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<document id="D60D6255D2F70908D988EB45A8E55192" ID-DOI="10.2110/palo.2009.p09-056r" ID-GBIF-Dataset="f8eb471f-471a-4b29-8a58-ef3da4eb2c87" ID-Zenodo-Dep="3942885" IM.bibliography_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1592905825948" checkinUser="jeremy" docAuthor="Joseph E. Peterson, Michael D. Henderson, Reed P. Scherer &amp; Christopher P. Vittore" docDate="2009" docId="03F987D85A3EFFFBD9E6FC051AF4FC10" docLanguage="en" docName="Petersonetal2009.pdf.imf" docOrigin="Palaios 24" docStyle="DocumentStyle{}" docTitle="Tyrannosaurus rex Osborn 1905" docType="treatment" docVersion="10" lastPageNumber="783" masterDocId="FFC0FFA05A3EFFF8D97CF9081F50FFEB" masterDocTitle="Face biting on a juvenile tyrannosaurid and behavioral implications" masterLastPageNumber="784" masterPageNumber="780" pageNumber="780" updateTime="1736513676254" updateUser="felipe">
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<mods:title id="7261335D5D1DD8F95F9CB863930F68CB">Face biting on a juvenile tyrannosaurid and behavioral implications</mods:title>
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<mods:namePart id="F3F2680F2C1E0448FAC40AD9F075CED2">Joseph E. Peterson</mods:namePart>
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<mods:namePart id="4FACE4340BCD12576FAA37891EC0609B">Michael D. Henderson</mods:namePart>
<mods:affiliation id="32F3E65C9F1B3815D491D4B4820751A1">Department of Earth and Environmental Sciences, Northern Illinois University, DeKalb, Illinois, 60115, USA, Burpee Museum of Natural History, 737 North Main Street, Rockford, Illinois, 61103, USA</mods:affiliation>
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<mods:namePart id="00B84526096118C2397D355A37FD7608">Reed P. Scherer</mods:namePart>
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<mods:namePart id="CA10FE3430B4017FFC0BBEF10FF4496D">Christopher P. Vittore</mods:namePart>
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Bite marks preserved on bone are not uncommon in the vertebrate fossil record and have the potential to yield insights into aspects of behavioral evolution and ecology (
<bibRefCitation id="EFC14B3F5A3EFFF8D88DFC4E1DDEFAB7" author="BUFFETAUT, E." box="[497,654,1350,1372]" journalOrPublisher="Palaontologische Zeitschrift" pageId="0" pageNumber="780" pagination="143 - 145" part="57" refId="ref3266" refString="BUFFETAUT, E., 1983, Wounds on the jaw of an Eocene mesosuchian crocodilian as possible evidence for the antiquity of crocodile intraspecific fighting behavior: Palaontologische Zeitschrift, v. 57, p. 143 - 145." title="Wounds on the jaw of an Eocene mesosuchian crocodilian as possible evidence for the antiquity of crocodile intraspecific fighting behavior" type="book chapter" year="1983">Buffetaut, 1983</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3EFFF8DBDFFC4E1E41FA93" author="CURRIE, P. J. &amp; JACOBSEN, A. R." journalOrPublisher="Canadian Journal of Earth Sciences" pageId="0" pageNumber="780" pagination="922 - 925" part="32" refId="ref3534" refString="CURRIE, P. J., and JACOBSEN, A. R., 1995, An azhdarchid pterosaur eaten by a velociraptorine theropod: Canadian Journal of Earth Sciences, v. 32, p. 922 - 925." title="An azhdarchid pterosaur eaten by a velociraptorine theropod" type="book chapter" year="1995">Currie and Jacobsen, 1995</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3EFFF8D860FC6A1E8BFA93" author="ERICKSON, G. M. &amp; VAN KIRK, S. D. &amp; SU, J. &amp; LEVENSON, M. E. &amp; CALER, W. E. &amp; CARTER, D. R." box="[284,475,1378,1400]" journalOrPublisher="Nature" pageId="0" pageNumber="780" pagination="706 - 708" part="382" refId="ref3675" refString="ERICKSON, G. M., VAN KIRK, S. D., SU, J., LEVENSON, M. E., CALER, W. E., and CARTER, D. R., 1996, Bite-force estimation for Tyrannosaurus rex from tooth-marked bones: Nature, v. 382, p. 706 - 708." title="Bite-force estimation for Tyrannosaurus rex from tooth-marked bones" type="book chapter" year="1996">Erickson et al., 1996</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3EFFF8D899FC6A1D93FA93" author="TANKE, D. H. &amp; CURRIE, P. J." box="[485,707,1378,1400]" journalOrPublisher="Gaia" pageId="0" pageNumber="780" pagination="167 - 184" part="15" refId="ref4344" refString="TANKE, D. H., and CURRIE, P. J., 1998, Head-biting behavior in theropod dinosaurs: Paleopathological evidence: Gaia, v. 15, p. 167 - 184." title="Head-biting behavior in theropod dinosaurs: Paleopathological evidence" type="book chapter" year="1998">Tanke and Currie, 1998</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3EFFF8DBB2FC6B1FE3FA7E" author="HANNA, R. R." journalOrPublisher="Journal of Vertebrate Paleontology" pageId="0" pageNumber="780" pagination="76 - 90" part="22" refId="ref3777" refString="HANNA, R. R., 2002, Multiple injury and infection in a sub-adult theropod dinosaur Allosaurus fragilis with comparisons to allosaur pathology in the Cleveland-Lloyd Dinosaur Quarry collection: Journal of Vertebrate Paleontology, v. 22, p. 76 - 90." title="Multiple injury and infection in a sub-adult theropod dinosaur Allosaurus fragilis with comparisons to allosaur pathology in the Cleveland-Lloyd Dinosaur Quarry collection" type="book chapter" year="2002">Hanna, 2002</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3EFFF8D9BCFC771E23FA7E" author="AVILLA, L. S. &amp; FERNANDES, R. &amp; RAMOS, D. F. B." box="[192,371,1407,1429]" journalOrPublisher="Journal of Vertebrate Paleonotology" pageId="0" pageNumber="780" pagination="971 - 973" part="24" refId="ref3100" refString="AVILLA, L. S., FERNANDES, R., and RAMOS, D. F. B., 2004, Bite marks on a crocodylomorph from the Upper Cretaceous of Brazil: Evidence of social behavior?: Journal of Vertebrate Paleonotology, v. 24, p. 971 - 973." title="Bite marks on a crocodylomorph from the Upper Cretaceous of Brazil: Evidence of social behavior?" type="journal article" year="2004">Avilla et al., 2004</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3EFFF8D8FCFC771D5AFA7E" author="KATSURA, Y." box="[384,522,1407,1429]" journalOrPublisher="Historical Biology" pageId="0" pageNumber="780" pagination="93 - 97" part="16" refId="ref4203" refString="KATSURA, Y., 2004, Paleopathology of Toyotamaphimeia machikanensis (Diapsida, Crocodylia) from the Middle Pleistocene of Central Japan: Historical Biology, v. 16, p. 93 - 97." title="Paleopathology of Toyotamaphimeia machikanensis (Diapsida, Crocodylia) from the Middle Pleistocene of Central Japan" type="book chapter" year="2004">Katsura, 2004</bibRefCitation>
). Determining whether bite marks and other injuries preserved on fossil bones are the result of interspecific or intraspecific conflict, predation, post-mortem scavenging or disease, however, is often difficult. Furthermore, positive identification of the species that caused the injury and inferences about behavior from such traces are far more challenging tasks. Here we report a juvenile tyrannosaurid skeleton (
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) that exhibits a series of partially healed lesions along the left maxilla and nasal, which we interpret as a bite mark injury. By utilizing the Extant Phylogenetic Bracket (EPB) method (
<bibRefCitation id="EFC14B3F5A3EFFF8D899FF771D37F97E" author="WITMER, L. M." box="[485,615,1663,1685]" editor="Thomason, J." journalOrPublisher="Cambridge University Press, Cambridge, UK" pageId="0" pageNumber="780" pagination="1933" refId="ref4517" refString="WITMER, L. M., 1995, The extant phylogenetic bracket and the importance of reconstructing soft tissues in fossils, in Thomason, J., ed., Functional Morphology in Vertebrate Paleontology: Cambridge University Press, Cambridge, UK, p. 1933." title="The extant phylogenetic bracket and the importance of reconstructing soft tissues in fossils" type="book chapter" volumeTitle="Functional Morphology in Vertebrate Paleontology" year="1995">Witmer, 1995</bibRefCitation>
), we interpret this injury as resulting from intraspecific aggression between two similarsized juvenile tyrannosaurids and discuss the insights the specimen provides for juvenile tyrannosaurid behavior.
