" italics="true" pageId="14" pageNumber="180">),</emphasis>
|
||
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">
|
||
<emphasis id="B9080A26362AFF6287C2F740FD80F731" italics="true" pageId="14" pageNumber="180">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
indicates a need for modified terminology.
|
||
</paragraph>
|
||
</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>
|
||
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>
|
||
,
|
||
</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>
|
||
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>
|
||
(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>
|
||
<emphasis id="B9080A263629FF6181EEF4B0FABDF474" box="[1272,1284,2933,2971]" italics="true" pageId="13" pageNumber="779">)</emphasis>
|
||
and bird
|
||
<emphasis id="B9080A263629FF6180A7F4B0F986F474" box="[1457,1599,2933,2971]" italics="true" pageId="13" pageNumber="779">
|
||
(
|
||
<taxonomicName id="4C7CADB73629FF6180A8F4B0F98BF474" box="[1470,1586,2933,2971]" class="Aves" family="Phasianidae" genus="Gallus" kingdom="Animalia" order="Galliformes" pageId="13" pageNumber="779" phylum="Chordata" rank="genus">Gallus</taxonomicName>
|
||
)
|
||
</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>
|
||
(
|
||
<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">
|
||
<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">
|
||
<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>
|
||
. 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>
|
||
.
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph id="8BC3D634362AFF62813DFCEDF9CBF809" blockId="14.[976,2348,215,3024]" pageId="14" pageNumber="180">
|
||
The technique begins by scanning scapulocoracoid outlines on which the muscles have been mapped (
|
||
<figureCitation id="1347CAB1362AFF628335FCA6F96EFC66" box="[1571,1751,867,905]" 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>
|
||
B, E). Minimum thickness of the scapular neck was selected to standardize the scapulocoracoids of the
|
||
<emphasis id="B9080A26362AFF628DBAFC5BFB25FC11" italics="true" pageId="14" pageNumber="180">
|
||
<taxonomicName id="4C7CADB7362AFF628DBAFC5BFBBEFC11" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Alligator</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C7CADB7362AFF628108FC1DFB2CFC11" box="[1054,1173,984,1022]" class="Aves" family="Phasianidae" genus="Gallus" kingdom="Animalia" order="Galliformes" pageId="14" pageNumber="180" phylum="Chordata" rank="genus">Gallus</taxonomicName>
|
||
,
|
||
</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">
|
||
<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>
|
||
(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
|
||
<figureCitation id="1347CAB1362AFF6282F6FA6AF7F8FA3A" box="[2016,2113,1455,1493]" 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">10.13</figureCitation>
|
||
C, D, F). Because moving the nodes also moved the contents of each grid, the result
|
||
<emphasis id="B9080A26362AFF628DFDFA2CF6BCF9E0" box="[2283,2309,1513,1551]" italics="true" pageId="14" pageNumber="180">is</emphasis>
|
||
a prediction ofwhat the resultant muscle pattern would be like. As used, Mesh Warp is not mathematically as rigorous as the thin plate spline of
|
||
<bibRefCitation id="EFEDABC5362AFF628D78F99AFBF9F92F" author="Bookstein, F. L." firstAuthor="Bookstein" journalOrPublisher="Cambridge University Press, Cambridge" pageId="14" pageNumber="180" refId="ref8848" refString="Bookstein, F. L. 1991. Morphometric tools for landmark data. Geometry and Biology. Cambridge University Press, Cambridge," title="Morphometric tools for landmark data. Geometry and Biology" type="book" year="1991">Bookstein (1991)</bibRefCitation>
|
||
because measuring the change in landmark position
|
||
<emphasis id="B9080A26362AFF6282E5F95FF7B4F92F" box="[2035,2061,1690,1728]" italics="true" pageId="14" pageNumber="180">is</emphasis>
|
||
irrelevant. Two versions are presented for the deformed crocodile scapulocoracoid, with results differing in the acromion.
|
||
<figureCitation id="1347CAB1362AFF628330F8D5F8A9F8D9" box="[1574,1808,1808,1846]" 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 assumes the dorsal prominence just anterior to the scapular neck of the crocodile
|
||
<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>
|
||
,
|
||
</emphasis>
|
||
whereas
|
||
<figureCitation id="1347CAB1362AFF628DBEF840FB8CF809" 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>
|
||
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">
|
||
<emphasis id="B9080A26362AFF628175F7B5FADFF779" box="[1123,1382,2160,2198]" italics="true" pageId="14" pageNumber="180">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<materialsCitation id="3B14DC69362AFF628090F7B5F93FF779" ID-GBIF-Occurrence="2813095325" box="[1414,1670,2160,2198]" collectionCode="DMNH" pageId="14" pageNumber="180" specimenCode="DMNH 2827">DMNH 2827</materialsCitation>
|
||
). As can be seen in
|
||
<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">
|
||
<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">
|
||
<emphasis id="B9080A26362AFF628DB8F5CEFBCBF583" italics="true" pageId="14" pageNumber="180">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
seems to better match the pattern for the M. supracoracoideus intermedius in
|
||
<figureCitation id="1347CAB1362AFF6281FEF544FA77F548" box="[1256,1486,2689,2727]" 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>
|
||
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>
|
||
.
|
||
</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">
|
||
<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 (
|
||
<emphasis id="B9080A26362BFF638307FA6DF91AFA21" box="[1553,1699,1448,1486]" italics="true" pageId="15" pageNumber="181">DMNH</emphasis>
|
||
avian collection,
|
||
<emphasis id="B9080A26362BFF638798FA26FBE5F9E6" box="[654,1116,1507,1545]" italics="true" pageId="15" pageNumber="181">including Aechmophorus,</emphasis>
|
||
Buteo, Cygnus,
|
||
<taxonomicName id="4C7CADB7362BFF63809EFA26FD87F9AB" authority=", Aquila, Gymnogyps" authorityName="Aquila, Gymnogyps" class="Aves" family="Corvidae" genus="Corvus" kingdom="Animalia" order="Passeriformes" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">
|
||
<emphasis id="B9080A26362BFF63809EFA26FD87F9AB" italics="true" pageId="15" pageNumber="181">Corvus, Aquila, Gymnogyps</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C7CADB7362BFF638740F9DBFD68F9AB" box="[598,721,1566,1604]" class="Aves" family="Phasianidae" genus="Gallus" kingdom="Animalia" order="Galliformes" pageId="15" pageNumber="181" phylum="Chordata" rank="genus">Gallus</taxonomicName>
|
||
,
|
||
<emphasis id="B9080A26362BFF6387FFF9DBFC5BF9AB" box="[745,994,1566,1604]" italics="true" pageId="15" pageNumber="181">and Struthio),</emphasis>
|
||
a faint
|
||
<emphasis id="B9080A26362BFF638162F9DBFAE9F9AB" box="[1140,1360,1566,1604]" italics="true" pageId="15" pageNumber="181">longitudinal</emphasis>
|
||
trough is
|
||
<emphasis id="B9080A26362BFF638307F9DBF92CF9AB" box="[1553,1685,1566,1604]" italics="true" pageId="15" pageNumber="181">present</emphasis>
|
||
in
|
||
<emphasis id="B9080A26362BFF6383C3F9DBF8B7F9AB" box="[1749,1806,1566,1604]" italics="true" pageId="15" pageNumber="181">the</emphasis>
|
||
distal half of
|
||
<emphasis id="B9080A26362BFF6387B1F99CFD5BF990" box="[679,738,1625,1663]" italics="true" pageId="15" pageNumber="181">the</emphasis>
|
||
scapular blade. Interpreted
|
||
<emphasis id="B9080A26362BFF6381C0F99CFAD8F990" box="[1238,1377,1625,1663]" italics="true" pageId="15" pageNumber="181">another</emphasis>
|
||
way, there is thickening ofthe
|
||
<emphasis id="B9080A26362BFF638739F951FD7FF955" box="[559,710,1684,1722]" italics="true" pageId="15" pageNumber="181">scapular</emphasis>
|
||
surface along the origins of the M. terres major and M. rhomboideus
|
||
<emphasis id="B9080A26362BFF638740F90AFC8AF91A" box="[598,819,1743,1781]" italics="true" pageId="15" pageNumber="181">superficialis</emphasis>
|
||
; the trough is the unthickened bone between these origins.