</paragraph>
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<heading id="D0A781A25A3EFFF8D826FE1D1D6DF8C0" allCaps="true" bold="true" box="[346,573,1813,1835]" centered="true" fontSize="9" level="1" pageId="0" pageNumber="780" reason="0">GEOLOGIC SETTING</heading>
</paragraph>
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Specimen
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was collected from the upper half of the Hell Creek Formation of western Carter County, Montana in 2002 (
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) and is latest Maastrichtian in age. Exact coordinates of the collection site are on file in the paleontology collections at the Burpee Museum of Natural History, Rockford, Illinois (BMR), where the specimen is housed. The tyrannosaurid skeleton was buried in a lenticular clay-ball conglomerate unit composed of poorly sorted sand, silt, rounded green clay rip-up clasts, and diagenetically produced siderite nodules that partly encased several bones. The bone-bearing unit displayed a rapid lateral variation in thickness from 12 to 40 cm and was overlain by a massive, poorly cemented, cross-bedded sandstone (
<bibRefCitation id="EFC14B3F5A3EFFF8DAE7FA081B90FCFD" author="HENDERSON, M. D. &amp; HARRISON, W. H." box="[923,1216,768,790]" editor="Larson, P. &amp; Carpenter, K." journalOrPublisher="Indiana University Press, Bloomington" pageId="0" pageNumber="780" pagination="82 - 90" refId="ref3912" refString="HENDERSON, M. D., and HARRISON, W. H., 2008, Taphonomy and environment of deposition of juvenile tyrannosaurid skeleton from the Hell Creek Formation (latest Maastrichtian) of southeastern Montana, in: Larson, P., and Carpenter, K., eds., Tyrannosaurus rex, the Tyrant King: Indiana University Press, Bloomington, p. 82 - 90" title="Taphonomy and environment of deposition of juvenile tyrannosaurid skeleton from the Hell Creek Formation (latest Maastrichtian) of southeastern Montana" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">Henderson and Harrison, 2008</bibRefCitation>
).
</paragraph>
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In southeastern Montana the Hell Creek Formation averages 150 meters in thickness (
<bibRefCitation id="EFC14B3F5A3EFFF8DD60FA311B8DFCA4" author="JOHNSON, K. R. &amp; NICHOLS, D. J. &amp; HARTMAN, J. H." box="[1052,1245,825,847]" editor="Hartman, J. H. &amp; Johnson, K. R. &amp; Nichols, D. J." journalOrPublisher="Geological Society of America Special Paper" pageId="0" pageNumber="780" pagination="503 - 510" part="361" refId="ref4071" refString="JOHNSON, K. R., NICHOLS, D. J., and HARTMAN, J. H., 2002, Hell Creek Formation: A 2001 Synthesis, in Hartman, J. H., Johnson, K. R., and Nichols, D. J., eds., The Hell Creek Formation and the Cretaceous-Tertiary boundary in the Northern Great Plains: An Integrated Continental Record of the End of the Cretaceous, Geological Society of America Special Paper, v. 361, p. 503 - 510." title="Hell Creek Formation: A 2001 Synthesis" type="journal article" volumeTitle="The Hell Creek Formation and the Cretaceous-Tertiary boundary in the Northern Great Plains: An Integrated Continental Record of the End of the Cretaceous" year="2002">Johnson et al., 2002</bibRefCitation>
) and contains a diverse and well-preserved latest Cretaceous biota. The formation is especially well known for its fossil vertebrates, which include fish, amphibians, aquatic and terrestrial reptiles, mammals, dinosaurs, and rarely pterosaurs (
<bibRefCitation id="EFC14B3F5A3EFFF8DADDFAA21B6BFC2B" author="FASTOVSKY, D. E." box="[929,1083,938,960]" journalOrPublisher="PALAIOS" pageId="0" pageNumber="780" pagination="282 - 295" part="2" refId="ref3740" refString="FASTOVSKY, D. E., 1987, Paleoenvironments of vertebrate-bearing strata during the Cretaceous-Paleogene transition, eastern Montana and western North Dakota: PALAIOS, v. 2, p. 282 - 295." title="Paleoenvironments of vertebrate-bearing strata during the Cretaceous-Paleogene transition, eastern Montana and western North Dakota" type="journal article" year="1987">Fastovsky, 1987</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3EFFF8DD3BFAA21BA1FC2B" author="WHITE, P. D. &amp; FASTOVSKY, D. E. &amp; SHEEHAN, P. M." box="[1095,1265,938,960]" journalOrPublisher="PALAIOS" pageId="0" pageNumber="780" pagination="41 - 51" part="13" refId="ref4567" refString="WHITE, P. D., FASTOVSKY, D. E., and SHEEHAN, P. M., 1998, Taphonomy and suggested structure of the dinosaurian assemblage of the Hell Creek Formation (Maastrichtian), eastern Montana and western North Dakota: PALAIOS, v. 13, p. 41 - 51." title="Taphonomy and suggested structure of the dinosaurian assemblage of the Hell Creek Formation (Maastrichtian), eastern Montana and western North Dakota" type="journal article" year="1998">White et al., 1998</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3EFFF8DD81FAA21AEDFC2B" author="JOHNSON, K. R. &amp; NICHOLS, D. J. &amp; HARTMAN, J. H." box="[1277,1469,938,960]" editor="Hartman, J. H. &amp; Johnson, K. R. &amp; Nichols, D. J." journalOrPublisher="Geological Society of America Special Paper" pageId="0" pageNumber="780" pagination="503 - 510" part="361" refId="ref4071" refString="JOHNSON, K. R., NICHOLS, D. J., and HARTMAN, J. H., 2002, Hell Creek Formation: A 2001 Synthesis, in Hartman, J. H., Johnson, K. R., and Nichols, D. J., eds., The Hell Creek Formation and the Cretaceous-Tertiary boundary in the Northern Great Plains: An Integrated Continental Record of the End of the Cretaceous, Geological Society of America Special Paper, v. 361, p. 503 - 510." title="Hell Creek Formation: A 2001 Synthesis" type="journal article" volumeTitle="The Hell Creek Formation and the Cretaceous-Tertiary boundary in the Northern Great Plains: An Integrated Continental Record of the End of the Cretaceous" year="2002">Johnson et al., 2002</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3EFFF8DA4EFACE1B0BFC37" author="HENDERSON, M. D. &amp; PETERSON, J. E." box="[818,1115,966,988]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="0" pageNumber="780" pagination="192 - 195" part="26" refId="ref3985" refString="HENDERSON, M. D., and PETERSON, J. E., 2006, An azhdarchid pterosaur vertebra from the Hell Creek Formation (Maastrichtian) of southeastern Montana: Journal of Vertebrate Paleontology, v. 26, p. 192 - 195." title="An azhdarchid pterosaur vertebra from the Hell Creek Formation (Maastrichtian) of southeastern Montana" type="book chapter" year="2006">Henderson and Peterson, 2006</bibRefCitation>
). A diverse and well-preserved fauna and flora was recovered from the
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excavation site (
<bibRefCitation id="EFC14B3F5A3EFFF8DA44FAF61CAEFBFF" author="HARRISON, W. F. &amp; NICHOLS, D. J. &amp; HENDERSON, M. D." box="[824,1022,1022,1044]" editor="Yang, Q. &amp; Wang, Y. D. &amp; Weldon, E. A." journalOrPublisher="University of Science and Technology of China Press, Beijing" pageId="0" pageNumber="780" pagination="199" refId="ref3824" refString="HARRISON, W. F., NICHOLS, D. J., and HENDERSON, M. D., 2006, Jane's world: A life community in the upper Cretaceous, in Yang, Q., Wang, Y. D., and Weldon, E. A., eds., Ancient Life and Modern Approaches: Abstracts of the Second International Paleontological Congress: University of Science and Technology of China Press, Beijing, p. 199." title="Jane's world: A life community in the upper Cretaceous" type="proceedings paper" volumeTitle="Ancient Life and Modern Approaches: Abstracts of the Second International Paleontological Congress" year="2006">Harrison et al., 2006</bibRefCitation>
).