|
||
<emphasis id="B9080A26362BFF638736F8CFFDFEF8DF" box="[544,583,1802,1840]" italics="true" pageId="15" pageNumber="181">In</emphasis>
|
||
contrast, the crocodile has a single longitudinal
|
||
<emphasis id="B9080A26362BFF6380A0F8CFF9C1F8DF" box="[1462,1656,1802,1840]" italics="true" pageId="15" pageNumber="181">thickening</emphasis>
|
||
or
|
||
<emphasis id="B9080A26362BFF6383A4F8CFF8B4F8DF" box="[1714,1805,1802,1840]" italics="true" pageId="15" pageNumber="181">ridge</emphasis>
|
||
on the scapular surface that corresponds roughly to the
|
||
<emphasis id="B9080A26362BFF6380B9F881F9E8F885" box="[1455,1617,1860,1898]" italics="true" pageId="15" pageNumber="181">common</emphasis>
|
||
margin of the
|
||
<emphasis id="B9080A26362BFF638710F8BAFD8DF84A" box="[518,564,1919,1957]" italics="true" pageId="15" pageNumber="181">M</emphasis>
|
||
.
|
||
<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>
|
||
also has
|
||
</emphasis>
|
||
<emphasis id="B9080A26362BFF6387CFF87FFD55F80F" bold="true" box="[729,748,1978,2016]" pageId="15" pageNumber="181">a</emphasis>
|
||
<emphasis id="B9080A26362BFF6387E1F87FFCCEF80F" box="[759,887,1978,2016]" italics="true" pageId="15" pageNumber="181">similar</emphasis>
|
||
ridge, not the
|
||
<emphasis id="B9080A26362BFF63816BF87FFB4DF80F" box="[1149,1268,1978,2016]" italics="true" pageId="15" pageNumber="181">trough</emphasis>
|
||
seen in birds.
|
||
</paragraph>
|
||
<caption id="DF0386BC362BFF63827CFF0DF7F4F7B3" ID-DOI="http://doi.org/10.5281/zenodo.3942841" ID-Zenodo-Dep="3942841" httpUri="https://zenodo.org/record/3942841/files/figure.png" pageId="15" pageNumber="181" startId="15.[1898,2001,200,235]" subCaptionStartIDs="15.[1991,2098,462,497]" subCaptionStarts="Figure 10" targetBox="[460,1809,196,1277]" targetPageId="15">
|
||
<paragraph id="8BC3D634362BFF63827CFF0DF7F4F7B3" blockId="15.[1888,2342,193,2140]" pageId="15" pageNumber="181">
|
||
Figure 10.14.
|
||
<emphasis id="B9080A26362BFF638D42FF0DF74EFF04" box="[2132,2295,200,235]" italics="true" pageId="15" pageNumber="181">Resultant</emphasis>
|
||
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>
|
||
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
|
||
<emphasis id="B9080A26362BFF638DCCFDC6F810FDB5" italics="true" pageId="15" pageNumber="181">actual</emphasis>
|
||
and predicted
|
||
<emphasis id="B9080A26362BFF638DA5FDF2F7F3FD60" italics="true" pageId="15" pageNumber="181">include relative</emphasis>
|
||
<emphasis id="B9080A26362BFF638D42FDA9F71EFD60" bold="true" box="[2132,2215,620,655]" pageId="15" pageNumber="181">sizes</emphasis>
|
||
of scars
|
||
<emphasis id="B9080A26362BFF6382DEFD64F84BFD2B" box="[1992,2034,673,708]" italics="true" pageId="15" pageNumber="181">(e.</emphasis>
|
||
g., m. deltoideus clavicularis
|
||
<emphasis id="B9080A26362BFF638D0AFD13F794FD16" box="[2076,2093,726,761]" italics="true" pageId="15" pageNumber="181">),</emphasis>
|
||
as well
|
||
<emphasis id="B9080A26362BFF638DD6FD13F75CFD16" bold="true" box="[2240,2277,726,761]" pageId="15" pageNumber="181">as</emphasis>
|
||
<emphasis id="B9080A26362BFF638272FCCEF7B5FCC1" box="[1892,2060,779,814]" italics="true" pageId="15" pageNumber="181">position (</emphasis>
|
||
<emphasis id="B9080A26362BFF638D1AFCCEF799FCC1" bold="true" box="[2060,2080,779,814]" pageId="15" pageNumber="181">e</emphasis>
|
||
.g., M. terres
|
||
<emphasis id="B9080A26362BFF638270FC85F855FC8C" box="[1894,2028,832,867]" italics="true" pageId="15" pageNumber="181">major +</emphasis>
|
||
M. latissimus dorsi
|
||
<emphasis id="B9080A26362BFF6382ADFCB3F875FC76" box="[1979,1996,886,921]" italics="true" pageId="15" pageNumber="181">).</emphasis>
|
||
Where muscle
|
||
<emphasis id="B9080A26362BFF638275FC6EF842FC21" box="[1891,2043,939,974]" italics="true" pageId="15" pageNumber="181">
|
||
scars
|
||
<emphasis id="B9080A26362BFF6382D2FC6EF842FC21" bold="true" box="[1988,2043,939,974]" italics="true" pageId="15" pageNumber="181">are</emphasis>
|
||
</emphasis>
|
||
ambiguous, the
|
||
<emphasis id="B9080A26362BFF638272FC1AF7ACFBED" box="[1892,2069,991,1026]" italics="true" pageId="15" pageNumber="181">prediction</emphasis>
|
||
was used
|
||
<emphasis id="B9080A26362BFF638DC6FC1AF74FFBED" box="[2256,2294,991,1026]" italics="true" pageId="15" pageNumber="181">as</emphasis>
|
||
a guide constrained
|
||
<emphasis id="B9080A26362BFF638DBEFBD1F76CFBD8" box="[2216,2261,1044,1079]" italics="true" pageId="15" pageNumber="181">by</emphasis>
|
||
unambiguous scars
|
||
<emphasis id="B9080A26362BFF638D85FB8CF71BFB83" box="[2195,2210,1097,1132]" italics="true" pageId="15" pageNumber="181">(</emphasis>
|
||
<emphasis id="B9080A26362BFF638DB4FB8CF70EFB83" bold="true" box="[2210,2231,1097,1132]" pageId="15" pageNumber="181">e</emphasis>
|
||
.g., M.
|
||
<emphasis id="B9080A26362BFF638273FBBBF800FB38" italics="true" pageId="15" pageNumber="181">triceps brevis). Abbreviations</emphasis>
|
||
: b, M. brachialis; cb, M. coracobrachialis
|
||
<emphasis id="B9080A26362BFF638DD6FB2DF821FAAE" italics="true" pageId="15" pageNumber="181">brevis;</emphasis>
|
||
cbd, M. coracobrachialis
|
||
<emphasis id="B9080A26362BFF6382C0FA96F76CFA99" box="[2006,2261,1363,1398]" italics="true" pageId="15" pageNumber="181">brevis dorsalis;</emphasis>
|
||
dc, M. deltoideus clavicularis; hr, M. humeroradialis;
|
||
<emphasis id="B9080A26362BFF638DF9FA78F6B4FA0F" box="[2287,2317,1469,1504]" italics="true" pageId="15" pageNumber="181">p,</emphasis>
|
||
M. pectoralis; sb, M.
|
||
<emphasis id="B9080A26362BFF638DC3FA34F7D2F9A5" italics="true" pageId="15" pageNumber="181">supracoracoideus</emphasis>
|
||
<emphasis id="B9080A26362BFF638D63F9E2F765F9A5" bold="true" box="[2165,2268,1575,1610]" pageId="15" pageNumber="181">brevis</emphasis>
|
||
;
|
||
<emphasis id="B9080A26362BFF638DE7F9E2F6A3F9A5" box="[2289,2330,1575,1610]" italics="true" pageId="15" pageNumber="181">sc,</emphasis>
|
||
M. scapulohumeralis
|
||
<emphasis id="B9080A26362BFF638DCEF998F83AF906" italics="true" pageId="15" pageNumber="181">cranialis; sc, scapulohumeralis</emphasis>
|
||
caudalis;
|
||
<emphasis id="B9080A26362BFF638D38F903F788F8F0" italics="true" pageId="15" pageNumber="181">scc, supracoracoideus</emphasis>
|
||
complex; sl, M.
|
||
<emphasis id="B9080A26362BFF63828CF8F4F774F8BB" box="[1946,2253,1841,1876]" italics="true" pageId="15" pageNumber="181">supracoracoideus</emphasis>
|
||
longus; tbi, M.
|
||
<emphasis id="B9080A26362BFF638D20F8A3F71EF866" box="[2102,2215,1894,1929]" italics="true" pageId="15" pageNumber="181">triceps</emphasis>
|
||
brevis intermedius
|
||
<emphasis id="B9080A26362BFF638D2CF85FF7E3F852" box="[2106,2138,1946,1981]" italics="true" pageId="15" pageNumber="181">(+</emphasis>
|
||
cranialis
|
||
<emphasis id="B9080A26362BFF638DE2F85FF6A1F852" box="[2292,2328,1946,1981]" italics="true" pageId="15" pageNumber="181">?);</emphasis>
|
||
tm, M. terres major.