</paragraph>
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</paragraph>
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consists of 145 bones of a small tyrannosaurid, approximately seven meters in length. Based on histological characters, degree of skeletal fusion, and thin-walled bone structure,
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is considered a late juvenile animal rather than a subadult since it had not yet reached a phase of rapid growth (
<bibRefCitation id="EFC14B3F5A3EFFF8DC4CFDE91C30FAF8" author="ERICKSON, G. M. &amp; CURRIE, P. J. &amp; INOUYE, B. D. &amp; WINN, A. A." journalOrPublisher="Science" pageId="0" pageNumber="780" pagination="213 - 217" part="313" refId="ref3576" refString="ERICKSON, G. M., CURRIE, P. J., INOUYE, B. D., and WINN, A. A., 2006, Tyrannosaur life tables: An example of nonavian dinosaur population biology: Science, v. 313, p. 213 - 217." title="Tyrannosaur life tables: An example of nonavian dinosaur population biology" type="journal article" year="2006">Erickson et al., 2006</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3EFFF8DA16FDF51BDEFAF8" author="HENDERSON, M. D. &amp; HARRISON, W. H." box="[874,1166,1277,1299]" editor="Larson, P. &amp; Carpenter, K." journalOrPublisher="Indiana University Press, Bloomington" pageId="0" pageNumber="780" pagination="82 - 90" refId="ref3912" refString="HENDERSON, M. D., and HARRISON, W. H., 2008, Taphonomy and environment of deposition of juvenile tyrannosaurid skeleton from the Hell Creek Formation (latest Maastrichtian) of southeastern Montana, in: Larson, P., and Carpenter, K., eds., Tyrannosaurus rex, the Tyrant King: Indiana University Press, Bloomington, p. 82 - 90" title="Taphonomy and environment of deposition of juvenile tyrannosaurid skeleton from the Hell Creek Formation (latest Maastrichtian) of southeastern Montana" type="book chapter" volumeTitle="Tyrannosaurus rex, the Tyrant King" year="2008">Henderson and Harrison, 2008</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C34A65455A3EFFF9DA34FC121C7DF839" lastPageId="1" lastPageNumber="781" pageId="0" pageNumber="780" type="nomenclature">
<paragraph id="8BEF36CE5A3EFFF9DA34FC121C7DF839" blockId="0.[818,1473,1136,1357]" lastBlockId="1.[159,814,1667,2002]" lastPageId="1" lastPageNumber="781" pageId="0" pageNumber="780">
The skeleton was partially articulated when discovered, and consists of a nearly complete and well-preserved skull, found disarticulated above the animal’s hips, and a large portion of the postcranial skeleton. The depositional setting indicates rapid burial of the skeleton, and there is no evidence for predation or extensive scavenging. Cranial characters closely resemble those of another late Maastrichtian specimen (CMNH 7541 [Cleveland Museum of Natural History]), which has been interpreted as belonging to either a juvenile
<taxonomicName id="4C504D4D5A3FFFF9DB3EFE1A1DBEF8CC" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[578,750,1810,1831]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="781" phylum="Chordata" rank="species" species="rex">
<emphasis id="B924EADC5A3FFFF9DB3EFE1A1DBEF8CC" box="[578,750,1810,1831]" italics="true" pageId="1" pageNumber="781">Tyrannosaurus rex</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFC14B3F5A3FFFF9DB80FE191F9CF8AF" author="CARR, T. D." journalOrPublisher="Journal of Vertebrate Paleontology" pageId="1" pageNumber="781" pagination="497 - 520" part="19" refId="ref3307" refString="CARR, T. D., 1999, Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria): Journal of Vertebrate Paleontology, v. 19, p. 497 - 520." title="Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Coelurosauria)" type="book chapter" year="1999">Carr, 1999</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3FFFF9D9A5FE261E8AF8AF" author="CARR, T. D. &amp; WILLIAMSON, T. E." box="[217,474,1838,1860]" journalOrPublisher="Zoological Journal of the Linnean Society" pageId="1" pageNumber="781" pagination="479 - 523" part="142" refId="ref3341" refString="CARR, T. D., and WILLIAMSON, T. E., 2004. Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America: Zoological Journal of the Linnean Society, v. 142, p. 479 - 523." title="Diversity of late Maastrichtian Tyrannosauridae (Dinosauria: Theropoda) from western North America" type="book chapter" year="2004">Carr and Williamson, 2004</bibRefCitation>
) or a separate taxon,
<taxonomicName id="4C504D4D5A3FFFF9DBD3FE271FBEF8B4" baseAuthorityName="Gilmore" baseAuthorityYear="1946" class="Reptilia" family="Tyrannosauridae" genus="Nanotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="781" phylum="Chordata" rank="species" species="lancensis">
<emphasis id="B924EADC5A3FFFF9DBD3FE271FBEF8B4" italics="true" pageId="1" pageNumber="781">Nanotyrannus lancensis</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFC14B3F5A3FFFF9D87EFE421E93F88B" author="BAKKER, R. T. &amp; WILLIAMS, M. &amp; CURRIE, P. J." box="[258,451,1866,1888]" journalOrPublisher="Hunteria" pageId="1" pageNumber="781" pagination="1 - 30" part="1" refId="ref3157" refString="BAKKER, R. T., WILLIAMS, M., and CURRIE, P. J., 1988, Nanotyrannus, a new genus of pygmy tyrannosaur, from the latest Cretaceous of Montana, Hunteria, v. 1, no. 5, p. 1 - 30." title="Nanotyrannus, a new genus of pygmy tyrannosaur, from the latest Cretaceous of Montana" type="journal article" year="1988">Bakker et al, 1988</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3FFFF9D8A8FE421D0BF88B" author="CURRIE, P. J." box="[468,603,1866,1888]" journalOrPublisher="Canadian Journal of Earth Science" pageId="1" pageNumber="781" pagination="651 - 665" part="40" refId="ref3452" refString="CURRIE, P. J., 2003 a. Allometric growth in tyrannosaurids (Dinosauria: Theropoda) from the Upper Cretaceous of North America and Asia: Canadian Journal of Earth Science, v. 40, p. 651 - 665." title="Allometric growth in tyrannosaurids (Dinosauria: Theropoda) from the Upper Cretaceous of North America and Asia" type="book chapter" year="2003">Currie, 2003a</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3FFFF9DB10FE421DF7F88B" author="CURRIE, P. J." box="[620,679,1866,1888]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="1" pageNumber="781" pagination="191 - 226" part="48" refId="ref3497" refString="CURRIE, P. J., 2003 b, Cranial anatomy of tyrannosaurid dinosaurs from the late Cretaceous Alberta, Canada: Acta Palaeontologica Polonica, v. 48, p. 191 - 226." title="Cranial anatomy of tyrannosaurid dinosaurs from the late Cretaceous Alberta, Canada" type="journal article" year="2003">2003b</bibRefCitation>
). Research is currently ongoing into the systematic position of both specimens. Four lesions are present on the left side of the skull of
<materialsCitation id="3B383C935A3FFFF9DBE9FE8C1C7DF872" ID-GBIF-Occurrence="2813097312" box="[661,813,1923,1945]" collectionCode="BMR" pageId="1" pageNumber="781" specimenCode="BMP P2002.4.1">BMR P2002.4.1</materialsCitation>
(
<figureCitation id="136B2A4B5A3FFFF9D9DAFE971F8EF85E" box="[166,222,1951,1973]" captionStart="FIGURE 2" captionStartId="1.[159,229,1561,1578]" captionTargetBox="[162,1494,181,1542]" captionTargetId="figure@1.[162,1495,181,1542]" captionTargetPageId="1" captionText="FIGURE 2—Lesions present on the face of BMR P2002.4.1. A) An index drawing of the lesions (1–4). B) The four lesions on the left maxilla and nasal of BMR P2002.4.1. C) A dorsal view of the anterior nasal, showing asymmetry as a result of the puncture on the left side. D) The first puncture (arrow = lesion 1) located on the articular surface of the anterior nasal and left maxilla. E–F) The three lesions on the left maxilla of BMR P2002.4.1 with a close-up of lesion 4 (F). Scale bars: A–B = 10 cm; C–E = 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3942889" httpUri="https://zenodo.org/record/3942889/files/figure.png" pageId="1" pageNumber="781">Fig. 2</figureCitation>
A–B). The lesions are lenticular shaped, rostrocaudally long, and fully penetrate the left nasal and maxilla, which is 6 mm thick near the
</paragraph>
</subSubSection>
<caption id="DF2F66465A3EFFF8DA4EFEA21CEDF839" ID-DOI="http://doi.org/10.5281/zenodo.3942887" ID-Zenodo-Dep="3942887" httpUri="https://zenodo.org/record/3942887/files/figure.png" pageId="0" pageNumber="780" startId="0.[818,888,1962,1979]" targetBox="[821,1469,1394,1943]" targetPageId="0">
<paragraph id="8BEF36CE5A3EFFF8DA4EFEA21CEDF839" blockId="0.[818,1471,1962,2003]" pageId="0" pageNumber="780">
<emphasis id="B924EADC5A3EFFF8DA4EFEA21CCDF850" bold="true" box="[818,925,1962,1979]" pageId="0" pageNumber="780">FIGURE 1—</emphasis>
Map of Carter County, MT and the location of the
<materialsCitation id="3B383C935A3EFFF8DCE8FEA21C2CF838" ID-GBIF-Occurrence="2813097310" collectionCode="BMR" pageId="0" pageNumber="780" specimenCode="BMP P2002.4.1">BMR P2002.4.1</materialsCitation>
quarry.
</paragraph>
</caption>
<footnote id="E84B2AC05A3EFFF8D9F9FECB1C53F7ED" pageId="0" pageNumber="780">
<paragraph id="8BEF36CE5A3EFFF8D9F9FECB1E15F83F" blockId="0.[133,325,1987,2004]" box="[133,325,1987,2004]" pageId="0" pageNumber="780">* Corresponding author.</paragraph>
<paragraph id="8BEF36CE5A3EFFF8D9F9FEFA1C53F7ED" blockId="0.[133,771,2033,2054]" box="[133,771,2033,2054]" pageId="0" pageNumber="780">
<emphasis id="B924EADC5A3EFFF8D9F9FEFA1FBEF7ED" bold="true" box="[133,238,2034,2054]" pageId="0" pageNumber="780">Copyright</emphasis>
<emphasis id="B924EADC5A3EFFF8D98AFEFE1E53F7E8" bold="true" box="[246,259,2038,2051]" italics="true" pageId="0" pageNumber="780">©</emphasis>
2009, SEPM (Society for Sedimentary Geology)
</paragraph>
</footnote>
<footnote id="E84B2AC05A3EFFF8DA4EFEF91B3BF7ED" box="[818,1131,2033,2054]" pageId="0" pageNumber="780">
<paragraph id="8BEF36CE5A3EFFF8DA4EFEF91B3BF7ED" blockId="0.[818,1131,2033,2054]" box="[818,1131,2033,2054]" pageId="0" pageNumber="780">0883-1351/09/002#-0780/$3.00</paragraph>
</footnote>
<caption id="DF2F66465A3FFFF9D9E3FF111AF7F9B1" ID-DOI="http://doi.org/10.5281/zenodo.3942889" ID-Zenodo-Dep="3942889" httpUri="https://zenodo.org/record/3942889/files/figure.png" pageId="1" pageNumber="781" startId="1.[159,229,1561,1578]" targetBox="[162,1494,181,1542]" targetPageId="1">
<paragraph id="8BEF36CE5A3FFFF9D9E3FF111AF7F9B1" blockId="1.[159,1499,1560,1627]" pageId="1" pageNumber="781">
<emphasis id="B924EADC5A3FFFF9D9E3FF111E59F9C1" bold="true" box="[159,265,1561,1578]" pageId="1" pageNumber="781">FIGURE 2—</emphasis>
Lesions present on the face of
<materialsCitation id="3B383C935A3FFFF9D893FF121D3CF9C0" ID-GBIF-Occurrence="2813097327" box="[495,620,1562,1579]" collectionCode="BMR" pageId="1" pageNumber="781" specimenCode="BMP P2002.4.1">BMR P2002.4.1</materialsCitation>
. A) An index drawing of the lesions (1–4). B) The four lesions on the left maxilla and nasal of
<materialsCitation id="3B383C935A3FFFF9DC4EFF121AFCF9C0" ID-GBIF-Occurrence="2813097319" box="[1330,1452,1562,1579]" collectionCode="BMR" pageId="1" pageNumber="781" specimenCode="BMP P2002.4.1">BMR P2002.4.1</materialsCitation>
. C) A dorsal view of the anterior nasal, showing asymmetry as a result of the puncture on the left side. D) The first puncture (arrow = lesion 1) located on the articular surface of the anterior nasal and left maxilla. E–F) The three lesions on the left maxilla of
<materialsCitation id="3B383C935A3FFFF9DB8AFF421C22F9B1" ID-GBIF-Occurrence="2813097302" box="[758,882,1609,1626]" collectionCode="BMR" pageId="1" pageNumber="781" specimenCode="BMP P2002.4.1">BMR P2002.4.1</materialsCitation>
with a close-up of lesion 4 (F). Scale bars: A–B = 10 cm; C–E = 5 cm.
</paragraph>
</caption>
<subSubSection id="C34A65455A3FFFFADA31FF8C1C43FC27" lastPageId="2" lastPageNumber="782" pageId="1" pageNumber="781" type="description">
<paragraph id="8BEF36CE5A3FFFF9DA31FF8C1C2BF905" blockId="1.[845,1500,1667,2002]" pageId="1" pageNumber="781">puncture marks. Several of the preserved lesions display bone repair (cicatrisation), indicating that the animal survived these injuries to its skull. There is no osteological evidence for infections at these puncture sites.</paragraph>
<paragraph id="8BEF36CE5A3FFFFADA1EFFFD1EA3FEB1" blockId="1.[845,1500,1667,2002]" lastBlockId="2.[133,788,182,973]" lastPageId="2" lastPageNumber="782" pageId="1" pageNumber="781">
The anteriormost puncture is located ~15 mm posterior to the left bony naris, at the nasomaxillary suture (
<figureCitation id="136B2A4B5A3FFFF9DDBCFE1A1BA7F8C3" box="[1216,1271,1810,1832]" captionStart="FIGURE 2" captionStartId="1.[159,229,1561,1578]" captionTargetBox="[162,1494,181,1542]" captionTargetId="figure@1.[162,1495,181,1542]" captionTargetPageId="1" captionText="FIGURE 2—Lesions present on the face of BMR P2002.4.1. A) An index drawing of the lesions (1–4). B) The four lesions on the left maxilla and nasal of BMR P2002.4.1. C) A dorsal view of the anterior nasal, showing asymmetry as a result of the puncture on the left side. D) The first puncture (arrow = lesion 1) located on the articular surface of the anterior nasal and left maxilla. E–F) The three lesions on the left maxilla of BMR P2002.4.1 with a close-up of lesion 4 (F). Scale bars: A–B = 10 cm; C–E = 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3942889" httpUri="https://zenodo.org/record/3942889/files/figure.png" pageId="1" pageNumber="781">Fig. 2</figureCitation>
C–D). This puncture has a maximum length of 15 mm and a maximum height of 5 mm. The edges are fractured with partial bone repair. The left maxillary process of the nasal is absent, apparently excised during the injury (
<figureCitation id="136B2A4B5A3FFFF9DCFBFE6F1A81F896" box="[1415,1489,1895,1917]" captionStart="FIGURE 2" captionStartId="1.[159,229,1561,1578]" captionTargetBox="[162,1494,181,1542]" captionTargetId="figure@1.[162,1495,181,1542]" captionTargetPageId="1" captionText="FIGURE 2—Lesions present on the face of BMR P2002.4.1. A) An index drawing of the lesions (1–4). B) The four lesions on the left maxilla and nasal of BMR P2002.4.1. C) A dorsal view of the anterior nasal, showing asymmetry as a result of the puncture on the left side. D) The first puncture (arrow = lesion 1) located on the articular surface of the anterior nasal and left maxilla. E–F) The three lesions on the left maxilla of BMR P2002.4.1 with a close-up of lesion 4 (F). Scale bars: A–B = 10 cm; C–E = 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3942889" httpUri="https://zenodo.org/record/3942889/files/figure.png" pageId="1" pageNumber="781">Fig. 2C</figureCitation>
). Three additional lesions are preserved on the left maxilla (
<figureCitation id="136B2A4B5A3FFFF9DC13FE8C1AF6F871" box="[1391,1446,1924,1946]" captionStart="FIGURE 2" captionStartId="1.[159,229,1561,1578]" captionTargetBox="[162,1494,181,1542]" captionTargetId="figure@1.[162,1495,181,1542]" captionTargetPageId="1" captionText="FIGURE 2—Lesions present on the face of BMR P2002.4.1. A) An index drawing of the lesions (1–4). B) The four lesions on the left maxilla and nasal of BMR P2002.4.1. C) A dorsal view of the anterior nasal, showing asymmetry as a result of the puncture on the left side. D) The first puncture (arrow = lesion 1) located on the articular surface of the anterior nasal and left maxilla. E–F) The three lesions on the left maxilla of BMR P2002.4.1 with a close-up of lesion 4 (F). Scale bars: A–B = 10 cm; C–E = 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3942889" httpUri="https://zenodo.org/record/3942889/files/figure.png" pageId="1" pageNumber="781">Fig. 2</figureCitation>
E–F). The rostralmost puncture is present directly above the maxillary fenestra and is 10 mm in length and 9 mm in maximum height. The edges are fractured without evidence of cicatrisation. The second puncture is positioned 8 mm posterior to the first and is 18 mm in length and 4 mm at maximum height. The edges of the puncture consist of smooth cortical bone. The third puncture is 12 mm posterior to the second hole and is 19 mm in length and 5 mm at maximum height; its edges are also of smooth cortical bone.