|
||
<emphasis id="B9080A26362BFF638DDBF80AF7BDF7C8" italics="true" pageId="15" pageNumber="181">Terminology</emphasis>
|
||
adapted from Meers (2003).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BC3D634362BFF638706F830FA20F463" blockId="15.[443,1810,1351,2956]" pageId="15" pageNumber="181">
|
||
The deformation method outlined above
|
||
<emphasis id="B9080A26362BFF638004F830FAEDF7F4" box="[1298,1364,2037,2075]" italics="true" pageId="15" pageNumber="181">was</emphasis>
|
||
applied
|
||
<emphasis id="B9080A26362BFF6380E2F830F9ACF7F4" box="[1524,1557,2037,2075]" italics="true" pageId="15" pageNumber="181">to</emphasis>
|
||
the humerus (
|
||
<figureCitation id="1347CAB1362BFF6384DBF7EAFDC2F7BA" box="[461,635,2095,2133]" captionStart="Fig. 10.9" captionText="Figure 10.9. Muscle maps for humerus in Tyrannosaurus, Alligator, and Gallus. Top row is anterior, bottom row is posterior. Muscle map based on scars (A, F) Tyrannosaurus. Map for Alligator (B, G) and predicted for Tyrannosaurus (C, D) based on deformation of Alligator humerus. Map for Gallus (D, I) and predicted for Tyrannosaurus (E, J) based on deformation of Gallus humerus. Note that predicted scars for deformed Alligator (C, H) are a better match for the scars of Tyrannosaurus (A, F). This prediction is also supported by the pattern ofavulsion seen in a Tyrannosaurus humerus (K, L). See Figure 10.13 and text for further explanation. " figureDoi="http://doi.org/10.5281/zenodo.3942831" httpUri="https://zenodo.org/record/3942831/files/figure.png" pageId="15" pageNumber="181" targetBox="[480,1829,206,1482]" targetPageId="9">Fig. 10.9</figureCitation>
|
||
B-E,
|
||
<emphasis id="B9080A26362BFF6387F4F7EAFC96F7BA" box="[738,815,2095,2133]" italics="true" pageId="15" pageNumber="181">G-J</emphasis>
|
||
) and tested against the muscle scars (
|
||
<figureCitation id="1347CAB1362BFF638314F7EAF91BF7BA" box="[1538,1698,2095,2133]" captionStart="Fig. 10.9" captionText="Figure 10.9. Muscle maps for humerus in Tyrannosaurus, Alligator, and Gallus. Top row is anterior, bottom row is posterior. Muscle map based on scars (A, F) Tyrannosaurus. Map for Alligator (B, G) and predicted for Tyrannosaurus (C, D) based on deformation of Alligator humerus. Map for Gallus (D, I) and predicted for Tyrannosaurus (E, J) based on deformation of Gallus humerus. Note that predicted scars for deformed Alligator (C, H) are a better match for the scars of Tyrannosaurus (A, F). This prediction is also supported by the pattern ofavulsion seen in a Tyrannosaurus humerus (K, L). See Figure 10.13 and text for further explanation. " figureDoi="http://doi.org/10.5281/zenodo.3942831" httpUri="https://zenodo.org/record/3942831/files/figure.png" pageId="15" pageNumber="181" targetBox="[480,1829,206,1482]" targetPageId="9">Fig. 10.9</figureCitation>
|
||
A, F). Overall, the pattern most closely resembles
|
||
<emphasis id="B9080A26362BFF6381F1F7AFFA96F77F" box="[1255,1327,2154,2192]" italics="true" pageId="15" pageNumber="181">that</emphasis>
|
||
of the crocodile,
|
||
<emphasis id="B9080A26362BFF638367F7AFF97AF77F" box="[1649,1731,2154,2192]" italics="true" pageId="15" pageNumber="181">with</emphasis>
|
||
notable exceptions. Aside from differences in relative
|
||
<emphasis id="B9080A26362BFF638085F760F9DEF724" box="[1427,1639,2213,2251]" italics="true" pageId="15" pageNumber="181">proportions</emphasis>
|
||
of some muscles between the actual and the predicted (cf.
|
||
<figureCitation id="1347CAB1362BFF63804CF725FA4DF6E9" box="[1370,1524,2272,2310]" captionStart="Fig. 10.9" captionText="Figure 10.9. Muscle maps for humerus in Tyrannosaurus, Alligator, and Gallus. Top row is anterior, bottom row is posterior. Muscle map based on scars (A, F) Tyrannosaurus. Map for Alligator (B, G) and predicted for Tyrannosaurus (C, D) based on deformation of Alligator humerus. Map for Gallus (D, I) and predicted for Tyrannosaurus (E, J) based on deformation of Gallus humerus. Note that predicted scars for deformed Alligator (C, H) are a better match for the scars of Tyrannosaurus (A, F). This prediction is also supported by the pattern ofavulsion seen in a Tyrannosaurus humerus (K, L). See Figure 10.13 and text for further explanation. " figureDoi="http://doi.org/10.5281/zenodo.3942831" httpUri="https://zenodo.org/record/3942831/files/figure.png" pageId="15" pageNumber="181" targetBox="[480,1829,206,1482]" targetPageId="9">
|
||
<emphasis id="B9080A26362BFF63804CF725FA20F6E9" box="[1370,1433,2272,2310]" italics="true" pageId="15" pageNumber="181">Fig</emphasis>
|
||
. 10.9
|
||
</figureCitation>
|
||
A and C, F and H), there are also some positional differences. For example, the common
|
||
<emphasis id="B9080A26362BFF6384A8F690FDDAF694" box="[446,611,2389,2427]" italics="true" pageId="15" pageNumber="181">insertion</emphasis>
|
||
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">
|
||
<emphasis id="B9080A26362BFF638166F655FAC0F659" box="[1136,1401,2448,2486]" italics="true" pageId="15" pageNumber="181">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
(cf.
|
||
<figureCitation id="1347CAB1362BFF6380DEF655F9DCF659" box="[1480,1637,2448,2486]" captionStart="Fig. 10.9" captionText="Figure 10.9. Muscle maps for humerus in Tyrannosaurus, Alligator, and Gallus. Top row is anterior, bottom row is posterior. Muscle map based on scars (A, F) Tyrannosaurus. Map for Alligator (B, G) and predicted for Tyrannosaurus (C, D) based on deformation of Alligator humerus. Map for Gallus (D, I) and predicted for Tyrannosaurus (E, J) based on deformation of Gallus humerus. Note that predicted scars for deformed Alligator (C, H) are a better match for the scars of Tyrannosaurus (A, F). This prediction is also supported by the pattern ofavulsion seen in a Tyrannosaurus humerus (K, L). See Figure 10.13 and text for further explanation. " figureDoi="http://doi.org/10.5281/zenodo.3942831" httpUri="https://zenodo.org/record/3942831/files/figure.png" pageId="15" pageNumber="181" targetBox="[480,1829,206,1482]" targetPageId="9">Fig. 10.9</figureCitation>
|
||
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">
|
||
<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
|
||
<emphasis id="B9080A26362BFF6384ABF585FDDBF589" box="[445,610,2624,2662]" italics="true" pageId="15" pageNumber="181">insertion</emphasis>
|
||
slightly below the peak of the deltopectoral crest (
|
||
<figureCitation id="1347CAB1362BFF6380E0F585F910F589" box="[1526,1705,2624,2662]" captionStart="Fig. 10.10" captionText="Figure 10.10. Distal carpal (BHI6230) of Tyrannosaurus in multiple views: proximal or dorsal (A); distal or ventral (B); anterior (C); posterior (D); extensor side (E); palmar side (F). Metacarpal III of BHI 6230 in lateral (G) and extensor side (H). Metacarpal III of MOR 690 in lateral (I) and extensor side (J). Scale in centimeters. " figureDoi="http://doi.org/10.5281/zenodo.3942833" httpUri="https://zenodo.org/record/3942833/files/figure.png" pageId="15" pageNumber="181" targetBox="[457,1819,194,2046]" targetPageId="11">Fig. 10.10</figureCitation>
|
||
)—either that or the pectoralis inserted more lateral
|
||
<emphasis id="B9080A26362BFF63802DF5BEFAE4F54E" box="[1339,1373,2683,2721]" italics="true" pageId="15" pageNumber="181">to</emphasis>
|
||
the deltopectoral crest than
|
||
<emphasis id="B9080A26362BFF63870AF573FD19F533" box="[540,672,2742,2780]" italics="true" pageId="15" pageNumber="181">medial</emphasis>
|
||
, but that seems highly unlikely. As noted above, the pathology on
|
||
<materialsCitation id="3B14DC69362BFF6384EEF535FD0AF4F9" ID-GBIF-Occurrence="2813095316" box="[504,691,2800,2838]" collectionCode="MOR" pageId="15" pageNumber="181" specimenCode="MOR 980">MOR 980</materialsCitation>
|
||
<emphasis id="B9080A26362BFF6387D6F535FC97F4F9" box="[704,814,2800,2838]" italics="true" pageId="15" pageNumber="181">better</emphasis>
|
||
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>
|
||
(
|
||
<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>
|
||
).
|
||
</paragraph>
|
||
<caption id="DF0386BC3634FF7C84DBFF2BFD08F76A" ID-DOI="http://doi.org/10.5281/zenodo.3942843" ID-Zenodo-Dep="3942843" httpUri="https://zenodo.org/record/3942843/files/figure.png" pageId="16" pageNumber="182" startId="16.[461,565,238,271]" subCaptionStartIDs="16.[688,810,448,483]" subCaptionStarts="Figures 10" targetBox="[993,2179,246,3040]" targetPageId="16">
|
||
<paragraph id="8BC3D6343634FF7C84DBFF2BFD08F76A" blockId="16.[449,908,235,2181]" pageId="16" pageNumber="182">
|
||
<emphasis id="B9080A263634FF7C84DBFF2BFD8CFEE0" box="[461,565,238,271]" italics="true" pageId="16" pageNumber="182">Figure</emphasis>
|
||
10.15.
|
||
<emphasis id="B9080A263634FF7C87AFFF28FDB5FEAB" italics="true" pageId="16" pageNumber="182">Reconstructed</emphasis>
|
||
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>
|
||
based on results of Figures
|
||
<emphasis id="B9080A263634FF7C84D9FE30FD9FFDF7" box="[463,550,501,536]" italics="true" pageId="16" pageNumber="182">10.13</emphasis>
|
||
and
|
||
<emphasis id="B9080A263634FF7C879FFE30FCEAFDF7" box="[649,851,501,536]" italics="true" pageId="16" pageNumber="182">10.14. Deep</emphasis>
|
||
muscles
|
||
<emphasis id="B9080A263634FF7C8748FDEFFDC4FDA2" box="[606,637,554,589]" italics="true" pageId="16" pageNumber="182">in</emphasis>
|
||
lateral
|
||
<emphasis id="B9080A263634FF7C8614FDEFFC8DFDA2" box="[770,820,554,589]" italics="true" pageId="16" pageNumber="182">(A)</emphasis>
|
||
and anterior
|
||
<emphasis id="B9080A263634FF7C8749FD9AFCB9FD6D" box="[607,768,607,642]" italics="true" pageId="16" pageNumber="182">(B) views;</emphasis>
|
||
intermediate muscles in lateral
|
||
<emphasis id="B9080A263634FF7C84DCFD0CFE41FD03" box="[458,504,713,748]" italics="true" pageId="16" pageNumber="182">(C)</emphasis>
|
||
and
|
||
<emphasis id="B9080A263634FF7C8747FD0CFD4BFD03" box="[593,754,713,748]" italics="true" pageId="16" pageNumber="182">anterior (</emphasis>
|
||
D
|
||
<emphasis id="B9080A263634FF7C861AFD0CFCAFFD03" box="[780,790,713,748]" italics="true" pageId="16" pageNumber="182">)</emphasis>
|
||
views; surficial muscles
|
||
<emphasis id="B9080A263634FF7C8670FD3BFC3BFCCE" box="[870,898,766,801]" italics="true" pageId="16" pageNumber="182">in</emphasis>
|
||
lateral
|
||
<emphasis id="B9080A263634FF7C8756FCF6FDD3FCB9" box="[576,618,819,854]" italics="true" pageId="16" pageNumber="182">(E)</emphasis>
|
||
and anterior
|
||
<emphasis id="B9080A263634FF7C864EFCF6FCC6FCB9" box="[856,895,819,854]" italics="true" pageId="16" pageNumber="182">(F)</emphasis>
|
||
views. Scale
|
||
<emphasis id="B9080A263634FF7C87B0FCADFDB6FC2F" italics="true" pageId="16" pageNumber="182">in centimeters.</emphasis>
|
||
Abbreviations: b, M. brachialis; bb,
|
||
<emphasis id="B9080A263634FF7C87D3FC17FCD4FC1A" box="[709,877,978,1013]" italics="true" pageId="16" pageNumber="182">M. biceps</emphasis>
|
||
brachii; cb,
|
||
<emphasis id="B9080A263634FF7C8785FBC2FD06FBC5" box="[659,703,1031,1066]" italics="true" pageId="16" pageNumber="182">M.</emphasis>
|
||
coracobrachialis
|
||
<emphasis id="B9080A263634FF7C872CFBF9FD13FBB0" box="[570,682,1084,1119]" italics="true" pageId="16" pageNumber="182">brevis;</emphasis>
|
||
cbd,
|
||
<emphasis id="B9080A263634FF7C861BFBF9FC80FBB0" box="[781,825,1084,1119]" italics="true" pageId="16" pageNumber="182">M.</emphasis>
|
||
coracobrachialis
|
||
<emphasis id="B9080A263634FF7C87FEFBB4FCF4FB7B" box="[744,845,1137,1172]" italics="true" pageId="16" pageNumber="182">brevis</emphasis>
|
||
dorsalis; dc, M. deltoideus davicularis; ds, M. deltoideus
|
||
<emphasis id="B9080A263634FF7C8790FAD4FC80FADB" box="[646,825,1297,1332]" italics="true" pageId="16" pageNumber="182">scapularis;</emphasis>
|
||
hr, M. humeroradialis;
|
||
<emphasis id="B9080A263634FF7C860CFA80FCF9FA87" box="[794,832,1349,1384]" italics="true" pageId="16" pageNumber="182">Id,</emphasis>
|
||
tendon for
|
||
<emphasis id="B9080A263634FF7C8798FABFFD00FA72" box="[654,697,1402,1437]" italics="true" pageId="16" pageNumber="182">M.</emphasis>
|
||
latissimus dorsi;
|
||
<emphasis id="B9080A263634FF7C8739FA75FDF4FA3C" box="[559,589,1456,1491]" italics="true" pageId="16" pageNumber="182">p,</emphasis>
|
||
M. pectoralis; sb, M.
|
||
<emphasis id="B9080A263634FF7C872AFA21FD8CF9D3" italics="true" pageId="16" pageNumber="182">supracoracoideus brevis;</emphasis>
|
||
sc,
|
||
<emphasis id="B9080A263634FF7C876DF9DCFCB6F99D" italics="true" pageId="16" pageNumber="182">M. scapulohumeralis cranialis;</emphasis>
|
||
sed, M. scapulohumeralis
|
||
<emphasis id="B9080A263634FF7C862FF941FDA8F933" italics="true" pageId="16" pageNumber="182">caudalis</emphasis>
|
||
;
|
||
<emphasis id="B9080A263634FF7C8730F97CFDE0F933" box="[550,601,1721,1756]" italics="true" pageId="16" pageNumber="182">sci,</emphasis>
|
||
M.
|
||
<emphasis id="B9080A263634FF7C87B2F97CFDF0F8FE" bold="true" italics="true" pageId="16" pageNumber="182">supracoracoideus</emphasis>
|
||
intermedius;
|
||
<emphasis id="B9080A263634FF7C8621F92BFCEFF8FE" box="[823,854,1774,1809]" italics="true" pageId="16" pageNumber="182">si,</emphasis>
|
||
M.
|
||
<emphasis id="B9080A263634FF7C84EBF8E6FC89F8A9" bold="true" box="[509,816,1827,1862]" pageId="16" pageNumber="182">supracoracoideus</emphasis>
|
||
longus; tbi, M.
|
||
<emphasis id="B9080A263634FF7C878CF89DFCC3F894" box="[666,890,1880,1915]" italics="true" pageId="16" pageNumber="182">triceps brevis</emphasis>
|
||
intermedius;
|
||
<emphasis id="B9080A263634FF7C87BFF84BFDB6F80A" italics="true" pageId="16" pageNumber="182">
|
||
tll, M.
|
||
<emphasis id="B9080A263634FF7C860AF84BFDB6F80A" bold="true" italics="true" pageId="16" pageNumber="182">triceps</emphasis>
|
||
</emphasis>
|
||
longus lateralis; tm, M. terres major.