</paragraph>
<paragraph id="8BEF36CE5A3CFFFAD9E6F86A1D70FCB1" blockId="2.[133,788,182,973]" pageId="2" pageNumber="782">
In addition to the lesions, the reconstructed skull of
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demonstrates abnormal asymmetry, with the distal portion of the muzzle curved laterally to the left side (
<figureCitation id="136B2A4B5A3CFFFADB7EF8921D6BFE5B" box="[514,571,410,432]" captionStart="FIGURE 3" captionStartId="2.[818,888,1512,1529]" captionTargetBox="[853,1437,181,1492]" captionTargetId="figure@2.[853,1437,181,1493]" captionTargetPageId="2" captionText="FIGURE 3—Dorsal view of the skull of BMR P2002.4.1. The central line illustrates the curved nasal and resulting deformation of the skull due to the injury. Scale bar = 10 cm." figureDoi="http://doi.org/10.5281/zenodo.3942891" httpUri="https://zenodo.org/record/3942891/files/figure.png" pageId="2" pageNumber="782">Fig. 3</figureCitation>
). The curvature is not considered severe enough to have affected the animal’s ability to bite or capture prey. Post-depositional deformation is a common cause of bone distortion in fossil specimens, but the asymmetry of this specimen is considered primary—that is, caused by the injury, with the distortion propagated by subsequent growth—rather than a taphonomic feature. When discovered, the skull of
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was disarticulated with its nasal lying dorsoventrally and both maxillae lying on their lateral sides. The nasal and maxillae fit well along their articular surfaces despite the disarticulation, and there is little evidence of postdepositional deformation in any of the other preserved cranial elements. This strongly suggests that the lateral curvature of the muzzle is the result of facial injury rather than diagenetic deformation. Furthermore, there is no sedimentological evidence of post-depositional compaction; the clay balls in the conglomerate that encase the bones are spherical, not compressed to oriented lenticular clasts.
</paragraph>
<paragraph id="8BEF36CE5A3CFFFAD9E6FA691C43FC27" blockId="2.[133,788,182,973]" pageId="2" pageNumber="782">A major pathology with evidence of active infection at the time of death is present on the first phalanx on the second pedal digit of the left pes (C.P. Vittore, personal communication, 2009). There is no evidence to suggest, however, that this pathology is related to the facial injuries.</paragraph>
</subSubSection>
<subSubSection id="C34A65455A3CFFFBD8FBFAF71AF4FC10" lastPageId="3" lastPageNumber="783" pageId="2" pageNumber="782" type="discussion">
<paragraph id="8BEF36CE5A3CFFFAD8FBFAF71D40FBFE" blockId="2.[391,528,1023,1045]" box="[391,528,1023,1045]" pageId="2" pageNumber="782">
<heading id="D0A781A25A3CFFFAD8FBFAF71D40FBFE" allCaps="true" bold="true" box="[391,528,1023,1045]" centered="true" fontSize="9" level="1" pageId="2" pageNumber="782" reason="0">DISCUSSION</heading>
</paragraph>
<paragraph id="8BEF36CE5A3CFFFAD9E6FD381C56FAC2" blockId="2.[133,787,1072,2005]" pageId="2" pageNumber="782">
There is little doubt that the loss of the left lateral process of the nasal and the lesions to the left maxilla of
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occurred at the same time. This is indicated by the uniform shape and size of the lesions and the presence of healed bone tissue along their edges. In addition, the distribution of the lesions along a gently arcuate line and the similar degree of healing supports the hypothesis that these injuries were inflicted simultaneously, probably from a bite to the face. The lateral curvature of the muzzle described above likely resulted from differential growth of the face following the loss of the left lateral nasal process.
</paragraph>
<paragraph id="8BEF36CE5A3CFFFAD9E6FC391E15F91A" blockId="2.[133,787,1072,2005]" pageId="2" pageNumber="782">
The only fossil vertebrates preserved in the Hell Creek Formation that are large enough to puncture the skull of
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are crocodilians and theropod dinosaurs. The lenticular-shaped lesions are inconsistent with the circular cross sections of crocodilian teeth, but they are consistent with the lenticular cross sections of juvenile theropod teeth. The maxillary teeth of
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are laterally compressed; a characteristic of juvenile tyrannosaurids in general. The size, shape, and spacing of the maxillary teeth of
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correspond well with the size, shape and spacing of the lesions, indicating that this animal was bitten by another juvenile tyrannosaurid of its size. The location and spacing of the facial injuries indicate that the bite was inflicted by the grazing anterior maxillary teeth of an adversary, which was in front of and facing
<materialsCitation id="3B383C935A3CFFFADB63FF8E1DE3F977" ID-GBIF-Occurrence="2813097325" box="[543,691,1670,1692]" collectionCode="BMR" pageId="2" pageNumber="782" specimenCode="BMP P2002.4.1">BMR P2002.4.1</materialsCitation>
(
<figureCitation id="136B2A4B5A3CFFFADBBCFF8E1DA7F977" box="[704,759,1670,1692]" captionStart="FIGURE 4" captionStartId="3.[159,229,783,800]" captionTargetBox="[165,810,181,761]" captionTargetId="figure@3.[162,810,181,764]" captionTargetPageId="3" captionText="FIGURE 4—Reconstruction of likely puncture origin of BMR P2002.4.1. A) Skeletal reconstruction of intraspecific conflict between BMR P2002.4.1 (right) and attacking individual (left). B) Life reconstruction of the intraspecific attack." figureDoi="http://doi.org/10.5281/zenodo.3942895" httpUri="https://zenodo.org/record/3942895/files/figure.png" pageId="2" pageNumber="782">Fig. 4</figureCitation>
A– B). The smooth cortical bone covering the edges of the lesions indicates that the bite was not fatal and healing was well underway or complete at the time of death.
</paragraph>
<caption id="DF2F66465A3CFFFADA4EFCE01C37F9C3" ID-DOI="http://doi.org/10.5281/zenodo.3942891" ID-Zenodo-Dep="3942891" httpUri="https://zenodo.org/record/3942891/files/figure.png" pageId="2" pageNumber="782" startId="2.[818,888,1512,1529]" targetBox="[853,1437,181,1492]" targetPageId="2">
<paragraph id="8BEF36CE5A3CFFFADA4EFCE01C37F9C3" blockId="2.[818,1472,1512,1577]" pageId="2" pageNumber="782">
<emphasis id="B924EADC5A3CFFFADA4EFCE01CCDFA12" bold="true" box="[818,925,1512,1529]" pageId="2" pageNumber="782">FIGURE 3—</emphasis>
Dorsal view of the skull of
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. The central line illustrates the curved nasal and resulting deformation of the skull due to the injury. Scale bar = 10 cm.
</paragraph>
</caption>
<paragraph id="8BEF36CE5A3CFFFAD9E6FFF01E0DF83E" blockId="2.[133,787,1072,2005]" pageId="2" pageNumber="782">
The lesions in
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provide insight into the social behavior of juvenile tyrannosaurids. Evidence for face biting is not uncommon in theropod dinosaurs (
<bibRefCitation id="EFC14B3F5A3CFFFAD8A0FE391D99F8AC" author="TANKE, D. H. &amp; CURRIE, P. J." box="[476,713,1841,1863]" journalOrPublisher="Gaia" pageId="2" pageNumber="782" pagination="167 - 184" part="15" refId="ref4344" refString="TANKE, D. H., and CURRIE, P. J., 1998, Head-biting behavior in theropod dinosaurs: Paleopathological evidence: Gaia, v. 15, p. 167 - 184." title="Head-biting behavior in theropod dinosaurs: Paleopathological evidence" type="book chapter" year="1998">Tanke and Currie, 1998</bibRefCitation>
). While intraspecific predation and cannibalism have been previously suggested in tyrannosaurids (
<bibRefCitation id="EFC14B3F5A3CFFFAD842FE611EECF894" author="TRACEY, C. R." box="[318,444,1897,1919]" journalOrPublisher="American Naturalist" pageId="2" pageNumber="782" pagination="1105 - 1106" part="110" refId="ref4382" refString="TRACEY, C. R., 1976, Tyrannosaurs: Evidence for endothermy?: American Naturalist, v. 110, p. 1105 - 1106." title="Tyrannosaurs: Evidence for endothermy?" type="journal article" year="1976">Tracey, 1976</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3CFFFAD8B0FE621D15F894" author="NAEYE, R." box="[460,581,1897,1919]" journalOrPublisher="Earth" pageId="2" pageNumber="782" pagination="12" part="5" refId="ref4321" refString="NAEYE, R., 1996, T. rex gets tougher: Earth, v. 5, p. 12." title="T. rex gets tougher" type="journal article" year="1996">Naeye, 1996</bibRefCitation>
), true predation and cannibalism usually involve a much larger individual attacking a smaller one (
<bibRefCitation id="EFC14B3F5A3CFFFAD982FEAA1E8FF853" author="TANKE, D. H. &amp; CURRIE, P. J." box="[254,479,1954,1976]" journalOrPublisher="Gaia" pageId="2" pageNumber="782" pagination="167 - 184" part="15" refId="ref4344" refString="TANKE, D. H., and CURRIE, P. J., 1998, Head-biting behavior in theropod dinosaurs: Paleopathological evidence: Gaia, v. 15, p. 167 - 184." title="Head-biting behavior in theropod dinosaurs: Paleopathological evidence" type="book chapter" year="1998">Tanke and Currie, 1998</bibRefCitation>
) and are not likely scenarios for the bite described here.
</paragraph>
<paragraph id="8BEF36CE5A3CFFFBDA34FF591D12FC39" blockId="2.[818,1473,1617,2005]" lastBlockId="3.[159,814,899,2004]" lastPageId="3" lastPageNumber="783" pageId="2" pageNumber="782">
While social behavior cannot be directly observed in or unequivocally inferred from the fossil record, it can be inferred from evidence of eating and from injuries (
<bibRefCitation id="EFC14B3F5A3CFFFADD5FFF811A57F974" author="TANKE, D. H. &amp; CURRIE, P. J." box="[1059,1287,1673,1695]" journalOrPublisher="Gaia" pageId="2" pageNumber="782" pagination="167 - 184" part="15" refId="ref4344" refString="TANKE, D. H., and CURRIE, P. J., 1998, Head-biting behavior in theropod dinosaurs: Paleopathological evidence: Gaia, v. 15, p. 167 - 184." title="Head-biting behavior in theropod dinosaurs: Paleopathological evidence" type="book chapter" year="1998">Tanke and Currie, 1998</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3CFFFADC6FFF811AEDF974" author="AVILLA, L. S. &amp; FERNANDES, R. &amp; RAMOS, D. F. B." box="[1299,1469,1673,1695]" journalOrPublisher="Journal of Vertebrate Paleonotology" pageId="2" pageNumber="782" pagination="971 - 973" part="24" refId="ref3100" refString="AVILLA, L. S., FERNANDES, R., and RAMOS, D. F. B., 2004, Bite marks on a crocodylomorph from the Upper Cretaceous of Brazil: Evidence of social behavior?: Journal of Vertebrate Paleonotology, v. 24, p. 971 - 973." title="Bite marks on a crocodylomorph from the Upper Cretaceous of Brazil: Evidence of social behavior?" type="journal article" year="2004">Avilla et al., 2004</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3CFFFADA4EFFAE1CEBF950" author="KATSURA, Y." box="[818,955,1701,1723]" journalOrPublisher="Historical Biology" pageId="2" pageNumber="782" pagination="93 - 97" part="16" refId="ref4203" refString="KATSURA, Y., 2004, Paleopathology of Toyotamaphimeia machikanensis (Diapsida, Crocodylia) from the Middle Pleistocene of Central Japan: Historical Biology, v. 16, p. 93 - 97." title="Paleopathology of Toyotamaphimeia machikanensis (Diapsida, Crocodylia) from the Middle Pleistocene of Central Japan" type="book chapter" year="2004">Katsura, 2004</bibRefCitation>
). The ability of tyrannosaurids to puncture and crush bone is well documented (
<bibRefCitation id="EFC14B3F5A3CFFFADD44FFC91BFCF93C" author="ABLER, W. L." box="[1080,1196,1729,1751]" journalOrPublisher="Paleobiology" pageId="2" pageNumber="782" pagination="161 - 183" part="18" refId="ref3065" refString="ABLER, W. L., 1992, The serrated teeth of tyrannosaurid dinosaurs, and biting structures in other animals: Paleobiology, v. 18, p. 161 - 183." title="The serrated teeth of tyrannosaurid dinosaurs, and biting structures in other animals" type="journal article" year="1992">Abler, 1992</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3CFFFADDC7FFC91AECF93C" author="ERICKSON, G. M. &amp; OLSON, K. H." box="[1211,1468,1729,1751]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="2" pageNumber="782" pagination="175 - 178" part="16" refId="ref3631" refString="ERICKSON, G. M., and OLSON, K. H., 1996, Bite marks attributable to Tyrannosaurus rex: Preliminary descriptions and implications: Journal of Vertebrate Paleontology, v. 16, p. 175 - 178." title="Bite marks attributable to Tyrannosaurus rex: Preliminary descriptions and implications" type="book chapter" year="1996">Erickson and Olson, 1996</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3CFFFADA4EFFD51CAFF918" author="ERICKSON, G. M. &amp; VAN KIRK, S. D. &amp; SU, J. &amp; LEVENSON, M. E. &amp; CALER, W. E. &amp; CARTER, D. R." box="[818,1023,1757,1779]" journalOrPublisher="Nature" pageId="2" pageNumber="782" pagination="706 - 708" part="382" refId="ref3675" refString="ERICKSON, G. M., VAN KIRK, S. D., SU, J., LEVENSON, M. E., CALER, W. E., and CARTER, D. R., 1996, Bite-force estimation for Tyrannosaurus rex from tooth-marked bones: Nature, v. 382, p. 706 - 708." title="Bite-force estimation for Tyrannosaurus rex from tooth-marked bones" type="book chapter" year="1996">Erickson et al., 1996</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3CFFFADD71FFD61A53F918" author="HURUM, J. H. &amp; CURRIE, P. J." box="[1037,1283,1757,1779]" journalOrPublisher="Journal of Vertebrate Paleontology" pageId="2" pageNumber="782" pagination="619 - 621" part="20" refId="ref4033" refString="HURUM, J. H., and CURRIE, P. J., 2000, The crushing bite of tyrannosaurids: Journal of Vertebrate Paleontology, v. 20, p. 619 - 621." title="The crushing bite of tyrannosaurids" type="book chapter" year="2000">Hurum and Currie, 2000</bibRefCitation>
;
<bibRefCitation id="EFC14B3F5A3CFFFADC6DFFD61ADAF918" author="MEERS, M. B." box="[1297,1418,1757,1779]" journalOrPublisher="Historical Biology" pageId="2" pageNumber="782" pagination="1 - 12" part="16" refId="ref4280" 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, v. 16, p. 1 - 12." title="Maximum bite force and prey size of Tyrannosaurus rex and their relationships to the inference of feeding behavior" type="book chapter" year="2002">Meers, 2002</bibRefCitation>
), and evidence of feeding is relatively common at fossil vertebrate sites (
<bibRefCitation id="EFC14B3F5A3CFFFADCFAFFF21CE0F8C7" author="CHURE, D. J. &amp; FIORILLO, A. R. &amp; JACOBSEN, A. R." journalOrPublisher="Gaia" pageId="2" pageNumber="782" pagination="227 - 232" part="15" refId="ref3390" refString="CHURE, D. J., FIORILLO, A. R., and JACOBSEN, A. R., 1998, Prey bone utilization by predatory dinosaurs in the Late Jurassic of North America, with comments on prey bone use by dinosaurs throughout the Mesozoic: Gaia, v. 15, p. 227 - 232." title="Prey bone utilization by predatory dinosaurs in the Late Jurassic of North America, with comments on prey bone use by dinosaurs throughout the Mesozoic" type="journal article" year="1998">Chure et al., 1998</bibRefCitation>
). Behavior can also be inferred using the Extant Phylogenetic Bracket (EPB) method (
<bibRefCitation id="EFC14B3F5A3CFFFADDE5FE3A1A4EF8A3" author="WITMER, L. M." box="[1177,1310,1842,1864]" editor="Thomason, J." journalOrPublisher="Cambridge University Press, Cambridge, UK" pageId="2" pageNumber="782" pagination="1933" refId="ref4517" refString="WITMER, L. M., 1995, The extant phylogenetic bracket and the importance of reconstructing soft tissues in fossils, in Thomason, J., ed., Functional Morphology in Vertebrate Paleontology: Cambridge University Press, Cambridge, UK, p. 1933." title="The extant phylogenetic bracket and the importance of reconstructing soft tissues in fossils" type="book chapter" volumeTitle="Functional Morphology in Vertebrate Paleontology" year="1995">Witmer, 1995</bibRefCitation>
). This method is generally used for soft-tissue reconstruction and behavioral inference by comparing the osteological correlates of living members of a group to infer a characteristic in an extinct member. The EPB has previously been used to interpret nesting and parental behavior in non-avian theropods (
<bibRefCitation id="EFC14B3F5A3CFFFADAE7FEB71B3CF83E" author="VARRICCHIO, D. J. &amp; MOORE, J. R. &amp; ERICKSON, G. M. &amp; NORELL, M. A. &amp; JACKSON, F. D. &amp; BORKOWSKI, J. J." box="[923,1132,1983,2005]" journalOrPublisher="Science" pageId="2" pageNumber="782" pagination="1826 - 1828" part="322" refId="ref4411" refString="VARRICCHIO, D. J., MOORE, J. R., ERICKSON, G. M., NORELL, M. A., JACKSON, F. D., and BORKOWSKI, J. J., 2008, Avian parental care had dinosaur origin: Science, v. 322, p. 1826 - 1828." title="Avian parental care had dinosaur origin" type="book chapter" year="2008">Varricchio et al., 2008</bibRefCitation>
), using extant crocodilians and birds as living members of Archosauria. A similar model was used for the purposes of this study, where intraspecific aggressive behaviors were investigated in extant crocodilians and birds.
</paragraph>
<caption id="DF2F66465A3DFFFBDA31FB7F1CDEFD4B" ID-DOI="http://doi.org/10.5281/zenodo.3942893" ID-Zenodo-Dep="3942893" httpUri="https://zenodo.org/record/3942893/files/figure.png" pageId="3" pageNumber="783" startId="3.[845,915,631,648]" targetBox="[850,1494,183,612]" targetPageId="3">
<paragraph id="8BEF36CE5A3DFFFBDA31FB7F1CDEFD4B" blockId="3.[845,1499,631,672]" pageId="3" pageNumber="783">
<emphasis id="B924EADC5A3DFFFBDA31FB7F1CE8FD63" bold="true" box="[845,952,631,648]" pageId="3" pageNumber="783">FIGURE 5—</emphasis>
Skull of
<taxonomicName id="4C504D4D5A3DFFFBDA84FB701BE3FD63" baseAuthorityName="Daudin" baseAuthorityYear="1802" box="[1016,1203,632,649]" class="Reptilia" family="Alligatoridae" genus="Alligator" kingdom="Animalia" order="Crocodylia" pageId="3" pageNumber="783" phylum="Chordata" rank="species" species="mississippiensis">
<emphasis id="B924EADC5A3DFFFBDA84FB701BE3FD63" box="[1016,1203,632,649]" italics="true" pageId="3" pageNumber="783">Alligator mississippiensis</emphasis>
</taxonomicName>
with facial injuries (arrows). Scale bar = 5 cm.
</paragraph>
</caption>
<caption id="DF2F66465A3DFFFBD9E3FA071DF0FCBB" ID-DOI="http://doi.org/10.5281/zenodo.3942895" ID-Zenodo-Dep="3942895" httpUri="https://zenodo.org/record/3942895/files/figure.png" pageId="3" pageNumber="783" startId="3.[159,229,783,800]" targetBox="[165,810,181,761]" targetPageId="3">
<paragraph id="8BEF36CE5A3DFFFBD9E3FA071DF0FCBB" blockId="3.[159,813,782,848]" pageId="3" pageNumber="783">
<emphasis id="B924EADC5A3DFFFBD9E3FA071E59FCCB" bold="true" box="[159,265,783,800]" pageId="3" pageNumber="783">FIGURE 4—</emphasis>
Reconstruction of likely puncture origin of
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. A) Skeletal reconstruction of intraspecific conflict between
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(right) and attacking individual (left). B) Life reconstruction of the intraspecific attack.
</paragraph>
</caption>
<paragraph id="8BEF36CE5A3DFFFBD9C9FAD21E0FFA71" blockId="3.[159,814,899,2004]" pageId="3" pageNumber="783">
<heading id="D0A781A25A3DFFFBD9C9FAD21C7DFC04" bold="true" box="[181,813,985,1007]" fontSize="9" level="3" pageId="3" pageNumber="783" reason="0">Puncture marks along the maxilla and nasal, and skewed jaws</heading>
similar to the injuries on
<materialsCitation id="3B383C935A3DFFFBD8D9FAFE1D10FBE7" ID-GBIF-Occurrence="2813097323" box="[421,576,1014,1036]" collectionCode="BMR" pageId="3" pageNumber="783" specimenCode="BMP P2002.4.1">BMR P2002.4.1</materialsCitation>
are commonly seen on extant crocodilians as the result of non-fatal intraspecific aggression (
<bibRefCitation id="EFC14B3F5A3DFFFBD9DAFD261E70FBAF" author="KALIN, J. A." box="[166,288,1070,1092]" journalOrPublisher="Zeitschrift fur Morphologie und Oekologie de Tiere" pageId="3" pageNumber="783" pagination="327 - 347" part="32" refId="ref4171" refString="KALIN, J. A., 1936, Uber Skeletanomalien der Crocodilien: Zeitschrift fur Morphologie und Oekologie de Tiere, v. 32, p. 327 - 347." title="Uber Skeletanomalien der Crocodilien" type="book chapter" year="1936">Kalin, 1936</bibRefCitation>
), especially on the saltwater crocodile,
<taxonomicName id="4C504D4D5A3DFFFBDBBAFD271FB4FB8A" authority="Schneider, 1801" baseAuthorityName="Schneider" baseAuthorityYear="1801" class="Reptilia" family="Crocodylidae" genus="Crocodylus" kingdom="Animalia" order="Crocodylia" pageId="3" pageNumber="783" phylum="Chordata" rank="species" species="porosus">
<emphasis id="B924EADC5A3DFFFBDBBAFD271FB4FB8A" italics="true" pageId="3" pageNumber="783">Crocodylus porosus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFC14B3F5A3DFFFBD984FD431EA1FB8A" author="WEBB, G. J. W. &amp; MESSEL, H." box="[248,497,1099,1121]" journalOrPublisher="Australian Wildlife Research" pageId="3" pageNumber="783" pagination="311 - 319" part="4" refId="ref4475" refString="WEBB, G. J. W., and MESSEL, H., 1977, Abnormalities and injuries in the estuarine crocodile, Crocodylus porosus: Australian Wildlife Research, v. 4, p. 311 - 319." title="Abnormalities and injuries in the estuarine crocodile, Crocodylus porosus" type="journal article" year="1977">Webb and Messel, 1977</bibRefCitation>
). Extant crocodilians display a variety of complex social behaviors that often result in intraspecific aggression due to conflicts in dominance hierarchies, territoriality, or courtship (
<bibRefCitation id="EFC14B3F5A3DFFFBD87BFDA81E28FB5D" author="LANG, J. W." box="[263,376,1184,1206]" editor="Ross, C. A." journalOrPublisher="Facts on File, New York" pageId="3" pageNumber="783" pagination="102 - 117" refId="ref4240" refString="LANG, J. W., 1989, Social behavior, in Ross, C. A., ed., Crocodiles and Alligators: Facts on File, New York, p. 102 - 117." title="Social behavior" type="book chapter" volumeTitle="Crocodiles and Alligators" year="1989">Lang, 1989</bibRefCitation>
). The injury described here lacks evidence of the shaking or rolling behaviors that extant crocodilians commonly display with prey capture. The lesions present on
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are consistent with injuries in extant crocodilians exhibiting intraspecific aggression (i.e., head-slapping) rather than prey capture (rolling, shaking). Because the puncture wounds present on
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are similar to those found on extant crocodilians (
<figureCitation id="136B2A4B5A3DFFFBDB08FC431DFDFA8A" box="[628,685,1355,1377]" captionStart="FIGURE 5" captionStartId="3.[845,915,631,648]" captionTargetBox="[850,1494,183,612]" captionTargetId="figure@3.[848,1495,181,613]" captionTargetPageId="3" captionText="FIGURE 5—Skull of Alligator mississippiensis with facial injuries (arrows). Scale bar = 5 cm." figureDoi="http://doi.org/10.5281/zenodo.3942893" httpUri="https://zenodo.org/record/3942893/files/figure.png" pageId="3" pageNumber="783">Fig. 5</figureCitation>
), we consider it likely that they both resulted from roughly comparable intraspecific aggressive behavior.
</paragraph>
<paragraph id="8BEF36CE5A3DFFFBD9C9FCA91C4DF9C3" blockId="3.[159,814,899,2004]" pageId="3" pageNumber="783">
Erickson at al. (
<bibRefCitation id="EFC14B3F5A3DFFFBD83BFCA91E24FA5C" author="ERICKSON, G. M. &amp; CURRIE, P. J. &amp; INOUYE, B. D. &amp; WINN, A. A." box="[327,372,1441,1463]" journalOrPublisher="Science" pageId="3" pageNumber="783" pagination="213 - 217" part="313" refId="ref3576" refString="ERICKSON, G. M., CURRIE, P. J., INOUYE, B. D., and WINN, A. A., 2006, Tyrannosaur life tables: An example of nonavian dinosaur population biology: Science, v. 313, p. 213 - 217." title="Tyrannosaur life tables: An example of nonavian dinosaur population biology" type="journal article" year="2006">2006</bibRefCitation>
) indicate that
<taxonomicName id="4C504D4D5A3DFFFBD885FCAA1DF2FA5C" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[505,674,1442,1463]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="783" phylum="Chordata" rank="species" species="rex">
<emphasis id="B924EADC5A3DFFFBD885FCAA1DF2FA5C" box="[505,674,1442,1463]" italics="true" pageId="3" pageNumber="783">Tyrannosaurus rex</emphasis>
</taxonomicName>
reached sexual maturity at 14 years of age.
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has been histologically determined to be 12 years old at the time of death (
<bibRefCitation id="EFC14B3F5A3DFFFBDBE7FCD11F9EF9E7" author="ERICKSON, G. M. &amp; CURRIE, P. J. &amp; INOUYE, B. D. &amp; WINN, A. A." journalOrPublisher="Science" pageId="3" pageNumber="783" pagination="213 - 217" part="313" refId="ref3576" refString="ERICKSON, G. M., CURRIE, P. J., INOUYE, B. D., and WINN, A. A., 2006, Tyrannosaur life tables: An example of nonavian dinosaur population biology: Science, v. 313, p. 213 - 217." title="Tyrannosaur life tables: An example of nonavian dinosaur population biology" type="journal article" year="2006">Erickson et al., 2006</bibRefCitation>
). Because
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was probably not sexually mature, we suggest that the injury did not likely result from courtship behavior.
</paragraph>
<paragraph id="8BEF36CE5A3DFFFBD9C9FF271C53F895" blockId="3.[159,814,899,2004]" pageId="3" pageNumber="783">
Aggression in dominance hierarchies and territoriality between extant crocodilians commonly result in conflict and injury (
<bibRefCitation id="EFC14B3F5A3DFFFBDBB0FF441E49F995" author="WEBB, G. J. W. &amp; MESSEL, H." journalOrPublisher="Australian Wildlife Research" pageId="3" pageNumber="783" pagination="311 - 319" part="4" refId="ref4475" refString="WEBB, G. J. W., and MESSEL, H., 1977, Abnormalities and injuries in the estuarine crocodile, Crocodylus porosus: Australian Wildlife Research, v. 4, p. 311 - 319." title="Abnormalities and injuries in the estuarine crocodile, Crocodylus porosus" type="journal article" year="1977">Webb and Messel, 1977</bibRefCitation>
). Territoriality is most common in small groups, however, whereas dominance hierarchies are typically seen in large communities and involve more massive individuals (
<bibRefCitation id="EFC14B3F5A3DFFFBDB3FFFA91DEBF95C" author="LANG, J. W." box="[579,699,1697,1719]" editor="Ross, C. A." journalOrPublisher="Facts on File, New York" pageId="3" pageNumber="783" pagination="102 - 117" refId="ref4240" refString="LANG, J. W., 1989, Social behavior, in Ross, C. A., ed., Crocodiles and Alligators: Facts on File, New York, p. 102 - 117." title="Social behavior" type="book chapter" volumeTitle="Crocodiles and Alligators" year="1989">Lang, 1989</bibRefCitation>
). Saltwater crocodiles are territorial and aggressively display and vocalize during confrontations, especially between two males, and these conflicts can include biting the face of an opponent. This aggression may be brought on by the defense of foraging areas, basking locations, access to mates, or a combination of these factors (
<bibRefCitation id="EFC14B3F5A3DFFFBD89AFE271D02F8AE" author="LANG, J. W." box="[486,594,1839,1861]" editor="Ross, C. A." journalOrPublisher="Facts on File, New York" pageId="3" pageNumber="783" pagination="102 - 117" refId="ref4240" refString="LANG, J. W., 1989, Social behavior, in Ross, C. A., ed., Crocodiles and Alligators: Facts on File, New York, p. 102 - 117." title="Social behavior" type="book chapter" volumeTitle="Crocodiles and Alligators" year="1989">Lang, 1989</bibRefCitation>
). We speculate that the facial injury of
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was the result of a territorial or dominance bout, similar to those observed in extant crocodilians.
</paragraph>
<paragraph id="8BEF36CE5A3DFFFBD9C9FE8D1BFFFCF3" blockId="3.[159,814,899,2004]" lastBlockId="3.[845,1499,713,1019]" pageId="3" pageNumber="783">
<bibRefCitation id="EFC14B3F5A3DFFFBD9C9FE8D1E36F870" author="BLANCO, G. &amp; TRAVERSO, J. M. &amp; MARCHAMALO, J. &amp; MARTINEZ, F." box="[181,358,1925,1947]" journalOrPublisher="Journal of Raptor Research" pageId="3" pageNumber="783" pagination="77 - 79" part="31" refId="ref3211" refString="BLANCO, G., TRAVERSO, J. M., MARCHAMALO, J., and MARTINEZ, F., 1997, Interspecific and intraspecific aggression among Griffon and Cinereous Vultures at nesting and foraging sites: Journal of Raptor Research, v. 31, p. 77 - 79." title="Interspecific and intraspecific aggression among Griffon and Cinereous Vultures at nesting and foraging sites" type="book chapter" year="1997">Blanco et al. (1997)</bibRefCitation>
documented inter- and intraspecific aggression in cinereous vultures (
<taxonomicName id="4C504D4D5A3DFFFBD823FEAA1D45F85C" baseAuthorityName="Linnaeus" baseAuthorityYear="1766" box="[351,533,1954,1975]" class="Aves" family="Accipitridae" genus="Aegypius" kingdom="Animalia" order="Accipitriformes" pageId="3" pageNumber="783" phylum="Chordata" rank="species" species="monachus">
<emphasis id="B924EADC5A3DFFFBD823FEAA1D45F85C" box="[351,533,1954,1975]" italics="true" pageId="3" pageNumber="783">Aegypius monachus</emphasis>
</taxonomicName>
) and griffon vultures (
<taxonomicName id="4C504D4D5A3DFFFBDA7CFEAA1F83F838" class="Aves" family="Accipitridae" genus="Gyps" kingdom="Animalia" order="Accipitriformes" pageId="3" pageNumber="783" phylum="Chordata" rank="species" species="fluvus">
<emphasis id="B924EADC5A3DFFFBDA7CFEAA1F83F838" italics="true" pageId="3" pageNumber="783">Gyps fluvus</emphasis>
</taxonomicName>
). Aggressive behaviors are commonly intraspecific in griffon
<heading id="D0A781A25A3DFFFBDA31FBC11BFFFCF3" bold="true" centered="true" fontSize="9" level="2" pageId="3" pageNumber="783" reason="0">
vultures, though conflicts rarely result in serious injury. The most violent interactions, where injuries are more likely, however, usually involve juveniles (
<bibRefCitation id="EFC14B3F5A3DFFFBDA8FFA0A1BF5FCF3" author="BLANCO, G. &amp; TRAVERSO, J. M. &amp; MARCHAMALO, J. &amp; MARTINEZ, F." box="[1011,1189,770,792]" journalOrPublisher="Journal of Raptor Research" pageId="3" pageNumber="783" pagination="77 - 79" part="31" refId="ref3211" refString="BLANCO, G., TRAVERSO, J. M., MARCHAMALO, J., and MARTINEZ, F., 1997, Interspecific and intraspecific aggression among Griffon and Cinereous Vultures at nesting and foraging sites: Journal of Raptor Research, v. 31, p. 77 - 79." title="Interspecific and intraspecific aggression among Griffon and Cinereous Vultures at nesting and foraging sites" type="book chapter" year="1997">Blanco et al., 1997</bibRefCitation>
).
</heading>
</paragraph>
<paragraph id="8BEF36CE5A3DFFFBDA1EFA171AF4FC10" blockId="3.[845,1499,713,1019]" pageId="3" pageNumber="783">
While juvenile tyrannosaurids are both rare and poorly known with little evidence upon which to infer their behavior, injuries that have been phylogenetically determined as bite marks suggest behaviors such as intraspecific aggression. Placement of
<materialsCitation id="3B383C935A3DFFFBDDB0FA7C1A31FC61" ID-GBIF-Occurrence="2813097328" box="[1228,1377,884,906]" collectionCode="BMR" pageId="3" pageNumber="783" specimenCode="BMP P2002.4.1">BMR P2002.4.1</materialsCitation>
on a logistic growth curve for
<taxonomicName id="4C504D4D5A3DFFFBDA97FA991BC4FC4D" authority="Osborn, 1905" authorityName="Osborn" authorityYear="1905" box="[1003,1172,913,934]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="783" phylum="Chordata" rank="species" species="rex">
<emphasis id="B924EADC5A3DFFFBDA97FA991BC4FC4D" box="[1003,1172,913,934]" italics="true" pageId="3" pageNumber="783">Tyrannosaurus rex</emphasis>
</taxonomicName>
indicates that the specimen was not yet sexually mature (
<bibRefCitation id="EFC14B3F5A3DFFFBDD75FAA41B97FC29" author="ERICKSON, G. M. &amp; CURRIE, P. J. &amp; INOUYE, B. D. &amp; WINN, A. A." box="[1033,1223,940,962]" journalOrPublisher="Science" pageId="3" pageNumber="783" pagination="213 - 217" part="313" refId="ref3576" refString="ERICKSON, G. M., CURRIE, P. J., INOUYE, B. D., and WINN, A. A., 2006, Tyrannosaur life tables: An example of nonavian dinosaur population biology: Science, v. 313, p. 213 - 217." title="Tyrannosaur life tables: An example of nonavian dinosaur population biology" type="journal article" year="2006">Erickson et al., 2006</bibRefCitation>
), suggesting that this behavior was not related to courtship or mating and that juvenile tyrannosaurids engaged in agonistic behavior prior to reaching sexual maturity.
</paragraph>
</subSubSection>
</treatment>
</document>
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