|
||
<emphasis id="B9080A263634FF7C87F1F832FD84F7A0" italics="true" pageId="16" pageNumber="182">Terminology</emphasis>
|
||
<emphasis id="B9080A263634FF7C8753F7E9FD61F7A0" bold="true" box="[581,728,2092,2127]" pageId="16" pageNumber="182">adapted</emphasis>
|
||
from Meers (2003).
|
||
</paragraph>
|
||
</caption>
|
||
<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">
|
||
<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>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343635FF7D84D8FF00F973FCDB" blockId="17.[452,1825,190,2293]" pageId="17" pageNumber="183">
|
||
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">
|
||
<emphasis id="B9080A263635FF7D8329FF3AFDF0FEB0" italics="true" pageId="17" pageNumber="183">Tyrannosaurus rex</emphasis>
|
||
</taxonomicName>
|
||
based mostly on
|
||
<materialsCitation id="3B14DC693635FF7D868CFEFCFB7EFEB0" ID-GBIF-Occurrence="2813095301" box="[922,1223,313,351]" collectionCode="FMNH" pageId="17" pageNumber="183" specimenCode="FMNH PR2081">FMNH PR2081</materialsCitation>
|
||
. In order to work out the forces acting on the forelimb, we start by modeling the forelimb as a third-class lever (
|
||
<figureCitation id="1347CAB13635FF7D8756FE6AFD43FE3A" box="[576,762,431,469]" captionStart="Fig. 10.16" captionText="Figure 10.16. Free-body diagram (simplified model) of the forelimb of FMNH PR2081. Abbreviations: MF, motive force; MFA, motive force arm; RF, resistive force; RFA, resistive force arm." figureDoi="http://doi.org/10.5281/zenodo.3942845" httpUri="https://zenodo.org/record/3942845/files/figure.png" pageId="17" pageNumber="183" targetBox="[1904,2352,853,1322]" targetPageId="17">Fig. 10.16</figureCitation>
|
||
). In a third-class lever, the effort force (M. biceps muscle) is applied between the fulcrum (elbow joint) and the resistance force. The elbow joint is a hinge where the humerus, ulna, and radius articulate. Of all of the muscles coordinating and controlling the movement of the elbow, the M. biceps
|
||
<emphasis id="B9080A263635FF7D862AFD5CFCEFFD50" box="[828,854,665,703]" italics="true" pageId="17" pageNumber="183">is</emphasis>
|
||
the most powerful flexor of the elbow joint (
|
||
<bibRefCitation id="EFEDABC53635FF7D8384FD5CFCA1FD15" author="Ozkaya, N. & Nordin, M." firstAuthor="Ozkaya" journalOrPublisher="Springer, New York" pageId="17" pageNumber="183" refId="ref10257" refString="Ozkaya, N., and Nordin, M. 1999. Fundamentals of Biomechanics: Equilibrium, Motion, and Deformation. Springer, New York." title="Fundamentals of Biomechanics: Equilibrium, Motion, and Deformation" type="book" year="1999">Ozkaya and Nordin 1999</bibRefCitation>
|
||
). Our model assumes that the M. biceps is the major flexor and that the line of action (the tension) at the biceps is vertical.
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343635FF7D870DFC8CFC09FAE4" blockId="17.[452,1825,190,2293]" pageId="17" pageNumber="183">
|
||
Anatomical measurements were used to derive the motive force arm (MFA) and the resistive force arm (
|
||
<emphasis id="B9080A263635FF7D812CFC41FB36FC45" box="[1082,1167,900,938]" italics="true" pageId="17" pageNumber="183">RFA</emphasis>
|
||
) (
|
||
<tableCitation id="C6FEE38F3635FF7D81A5FC41FAD5FC45" box="[1203,1388,900,938]" captionStart="Table 10.1" captionText="Table 10.1. Power AnalysisMeasurements Abbreviations.—MFA, motive force arm; RFA, restive force arm" pageId="17" pageNumber="183" targetBox="[1010,2246,256,606]" targetPageId="18">Table 10.1</tableCitation>
|
||
). For the ulna, the MFA was measured from the sigmoid notch to the midscar of the insertion point for the M. biceps (motive force, MF), and the
|
||
<emphasis id="B9080A263635FF7D81EEFC3FFAF2FBCF" bold="true" box="[1272,1355,1018,1056]" pageId="17" pageNumber="183">RFA</emphasis>
|
||
was derived from measuring the ulna from the sigmoid notch to the distal end (
|
||
<figureCitation id="1347CAB13635FF7D808AFBF0F9F7FBB4" box="[1436,1614,1077,1115]" captionStart="Fig. 10.16" captionText="Figure 10.16. Free-body diagram (simplified model) of the forelimb of FMNH PR2081. Abbreviations: MF, motive force; MFA, motive force arm; RF, resistive force; RFA, resistive force arm." figureDoi="http://doi.org/10.5281/zenodo.3942845" httpUri="https://zenodo.org/record/3942845/files/figure.png" pageId="17" pageNumber="183" targetBox="[1904,2352,853,1322]" targetPageId="17">Fig. 10.16</figureCitation>
|
||
).
|
||
<emphasis id="B9080A263635FF7D837DFBF0F925FBB4" box="[1643,1692,1077,1115]" italics="true" pageId="17" pageNumber="183">To</emphasis>
|
||
obtain the MFA of the radius, we measured from the radial head to the midscar of the insertion point for the M. biceps and from the radial head to the distal end to determine the RFA.
|
||
</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>
|
||
(
|
||
<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 (
|
||
<bibRefCitation id="EFEDABC53635FF7D87B9FA15FB91FA19" author="Van Snik, G. & Olmos, AL & Casinos, A. & Planell, J. A." box="[687,1064,1488,1526]" etAl="et al." firstAuthor="Van Snik" journalOrPublisher="Netherlands Journal of Zoology" pageId="17" pageNumber="183" pagination="1 - 14" part="44" refId="ref10762" refString="Van Snik, G., Olmos, AL, Casinos, A., and Planell, J. A. 1994. Stresses in leg tendons of birds. Netherlands Journal of Zoology 44: 1 - 14." title="Stresses in leg tendons of birds" type="journal article" year="1994">Van Snik et al. 1994</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFEDABC53635FF7D812CFA15FAEFFA19" author="Alexander, R. McN" box="[1082,1366,1488,1526]" firstAuthor="Alexander" journalOrPublisher="Science Progress" pageId="17" pageNumber="183" pagination="109 - 130" part="67" refId="ref8774" refString="Alexander, R. McN. 1981. Factors of safety in the structure of animals. Science Progress 67: 109 - 130." title="Factors of safety in the structure of animals" type="journal article" year="1981">Alexander 1981</bibRefCitation>
|
||
). A safety factor of 3 will be used
|
||
<emphasis id="B9080A263635FF7D8776F9C9FD3DF9DD" box="[608,644,1548,1586]" italics="true" pageId="17" pageNumber="183">in</emphasis>
|
||
this study.
|
||
</paragraph>
|
||
<caption id="DF0386BC3635FF7D8278FA65F717F8D9" ID-DOI="http://doi.org/10.5281/zenodo.3942845" ID-Zenodo-Dep="3942845" httpUri="https://zenodo.org/record/3942845/files/figure.png" pageId="17" pageNumber="183" startId="17.[1902,2006,1440,1475]" subCaptionStartIDs="17.[2173,2348,1599,1634]" subCaptionStarts="Abbr" targetBox="[1904,2352,853,1322]" targetPageId="17">
|
||
<paragraph id="8BC3D6343635FF7D8278FA65F717F8D9" blockId="17.[1899,2348,1370,1846]" pageId="17" pageNumber="183">
|
||
<emphasis id="B9080A263635FF7D8278FA65F86FFA2C" box="[1902,2006,1440,1475]" italics="true" pageId="17" pageNumber="183">Figure</emphasis>
|
||
<emphasis id="B9080A263635FF7D82F0FA65F7FDFA2C" bold="true" box="[2022,2116,1440,1475]" pageId="17" pageNumber="183">10.16</emphasis>
|
||
.
|
||
<emphasis id="B9080A263635FF7D8D41FA65F7ACFA18" italics="true" pageId="17" pageNumber="183">Free-body diagram (</emphasis>
|
||
<emphasis id="B9080A263635FF7D8D03FA11F705FA18" bold="true" box="[2069,2236,1492,1527]" pageId="17" pageNumber="183">simplified</emphasis>
|
||
model
|
||
<emphasis id="B9080A263635FF7D82CFF9CFF85FF9C2" box="[2009,2022,1546,1581]" italics="true" pageId="17" pageNumber="183">)</emphasis>
|
||
of the forelimb of
|
||
<emphasis id="B9080A263635FF7D827DF9FAF723F978" italics="true" pageId="17" pageNumber="183">
|
||
<materialsCitation id="3B14DC693635FF7D827DF9FAF7D2F98D" ID-GBIF-Occurrence="2813095317" box="[1899,2155,1599,1634]" collectionCode="FMNH" pageId="17" pageNumber="183" specimenCode="FMNH PR2081">
|
||
FMNH PR
|
||
<emphasis id="B9080A263635FF7D8D0EF9FAF7D2F98D" bold="true" box="[2072,2155,1599,1634]" italics="true" pageId="17" pageNumber="183">2081</emphasis>
|
||
</materialsCitation>
|
||
. Abbreviations: MF, motive
|
||
</emphasis>
|
||
<emphasis id="B9080A263635FF7D8DB3F9B1F746F978" bold="true" box="[2213,2303,1652,1687]" pageId="17" pageNumber="183">force</emphasis>
|
||
;
|
||
<emphasis id="B9080A263635FF7D827DF96CF87CF923" box="[1899,1989,1705,1740]" italics="true" pageId="17" pageNumber="183">MFA,</emphasis>
|
||
motive
|
||
<emphasis id="B9080A263635FF7D8D40F96CF6B0F923" box="[2134,2313,1705,1740]" italics="true" pageId="17" pageNumber="183">force arm;</emphasis>
|
||
RF,
|
||
<emphasis id="B9080A263635FF7D82BCF91BF78AF8EE" box="[1962,2099,1758,1793]" italics="true" pageId="17" pageNumber="183">resistive</emphasis>
|
||
<emphasis id="B9080A263635FF7D8D29F91BF721F8EE" bold="true" box="[2111,2200,1758,1793]" pageId="17" pageNumber="183">force</emphasis>
|
||
;
|
||
<emphasis id="B9080A263635FF7D8DB8F91BF740F8EE" box="[2222,2297,1758,1793]" italics="true" pageId="17" pageNumber="183">RFA,</emphasis>
|
||
resistive force arm.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BC3D6343635FF7D870EF983FAF9F87D" blockId="17.[452,1825,190,2293]" pageId="17" pageNumber="183">
|
||
The normal working range (NWR) is one-third the safety factor (
|
||
<bibRefCitation id="EFEDABC53635FF7D83D1F983FCCDF948" author="Carpenter, K. & Smith, M." 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>
|
||
). Although the size, shape, and the biomechanical behavior of each tendon differs, the basic structure of tendons and their mechanical properties are similar (Jozsa and Kannus 1997). The
|
||
<emphasis id="B9080A263635FF7D8377F933F91CF8F3" box="[1633,1701,1782,1820]" italics="true" pageId="17" pageNumber="183">size</emphasis>
|
||
of the cross-sectional area of a tendon is directly related to the size of the load that can be carried before failure (
|
||
<bibRefCitation id="EFEDABC53635FF7D86E6F8A9FA96F87D" author="Butler. D. L. & Grood, E. S. & Noyes, F. R. & Zernicke, R. F." box="[1008,1327,1900,1938]" etAl="et al." firstAuthor="Butler" journalOrPublisher="Exercise Sports Science Review" pageId="17" pageNumber="183" pagination="125 - 181" part="6" refId="ref8930" refString="Butler. D. L., Grood, E. S., Noyes, F. R., and Zernicke, R. F. 1978. Biomechanics of ligaments and tendons. Exercise Sports Science Review 6: 125 - 181." title="Biomechanics of ligaments and tendons" type="journal article" year="1978">Butler et al. 1978</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343635FF7D870EF863FA66F71C" blockId="17.[452,1825,190,2293]" pageId="17" pageNumber="183">
|
||
The surface area of the scar for the insertion of the M. biceps is 122.11 mm2 on the radius and 192 mm2 on the ulna. The
|
||
<emphasis id="B9080A263635FF7D80C0F824F920F7E8" box="[1494,1689,2017,2055]" italics="true" pageId="17" pageNumber="183">conversion</emphasis>
|
||
for the tendon strength, expressed as MPa, is 1 MPa = 1,000,000 Pa, with 1 Pa = 1 N/m2. Therefore, 1 MPa = 1 N/mm2. The maximum working range (MWR) and the NWR are calculated from the tensile strength of the tendon. The formula for estimated tendon tensile strength is as follows:
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343635FF7D873BF6F5F8ADF67E" blockId="17.[524,1812,2345,2450]" pageId="17" pageNumber="183">
|
||
tendon tensile strength/area2
|
||
<emphasis id="B9080A263635FF7D8166F6F5FB3FF6B8" bold="true" box="[1136,1158,2352,2391]" pageId="17" pageNumber="183">x</emphasis>
|
||
surface area of the scar for insertion of the M. biceps = estimated tendon tensile strength (1)
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343635FF7D84D5F615FB1FF619" blockId="17.[451,1190,2506,2553]" box="[451,1190,2512,2550]" pageId="17" pageNumber="183">Tendon tensile strength for the radius is</paragraph>
|
||
<paragraph id="8BC3D6343635FF7D87E8F5F6FA69F5B6" blockId="17.[766,1488,2605,2649]" box="[766,1488,2611,2649]" pageId="17" pageNumber="183">100 N/mm2 x 122.11 mm2 = 12,211 N</paragraph>
|
||
<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">
|
||
<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>
|
||
AnalysisMeasurements Abbreviations.—MFA, motive force arm; RFA, restive force arm.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BC3D6343636FF7E86E3FEC5F77FFDB1" pageId="18" pageNumber="184">
|
||
<table id="F97C24943636009386E4FEC5F77FFDB1" box="[1010,2246,256,606]" gridcols="3" gridrows="5" pageId="18" pageNumber="184">
|
||
<tr id="354CD4763636009386E4FEC5F77FFEC7" box="[1010,2246,256,296]" gridrow="0" pageId="18" pageNumber="184">
|
||
<th id="769DBD0A3636009386E4FEC5FAE8FEC7" box="[1010,1361,256,296]" gridcol="0" gridrow="0" pageId="18" pageNumber="184">Measurement</th>
|
||
<th id="769DBD0A3636009380B5FEC5F8BFFEC7" box="[1443,1798,256,296]" gridcol="1" gridrow="0" pageId="18" pageNumber="184">Radius</th>
|
||
<th id="769DBD0A363600938275FEC5F77FFEC7" box="[1891,2246,256,296]" gridcol="2" gridrow="0" pageId="18" pageNumber="184">Ulna</th>
|
||
</tr>
|
||
<tr id="354CD4763636009386E4FE9CF77FFE94" box="[1010,2246,345,379]" gridrow="1" pageId="18" pageNumber="184">
|
||
<th id="769DBD0A3636009386E4FE9CFAE8FE94" box="[1010,1361,345,379]" gridcol="0" gridrow="1" pageId="18" pageNumber="184">MFA</th>
|
||
<td id="769DBD0A3636009380B5FE9CF8BFFE94" box="[1443,1798,345,379]" gridcol="1" gridrow="1" pageId="18" pageNumber="184">15.2 mm (0.0152 m)</td>
|
||
<td id="769DBD0A363600938275FE9CF77FFE94" box="[1891,2246,345,379]" gridcol="2" gridrow="1" pageId="18" pageNumber="184">45.8 mm (0.0458 m)</td>
|
||
</tr>
|
||
<tr id="354CD4763636009386E4FE60F77FFE28" box="[1010,2246,421,455]" gridrow="2" pageId="18" pageNumber="184">
|
||
<th id="769DBD0A3636009386E4FE60FAE8FE28" box="[1010,1361,421,455]" gridcol="0" gridrow="2" pageId="18" pageNumber="184">RFA</th>
|
||
<td id="769DBD0A3636009380B5FE60F8BFFE28" box="[1443,1798,421,455]" gridcol="1" gridrow="2" pageId="18" pageNumber="184">166.2 mm (0.166 m)</td>
|
||
<td id="769DBD0A363600938275FE60F77FFE28" box="[1891,2246,421,455]" gridcol="2" gridrow="2" pageId="18" pageNumber="184">186.6 mm (0.187 m)</td>
|
||
</tr>
|
||
<tr id="354CD4763636009386E4FE34F77FFDFC" box="[1010,2246,497,531]" gridrow="3" pageId="18" pageNumber="184">
|
||
<th id="769DBD0A3636009386E4FE34FAE8FDFC" box="[1010,1361,497,531]" gridcol="0" gridrow="3" pageId="18" pageNumber="184">MANUS</th>
|
||
<td id="769DBD0A3636009380B5FE34F8BFFDFC" box="[1443,1798,497,531]" gridcol="1" gridrow="3" pageId="18" pageNumber="184">177.6 mm (0.178 m)</td>
|
||
<td id="769DBD0A363600938275FE34F77FFDFC" box="[1891,2246,497,531]" gridcol="2" gridrow="3" pageId="18" pageNumber="184">177.6 mm (0.178 m)</td>
|
||
</tr>
|
||
<tr id="354CD4763636009386E4FDF9F77FFDB1" box="[1010,2246,572,606]" gridrow="4" pageId="18" pageNumber="184">
|
||
<th id="769DBD0A3636009386E4FDF9FAE8FDB1" box="[1010,1361,572,606]" gridcol="0" gridrow="4" pageId="18" pageNumber="184">RFA including manus</th>
|
||
<td id="769DBD0A3636009380B5FDF9F8BFFDB1" box="[1443,1798,572,606]" gridcol="1" gridrow="4" pageId="18" pageNumber="184">343.8 mm (0.344 m)</td>
|
||
<td id="769DBD0A363600938275FDF9F77FFDB1" box="[1891,2246,572,606]" gridcol="2" gridrow="4" pageId="18" pageNumber="184">364.2 mm (0.364 m)</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<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>
|
||
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">
|
||
<emphasis id="B9080A263636FF7E8D45FC6DFBA2FBE6" italics="true" pageId="18" pageNumber="184">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
can resist (resistive force, or RF):
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E80EFFB92F69BFB92" blockId="18.[1529,2353,1106,1149]" box="[1529,2338,1111,1149]" pageId="18" pageNumber="184">MFxMFA = T (2)</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E831FFB0BF69AFB1B" blockId="18.[1545,2353,1223,1268]" box="[1545,2339,1230,1268]" pageId="18" pageNumber="184">RF x RFA = T (3)</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E8120FA81F860FA0F" blockId="18.[996,2358,1341,1563]" pageId="18" pageNumber="184">
|
||
Measurement of the manus (177.6 mm) was taken from a cast of
|
||
<materialsCitation id="3B14DC693636FF7E86F3FABAFAA3FA4A" ID-GBIF-Occurrence="2813095329" box="[997,1306,1407,1445]" collectionCode="FMNH" pageId="18" pageNumber="184" specimenCode="FMNH PR2081">FMNH PR 2081</materialsCitation>
|
||
, from the proximal end of the wrist to the proximal end of the claws. It was then added to the RFA (166.2 mm).
|
||
</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">
|
||
T.
|
||
<emphasis id="B9080A263636FF7E8300FA30F9F2F9F4" box="[1558,1611,1525,1563]" italics="true" pageId="18" pageNumber="184">rex</emphasis>
|
||
</taxonomicName>
|
||
is as follows:
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E802AF9AFF87DF97F" blockId="18.[1340,1988,1637,1801]" box="[1340,1988,1642,1680]" pageId="18" pageNumber="184">
|
||
4, 070 N
|
||
<emphasis id="B9080A263636FF7E80E5F9AFF9B3F97F" bold="true" box="[1523,1546,1642,1680]" pageId="18" pageNumber="184">x</emphasis>
|
||
0.0152 m = 61.86 Nm
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E8097F960F838F924" blockId="18.[1340,1988,1637,1801]" box="[1409,1921,1701,1739]" pageId="18" pageNumber="184">RF x 03438 m = 61.86 Nm</paragraph>
|
||
<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">
|
||
T.
|
||
<emphasis id="B9080A263636FF7E80ACF893FA56F893" box="[1466,1519,1878,1916]" italics="true" pageId="18" pageNumber="184">rex</emphasis>
|
||
</taxonomicName>
|
||
is as follows:
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E8050F80EF804F81E" blockId="18.[1350,1981,1991,2151]" box="[1350,1981,1995,2033]" pageId="18" pageNumber="184">
|
||
1,357 N
|
||
<emphasis id="B9080A263636FF7E80FCF80EF9B9F81E" bold="true" box="[1514,1536,1995,2033]" pageId="18" pageNumber="184">x</emphasis>
|
||
0.0152 m = 20.63 Nm
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E8060F7C3F833F7C3" blockId="18.[1350,1981,1991,2151]" box="[1398,1930,2054,2092]" pageId="18" pageNumber="184">
|
||
RF
|
||
<emphasis id="B9080A263636FF7E80A7F7C3FA7FF7C3" bold="true" box="[1457,1478,2054,2092]" pageId="18" pageNumber="184">x</emphasis>
|
||
0.3438 m = 20.63 Nm
|
||
</paragraph>
|
||
<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">
|
||
T.
|
||
<emphasis id="B9080A263636FF7E80B3F772FA63F732" box="[1445,1498,2231,2269]" italics="true" pageId="18" pageNumber="184">rex</emphasis>
|
||
</taxonomicName>
|
||
is as follows:
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E8024F6E9F872F6BD" blockId="18.[1330,1995,2344,2504]" box="[1330,1995,2348,2386]" pageId="18" pageNumber="184">
|
||
6,400 N
|
||
<emphasis id="B9080A263636FF7E80F6F6E9FA4FF6BD" bold="true" box="[1504,1526,2348,2386]" pageId="18" pageNumber="184">x</emphasis>
|
||
0.0458 m = 293.12 Nm
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E807AF6A2F82BF662" blockId="18.[1330,1995,2344,2504]" box="[1388,1938,2407,2445]" pageId="18" pageNumber="184">RF x 0.3642 m = 293.12 Nm</paragraph>
|
||
<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">
|
||
T.
|
||
<emphasis id="B9080A263636FF7E808BF5D2FA69F5D2" box="[1437,1488,2583,2621]" italics="true" pageId="18" pageNumber="184">rex</emphasis>
|
||
</taxonomicName>
|
||
is as follows:
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E8054F548F800F55C" blockId="18.[1346,1977,2696,2857]" box="[1346,1977,2701,2739]" pageId="18" pageNumber="184">
|
||
2,133 N
|
||
<emphasis id="B9080A263636FF7E80F3F548FA42F55C" bold="true" box="[1509,1531,2701,2739]" pageId="18" pageNumber="184">x</emphasis>
|
||
0.0458 m = 97.69 Nm
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E8061F502F83FF502" blockId="18.[1346,1977,2696,2857]" box="[1399,1926,2759,2797]" pageId="18" pageNumber="184">
|
||
RF
|
||
<emphasis id="B9080A263636FF7E80A7F502FA7EF502" bold="true" box="[1457,1479,2759,2797]" pageId="18" pageNumber="184">x</emphasis>
|
||
0.3642 m = 97.69 Nm
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343636FF7E8064F4C7F833F4C7" blockId="18.[1346,1977,2696,2857]" box="[1394,1930,2818,2856]" pageId="18" pageNumber="184">RF = 268.23 N (or 27.37 kg)</paragraph>
|
||
<paragraph id="8BC3D6343636FF7F8126F4B2FD6AFEB9" blockId="18.[992,2349,2928,3036]" lastBlockId="19.[474,1840,181,1051]" lastPageId="19" lastPageNumber="185" pageId="18" pageNumber="184">
|
||
Adding the resistive forces of the radius and ulna results in 984.76 N (100.49 kg or 221.10 pounds) for the MWR (no safety factor) and 328.24 N (33.49 kg or 73.70 pounds) for the NWR (with safety factor). The conversion factor for kilograms to newtons is 1 kg = 9.8 N. These results are summarized in
|
||
<tableCitation id="C6FEE38F3637FF7F871BFEF5FD77FEB9" box="[525,718,304,342]" captionStart="Table 10.2" captionText="Table 10.2. Power Analysis Summary Abbreviations.—RF, resistive force; MWR, maximum working rang-; NWR, normal working range." pageId="19" pageNumber="185" targetBox="[478,1752,2662,2945]" targetPageId="19">Table 10.2</tableCitation>
|
||
.
|
||
</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>
|
||
(
|
||
<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>
|
||
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
|
||
<emphasis id="B9080A263637FF7F86A5FC72FC75FC32" box="[947,972,951,989]" italics="true" pageId="19" pageNumber="185">is</emphasis>
|
||
estimated to have been around 1150 kg, or 11,270 N. Of this, the biceps generated about 40%, and thus was a major muscle.
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343637FF7F8269FB6DF6AAFAB6" blockId="19.[1917,2358,1187,1369]" pageId="19" pageNumber="185">
|
||
<heading id="D08B61583637FF7F8269FB6DF6AAFAB6" bold="true" centered="true" fontSize="14" level="7" pageId="19" pageNumber="185" reason="0">
|
||
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>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343637FF7F84C0FB6CFA26FA1A" blockId="19.[469,1836,1186,2410]" pageId="19" pageNumber="185">
|
||
A small lever arm requires greater muscle tension to balance a load. Therefore, while resisting prey or holding prey, it is disadvantageous to have a muscle attachment close to the elbow joint. The advantage to having the muscle attachment close to elbow joint is that it will have a larger range of motion of the elbow flexion-extension, and therefore the hand can move faster toward the upper arm or shoulder (
|
||
<bibRefCitation id="EFEDABC53637FF7F86DCFA0AFA37FA1A" author="Ozkaya, N. & Nordin, M." box="[970,1422,1487,1525]" firstAuthor="Ozkaya" journalOrPublisher="Springer, New York" pageId="19" pageNumber="185" refId="ref10257" refString="Ozkaya, N., and Nordin, M. 1999. Fundamentals of Biomechanics: Equilibrium, Motion, and Deformation. Springer, New York." title="Fundamentals of Biomechanics: Equilibrium, Motion, and Deformation" type="book" year="1999">Ozkaya and Nordin 1999</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343637FF7F873FF9CFFB18F90E" blockId="19.[469,1836,1186,2410]" pageId="19" pageNumber="185">
|
||
The mechanical advantage is the amount of force a given effort can produce. It can be expressed as a ratio of the resistive force to the MF, or as a ratio of the MFA to the RFA (
|
||
<bibRefCitation id="EFEDABC53637FF7F8157F945F938F949" author="Kreighbaum, E. & Barthels, K. M." box="[1089,1665,1664,1702]" firstAuthor="Kreighbaum" journalOrPublisher="Burgess Publishing, Minneapolis, MN" pageId="19" pageNumber="185" refId="ref9837" refString="Kreighbaum, E., and Barthels, K. M. 1985. Biomechanics: A Qualitative Approach for Studying Human Movement. Burgess Publishing, Minneapolis, MN." title="Biomechanics: A Qualitative Approach for Studying Human Movement" type="book" year="1985">Kreighbaum and Barthels 1985</bibRefCitation>
|
||
). Both of the equations produce the same result.
|
||
</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>
|
||
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>
|
||
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">
|
||
<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>
|
||
Summary Abbreviations.—RF,
|
||
<emphasis id="B9080A263637FF7F8DC7F50BF816F4CF" italics="true" pageId="19" pageNumber="185">resistive</emphasis>
|
||
force; MWR,
|
||
<emphasis id="B9080A263637FF7F8D89F53AF871F4A0" italics="true" pageId="19" pageNumber="185">maximum</emphasis>
|
||
working rang-; NWR,
|
||
<emphasis id="B9080A263637FF7F82F5F4A5F7ECF46E" box="[2019,2133,2912,2945]" italics="true" pageId="19" pageNumber="185">normal</emphasis>
|
||
working range.
|
||
</paragraph>
|
||
</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>
|
||
<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="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="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">
|
||
<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>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343630FF7884D5FF2AFD6BFEF1" blockId="20.[451,723,239,286]" box="[451,722,239,286]" pageId="20" pageNumber="186">
|
||
<heading id="D08B61583630FF7884D5FF2AFD6BFEF1" bold="true" box="[451,722,239,286]" fontSize="16" level="5" pageId="20" pageNumber="186" reason="0">Acceleration</heading>
|
||
</paragraph>
|
||
<paragraph id="8BC3D6343630FF7886CDFF33F9B8FB28" blockId="20.[978,2345,240,1226]" pageId="20" pageNumber="186">
|
||
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">
|
||
<emphasis id="B9080A263630FF788234FE63F792FE23" box="[1826,2091,422,460]" italics="true" pageId="20" pageNumber="186">Tyrannosaurus</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="1347CAB13630FF788D5EFE63FB87FDE8" captionStart-0="Fig. 10.15" captionStart-1="Fig. 10.16" captionStart-2="Fig. 10.17" captionText-0="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). " captionText-1="Figure 10.16. Free-body diagram (simplified model) of the forelimb of FMNH PR2081. Abbreviations: MF, motive force; MFA, motive force arm; RF, resistive force; RFA, resistive force arm." captionText-2="Figure 10.17. A 3-D representation of the forearm and manus in the Tyrannosaurus rex FMNH PR2081. In (A) lateral, (B) anterior, and (C) reaching views." figureDoi-0="http://doi.org/10.5281/zenodo.3942843" figureDoi-1="http://doi.org/10.5281/zenodo.3942845" figureDoi-2="http://doi.org/10.5281/zenodo.3942847" httpUri-0="https://zenodo.org/record/3942843/files/figure.png" httpUri-1="https://zenodo.org/record/3942845/files/figure.png" httpUri-2="https://zenodo.org/record/3942847/files/figure.png" pageId="20" pageNumber="186" targetBox-0="[993,2179,246,3040]" targetBox-1="[1904,2352,853,1322]" targetBox-2="[969,2327,2172,3055]" targetPageId-0="16" targetPageId-1="17" targetPageId-2="20">Figs. 10.15- 10.17</figureCitation>
|
||
was converted to
|
||
<emphasis id="B9080A263630FF788086FE24FA18FDE8" box="[1424,1441,481,519]" italics="true" pageId="20" pageNumber="186">a</emphasis>
|
||
closely packed series
|
||
<emphasis id="B9080A263630FF78822CFE24F8E6FDE8" box="[1850,1887,481,519]" italics="true" pageId="20" pageNumber="186">of</emphasis>
|
||
elliptical cylinders. The cross sections of each elliptical cylinder were determined from
|
||
<figureCitation id="1347CAB13630FF788D2BFDDEF6A2FDAE" box="[2109,2331,539,577]" 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="20" pageNumber="186" targetBox="[993,2179,246,3040]" targetPageId="16">Figure 10.15</figureCitation>
|
||
. Assuming a density of 1000 kg/m2 for tissue, the data from these cylinders result in a mass of 1.8 kg for the forearm plus manus. The integral of the density times the perpendicular distance to the pivot point results in a moment of inertia of 0.06 kg m2. Because the NWR torque of the forearm and hand to be 118.3 Nm, the angular acceleration (torque divided by the moment of inertia) is 1983 s-2. The angular acceleration can be converted to a linear acceleration by multiplying it by the distance (0.354 m) from the pivot point to the claws, which results in a linear acceleration of 702 ms-2. This is likely an overestimate because the skin and the claw sheath are not factored in. Also, this only gives the initial acceleration. The force from muscles is known to rapidly reduce at high speeds (
|
||
<bibRefCitation id="EFEDABC53630FF78805EFB64FA48FB28" author="Hill, A. V." box="[1352,1521,1185,1223]" firstAuthor="Hill" journalOrPublisher="Proceedings of the Royal Society of London" pageId="20" pageNumber="186" pagination="314 - 320" part="141" refId="ref9674" refString="Hill, A. V. 1938. The heat of shorting and dynamic constants of muscle. Proceedings of the Royal Society of London 141: 314 - 320." title="The heat of shorting and dynamic constants of muscle" type="journal article" year="1938">Hill 1938</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C36685BF3630FF7984ABFA96F9A2F9D0" lastPageId="21" lastPageNumber="187" pageId="20" pageNumber="186" type="discussion">
|
||
<paragraph id="8BC3D6343630FF7884ABFA96FD06FA91" blockId="20.[445,703,1358,1406]" box="[445,703,1363,1406]" pageId="20" pageNumber="186">
|
||
<heading id="D08B61583630FF7884ABFA96FD06FA91" bold="true" box="[445,703,1363,1406]" fontSize="14" level="6" pageId="20" pageNumber="186" reason="0">Conclusions</heading>
|
||
</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>
|
||
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>
|
||
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>
|
||
,
|
||
</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
|
||
<taxonomicName id="4C7CADB73631FF798091FAADF906FA61" box="[1415,1727,1384,1422]" class="Reptilia" family="Tyrannosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="21" pageNumber="187" phylum="Chordata" rank="family">Tyrannosauridae</taxonomicName>
|
||
(
|
||
<figureCitation id="1347CAB13631FF7983F7FAA2FD84FA22" captionStart="Fig. 10.18" captionText="Figure 10.18. Comparison of forelimb length to hind limb length shows that a progressive reduction in forelimb length does not occur in the Tyrannosauridae. Abbreviations: Gu, Guanlong (basal tyrannosauroid); Go, Gorgosaurus; Da, Daspletosaurus; Ab, Albertosaurus; T, Tyrannosaurus " figureDoi="http://doi.org/10.5281/zenodo.3942849" httpUri="https://zenodo.org/record/3942849/files/figure.png" pageId="21" pageNumber="187" targetBox="[473,1825,173,1236]" targetPageId="21">Fig. 10.18</figureCitation>
|
||
), contrary to Paul (1988) and Lockley et al. (this volume). In point of fact, once the shortened forelimb of the tyrannosaurids was established, it remained proportionally stable relative to hindlimb length.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |