771 lines
105 KiB
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771 lines
105 KiB
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<document id="EC065FF6C8E28512B433D064B29B4102" ID-DOI="10.1371/journal.pone.0016574" ID-GBIF-Dataset="5a25e523-7350-43a2-b5cb-6cafa6259b25" ID-ISSN="1932-6203" ID-PMC="PMC3036655" ID-PubMed="21347420" ID-Zenodo-Dep="3744916" IM.tables_requiresApprovalFor="existingObjects,jeremy" IM.taxonomicNames_approvedBy="jeremy" IM.treatments_approvedBy="jeremy" approvalRequired="18" approvalRequired_for_illustrations="6" approvalRequired_for_textStreams="10" approvalRequired_for_treatments="2" checkinTime="1586363873496" checkinUser="jeremy" docAuthor="John R. Horner, Mark B. Goodwin & Nathan Myhrvold" docDate="2011" docId="F5541C423B26345FFCD2AAD6FC9022BE" docLanguage="en" docName="Horneretal2011.PDF.imf" docOrigin="PLoS ONE (e 16574) 6, No. 2" docStyle="DocumentStyle{}" docTitle="Tyrannosaurus rex Osborn 1905" docType="treatment" docVersion="5" lastPageNumber="7" masterDocId="096D643A3B253459FF9AAA70FFE7266A" masterDocTitle="Dinosaur Census Reveals Abundant Tyrannosaurus and Rare Ontogenetic Stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA" masterLastPageNumber="9" masterPageNumber="1" pageNumber="4" updateTime="1689602472708" updateUser="jeremy">
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<mods:titleInfo id="DF1AAA37C06BAB6DA29955004DD88FA2">
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<mods:title id="405C4A06AC838EA6168AF8F8929822B0">Dinosaur Census Reveals Abundant Tyrannosaurus and Rare Ontogenetic Stages in the Upper Cretaceous Hell Creek Formation (Maastrichtian), Montana, USA</mods:title>
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<mods:name id="B94F789907076446FD1B5A22E5DFDA6B" type="personal">
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<mods:roleTerm id="C569D062030078F4C0900FD676B5E15B">Author</mods:roleTerm>
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</mods:role>
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<mods:namePart id="EBE2B5003AC300F0EECDFADB370B2679">John R. Horner</mods:namePart>
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<mods:affiliation id="E2C317AA69A0A8979F661C4E3895E9A2">Museum of the Rockies, Montana State University, Bozeman, Montana, United States of America</mods:affiliation>
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<mods:nameIdentifier id="0B7E4487AEAADA90383FF10A2DDAB7DC" type="email">jhorner@montana.edu</mods:nameIdentifier>
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</mods:name>
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<mods:name id="89DE482E9D14CEE450259D6D388DE855" type="personal">
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</mods:role>
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<mods:namePart id="75919089F000549BB802BA9E7DB69CF8">Mark B. Goodwin</mods:namePart>
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<mods:affiliation id="B71DCBEA8899246154E8B3A0D39572EB">Museum of Paleontology, University of California, Berkeley, California, United States of America</mods:affiliation>
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</mods:name>
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<mods:name id="20923FF0051CE5A4C2F9096D708516F3" type="personal">
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</mods:role>
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<mods:namePart id="29FFA9AF0F41DDC399E4C211A5ADABA0">Nathan Myhrvold</mods:namePart>
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<mods:affiliation id="CBECB65BE4FE7A431ECE221628412A74">Intellectual Ventures, Bellevue, Washington, United States of America</mods:affiliation>
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<mods:title id="B4BA7A66B54F60D78D29D5912676E99B">PLoS ONE</mods:title>
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</mods:titleInfo>
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<mods:part id="91103EC6E38AD5FC45DCF163BD7DBB3C">
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<mods:date id="740355CE7D47AC69B9E0284B05545713">2011</mods:date>
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<mods:detail id="FA3C71884A0FFA5416F6CCAE0D0D49C0" type="series">
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<mods:title id="E9741E2D365D69C55BEFD148F1AC0A37">e 16574</mods:title>
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</mods:detail>
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<mods:detail id="199B69B30D08089C859C6D9F23FEACAD" type="pubDate">
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<mods:number id="A290A48ABEF05851BBB363B00B2BA7AB">2011-02-09</mods:number>
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<mods:detail id="F67F2A962F9E5FDC4D5274F47B3BF1C9" type="volume">
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<mods:number id="BA3633C5E8B7AC5BCA386941561F4CDE">6</mods:number>
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<mods:number id="738E70D58280A40A4685BA25274D2CBA">2</mods:number>
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<mods:classification id="B07B5DBE39289C76E07240FEABE825F2">journal article</mods:classification>
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<mods:identifier id="1DABA764D54CD442714FDA9A1D9A471E" type="DOI">10.1371/journal.pone.0016574</mods:identifier>
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<mods:identifier id="0503C31528CF388431F64062BFDAE391" type="GBIF-Dataset">5a25e523-7350-43a2-b5cb-6cafa6259b25</mods:identifier>
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<mods:identifier id="F647AB70A326DC515715A83510433E98" type="ISSN">1932-6203</mods:identifier>
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<mods:identifier id="6858DB1E75A49772C1B560CBBDB7CC33" type="PMC">PMC3036655</mods:identifier>
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<mods:identifier id="F23C7374AA0CC6B8C17525B3BE34FD6D" type="PubMed">21347420</mods:identifier>
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<mods:identifier id="83D9B6014471522C9890ADF5832AEABA" type="Zenodo-Dep">3744916</mods:identifier>
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<treatment id="F5541C423B26345FFCD2AAD6FC9022BE" ID-DOI="http://doi.org/10.5281/zenodo.3809607" ID-GBIF-Taxon="184621772" ID-Zenodo-Dep="3809607" LSID="urn:lsid:plazi:treatment:F5541C423B26345FFCD2AAD6FC9022BE" httpUri="http://treatment.plazi.org/id/F5541C423B26345FFCD2AAD6FC9022BE" lastPageId="6" lastPageNumber="7" pageId="3" pageNumber="4">
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<subSubSection id="35E7FEDF3B26345AFCD2AAD6FC4426D4" box="[840,931,166,190]" pageId="3" pageNumber="4" type="nomenclature">
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<paragraph id="7D42AD543B26345AFCD2AAD6FC4426D4" blockId="3.[840,931,166,190]" box="[840,931,166,190]" pageId="3" pageNumber="4">
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<heading id="260A1A383B26345AFCD2AAD6FC4426D4" bold="true" box="[840,931,166,190]" fontSize="10" level="1" pageId="3" pageNumber="4" reason="0">
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<taxonomicName id="BAFDD6D73B26345AFCD2AAD6FC4426D4" authorityName="Osborn" authorityYear="1905" box="[840,931,166,190]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="9" phylum="Chordata" rank="species" species="rex">
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<emphasis id="4F8971463B26345AFCD2AAD6FC4426D4" bold="true" box="[840,931,166,190]" pageId="3" pageNumber="4">Results</emphasis>
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</taxonomicName>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="35E7FEDF3B26345AFCD2AAA7FB7C2412" pageId="3" pageNumber="4" type="discussion">
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<paragraph id="7D42AD543B26345AFCD2AAA7FBF22685" blockId="3.[840,1478,215,632]" box="[840,1045,215,239]" pageId="3" pageNumber="4">
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<heading id="260A1A383B26345AFCD2AAA7FBF22685" box="[840,1045,215,239]" fontSize="10" level="2" pageId="3" pageNumber="4" reason="1">Geological Results</heading>
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</paragraph>
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<paragraph id="7D42AD543B26345AFCFAAA85FB7C2412" blockId="3.[840,1478,215,632]" pageId="3" pageNumber="4">
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<taxonomicName id="BAFDD6D73B26345AFCFAAA85FC3A2761" authorityName="Lambe" authorityYear="1917" box="[864,989,245,267]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="4F8971463B26345AFCFAAA85FC3A2761" box="[864,989,245,267]" italics="true" pageId="3" pageNumber="4">Edmontosaurus</emphasis>
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</taxonomicName>
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,
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<taxonomicName id="BAFDD6D73B26345AFC6DAA85FB8B2761" authorityName="Marsh" authorityYear="1890" box="[1015,1132,245,267]" class="Reptilia" family="Ornithomimidae" genus="Ornithomimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="4F8971463B26345AFC6DAA85FB8B2761" box="[1015,1132,245,267]" italics="true" pageId="3" pageNumber="4">Ornithomimus</emphasis>
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</taxonomicName>
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and
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<taxonomicName id="BAFDD6D73B26345AFB2DAA85FAC42761" authorityName="Brown" authorityYear="1908" box="[1207,1315,245,267]" class="Reptilia" family="Ankylosauridae" genus="Ankylosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="4F8971463B26345AFB2DAA85FAC42761" box="[1207,1315,245,267]" italics="true" pageId="3" pageNumber="4">Ankylosaurus</emphasis>
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</taxonomicName>
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are found in siltstones or sandstones, and
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<taxonomicName id="BAFDD6D73B26345AFBF2AB61FB05274D" authorityName="Gilmore" authorityYear="1913" box="[1128,1250,273,295]" class="Reptilia" family="Thescelosauridae" genus="Thescelosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="4F8971463B26345AFBF2AB61FB05274D" box="[1128,1250,273,295]" italics="true" pageId="3" pageNumber="4">Thescelosaurus</emphasis>
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</taxonomicName>
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is found exclusively in mudstones, but the relative number of specimens is small and subsequently questionable as a real pattern of sediment preference or taphonomic artifact (
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<tableCitation id="307F98EF3B26345AFBACAB16FB7E2716" box="[1078,1177,358,380]" captionStart="Table 1" captionStartId="4.[171,235,174,195]" captionTargetBox="[171,1414,252,856]" captionTargetId="graphics@4.[163,1478,151,962]" captionTargetPageId="4" captionText="Table 1. Hell Creek Formation dinosaur census." httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="3" pageNumber="4" tableUuid="2982FDDC3B21345DFF31AADEFD7F26A9">Tables S1</tableCitation>
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, S4). Other taxa are found in both channel and overbank sediments, but the majority of
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<taxonomicName id="BAFDD6D73B26345AFCD2ABEDFC4327D9" authorityName="Marsh" authorityYear="1889" box="[840,932,413,435]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="4F8971463B26345AFCD2ABEDFC4327D9" box="[840,932,413,435]" italics="true" pageId="3" pageNumber="4">Triceratops</emphasis>
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</taxonomicName>
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, and in particular juvenile specimens, come primarily from mudstones [18]. There was no apparent sediment preference for preserving articulation. The basal sand unit (L3.lBS) produced both an articulated specimen of
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<taxonomicName id="BAFDD6D73B26345AFB04AB81FAFC246D" authorityName="Lambe" authorityYear="1917" box="[1182,1307,497,519]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="4F8971463B26345AFB04AB81FAFC246D" box="[1182,1307,497,519]" italics="true" pageId="3" pageNumber="4">Edmontosaurus</emphasis>
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</taxonomicName>
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(‘‘X-rex’’/MOR 1142) and a disarticulated
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<taxonomicName id="BAFDD6D73B26345AFBDCA87DFB262449" box="[1094,1217,525,547]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="4F8971463B26345AFBDCA87DFB262449" box="[1094,1217,525,547]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
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</taxonomicName>
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(
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<materialsCitation id="CD95A7093B26345AFB54A87EFA7E244E" ID-GBIF-Occurrence="3336525305" box="[1230,1433,526,548]" collectionCode="MOR" country="United States" formation="Hell Creek Formation" pageId="3" pageNumber="4" specimenCode="MOR 1125" stateProvince="Montana">‘‘B-rex’’/MOR 1125</materialsCitation>
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). An articulated
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<taxonomicName id="BAFDD6D73B26345AFC2DA859FBD52455" box="[951,1074,553,575]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="4F8971463B26345AFC2DA859FBD52455" box="[951,1074,553,575]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
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</taxonomicName>
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(
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<materialsCitation id="CD95A7093B26345AFBDBA85AFA67242A" ID-GBIF-Occurrence="3336525302" box="[1089,1408,554,576]" collectionCode="USNM" country="United States" formation="Hell Creek Formation" pageId="3" pageNumber="4" specimenCode="N-rex" stateProvince="Montana">‘‘N-rex’’/Smithsonian Institution</materialsCitation>
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) and a disarticulated
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<taxonomicName id="BAFDD6D73B26345AFC48A835FBAA2431" box="[978,1101,581,603]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="4F8971463B26345AFC48A835FBAA2431" box="[978,1101,581,603]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
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</taxonomicName>
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(
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<materialsCitation id="CD95A7093B26345AFBC7A836FACB2436" ID-GBIF-Occurrence="3336525306" box="[1117,1324,582,604]" collectionCode="MOR" country="United States" formation="Hell Creek Formation" pageId="3" pageNumber="4" specimenCode="MOR 1128" stateProvince="Montana">‘‘G-rex’’/MOR 1128</materialsCitation>
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) were found in the lower mudstone unit (L3.lMS).
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</paragraph>
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</subSubSection>
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<subSubSection id="35E7FEDF3B26345DFCD2A8EDFB502360" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4" type="description">
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<paragraph id="7D42AD543B26345AFCD2A8EDFC0924DF" blockId="3.[840,1479,669,1956]" box="[840,1006,669,693]" pageId="3" pageNumber="4">
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<heading id="260A1A383B26345AFCD2A8EDFC0924DF" box="[840,1006,669,693]" fontSize="10" level="2" pageId="3" pageNumber="4" reason="1">Census Results</heading>
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</paragraph>
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<paragraph id="7D42AD543B26345AFCFAA8CCFAED22A0" blockId="3.[840,1479,669,1956]" pageId="3" pageNumber="4">
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The dinosaur census results are summarized in
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<tableCitation id="307F98EF3B26345AFAC8A8CCFA4524B8" box="[1362,1442,700,722]" captionStart="Table 1" captionStartId="4.[171,235,174,195]" captionTargetBox="[171,1414,252,856]" captionTargetId="graphics@4.[163,1478,151,962]" captionTargetPageId="4" captionText="Table 1. Hell Creek Formation dinosaur census." httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="3" pageNumber="4" tableUuid="2982FDDC3B21345DFF31AADEFD7F26A9">Table 1</tableCitation>
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by taxon with percentage of the fauna and absolute numbers given. Additional sedimentological details, more precise stratigraphic interval, preservation and ontogenetic designations are provided in
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<tableCitation id="307F98EF3B26345AFCD2A95CFC4D2528" box="[840,938,812,834]" captionStart="Table 1" captionStartId="4.[171,235,174,195]" captionTargetBox="[171,1414,252,856]" captionTargetId="graphics@4.[163,1478,151,962]" captionTargetPageId="4" captionText="Table 1. Hell Creek Formation dinosaur census." httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="3" pageNumber="4" tableUuid="2982FDDC3B21345DFF31AADEFD7F26A9">Tables S1</tableCitation>
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, S2, S3, S4, S5, S6. The isolated, uncollected
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<taxonomicName id="BAFDD6D73B26345AFAF0A95BFA21252B" authorityName="Marsh" authorityYear="1889" box="[1386,1478,811,833]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
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<emphasis id="4F8971463B26345AFAF0A95BFA21252B" box="[1386,1478,811,833]" italics="true" pageId="3" pageNumber="4">Triceratops</emphasis>
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</taxonomicName>
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skulls listed in Table S3 are not included in the census of skeletons from the lower Hell Creek Formation (
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<tableCitation id="307F98EF3B26345AFB77A914FAAD2510" box="[1261,1354,868,890]" captionStart="Table 1" captionStartId="4.[171,235,174,195]" captionTargetBox="[171,1414,252,856]" captionTargetId="graphics@4.[163,1478,151,962]" captionTargetPageId="4" captionText="Table 1. Hell Creek Formation dinosaur census." httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="3" pageNumber="4" tableUuid="2982FDDC3B21345DFF31AADEFD7F26A9">Table S1</tableCitation>
|
|||
|
|
) at present because there is no way to know if they consist of three or more disarticulated pieces until they are collected. Thirty-nine skeletons (not counting the isolated
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFBD8A9C7FB7925A7" authorityName="Marsh" authorityYear="1889" box="[1090,1182,951,973]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFBD8A9C7FB7925A7" box="[1090,1182,951,973]" italics="true" pageId="3" pageNumber="4">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
skulls) were recorded from the L3 strata. All but three of these skeletons were collected. The uncollected specimens were represented by at least three elements but were too severely eroded to yield data other than for this census. In addition, seven specimens (superscript
|
|||
|
|
<superScript id="8A88001C3B26345AFAA6AE53FAA2225B" attach="left" box="[1340,1349,1059,1073]" fontSize="6" pageId="3" pageNumber="4">3</superScript>
|
|||
|
|
numbers in
|
|||
|
|
<tableCitation id="307F98EF3B26345AFCD2AE34FC4D2230" box="[840,938,1092,1114]" captionStart="Table 1" captionStartId="4.[171,235,174,195]" captionTargetBox="[171,1414,252,856]" captionTargetId="graphics@4.[163,1478,151,962]" captionTargetPageId="4" captionText="Table 1. Hell Creek Formation dinosaur census." httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="3" pageNumber="4" tableUuid="2982FDDC3B21345DFF31AADEFD7F26A9">Table S1</tableCitation>
|
|||
|
|
) consisted of only three elements each. Limited excavation around the elements failed to yield more material, and the sites were abandoned. Five specimens were found with some articulation, and of these, only one (
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFB7EAEE7FA8622C7" authorityName="Lambe" authorityYear="1917" box="[1252,1377,1175,1197]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFB7EAEE7FA8622C7" box="[1252,1377,1175,1197]" italics="true" pageId="3" pageNumber="4">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
, ‘‘X-rex,’’ MOR 1142) was found with skin impressions.
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B26345AFCFAAEA1FBA9205D" blockId="3.[840,1479,669,1956]" pageId="3" pageNumber="4">
|
|||
|
|
The most interesting census result in the L3 is the high number of
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFCFBAE9CFC3B2368" box="[865,988,1260,1282]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFCFBAE9CFC3B2368" box="[865,988,1260,1282]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
skeletons (n = 11) that is nearly double the number of
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFCFBAF79FC392375" authorityName="Lambe" authorityYear="1917" box="[865,990,1289,1311]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFCFBAF79FC392375" box="[865,990,1289,1311]" italics="true" pageId="3" pageNumber="4">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
skeletons (n = 6) and equals
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFB68AF79FAA92375" authorityName="Marsh" authorityYear="1889" box="[1266,1358,1289,1311]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFB68AF79FAA92375" box="[1266,1358,1289,1311]" italics="true" pageId="3" pageNumber="4">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
(n = 11) (see
|
|||
|
|
<figureCitation id="E5C6B1D13B26345AFCD2AF55FC7A2351" box="[840,925,1317,1339]" captionStart="Figure 1" captionStartId="1.[155,220,1839,1858]" captionTargetBox="[160,1224,1232,1801]" captionTargetId="figure@1.[155,1229,1227,1806]" captionTargetPageId="1" captionText="Figure 1. Index map of the Upper Cretaceous Hell Creek Formation along Ft. Peck Lake in northeastern Montanaı USA. Contiguous outcrops are traceable over an area of about 1000 square km adjacent to Fort Peck Lake shown in blue. Dark orange represents the lower Hell Creek Formation and light orange represents the middle and upper Hell Creek Formation. The enlarged view of the study area is indicated by the rectangle in the northeast quarter of the map of Montana. doi:10.1371/journal.pone.0016574.g001" figureDoi="http://doi.org/10.5281/zenodo.3744918" httpUri="https://zenodo.org/record/3744918/files/figure.png" pageId="3" pageNumber="4">Figure 1</figureCitation>
|
|||
|
|
and
|
|||
|
|
<tableCitation id="307F98EF3B26345AFC42AF55FBC12351" box="[984,1062,1317,1339]" captionStart="Table 1" captionStartId="4.[171,235,174,195]" captionTargetBox="[171,1414,252,856]" captionTargetId="graphics@4.[163,1478,151,962]" captionTargetPageId="4" captionText="Table 1. Hell Creek Formation dinosaur census." httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="3" pageNumber="4" tableUuid="2982FDDC3B21345DFF31AADEFD7F26A9">Table 1</tableCitation>
|
|||
|
|
). However, as explained in the previous paragraph, it is likely the number of
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFB42AF30FAD3233C" authorityName="Marsh" authorityYear="1889" box="[1240,1332,1344,1366]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFB42AF30FAD3233C" box="[1240,1332,1344,1366]" italics="true" pageId="3" pageNumber="4">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
skeletons will increase as these sites in L3 are excavated.
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFAD1AF2DFA212319" box="[1355,1478,1373,1395]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFAD1AF2DFA212319" box="[1355,1478,1373,1395]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
contributes to 28% of the dinosaur skeletons recorded in L3 while
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFCD2AFE4FC2223C3" authorityName="Lambe" authorityYear="1917" box="[840,965,1428,1449]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFCD2AFE4FC2223C3" box="[840,965,1428,1449]" italics="true" pageId="3" pageNumber="4">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
makes up only 15%. Considering the fact that
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFCD2AFC1FC2423AC" box="[840,963,1457,1478]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFCD2AFC1FC2423AC" box="[840,963,1457,1478]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
,
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFC55AFC1FBCC23AC" authorityName="Marsh" authorityYear="1889" box="[975,1067,1457,1478]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFC55AFC1FBCC23AC" box="[975,1067,1457,1478]" italics="true" pageId="3" pageNumber="4">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
, and
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFBF8AFC0FB3823AF" authorityName="Lambe" authorityYear="1917" box="[1122,1247,1456,1477]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFBF8AFC0FB3823AF" box="[1122,1247,1456,1477]" italics="true" pageId="3" pageNumber="4">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
are all relatively similar in size as full-grown adults, we presume that there are few taphonomic biases that would amplify the
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFB75AF99FA8D2394" box="[1263,1386,1513,1534]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFB75AF99FA8D2394" box="[1263,1386,1513,1534]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
numbers to be greater than
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFB9DAC74FB632073" authorityName="Lambe" authorityYear="1917" box="[1031,1156,1540,1561]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFB9DAC74FB632073" box="[1031,1156,1540,1561]" italics="true" pageId="3" pageNumber="4">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
and this likely reflects a correct ratio of approximately 2:1.
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B26345DFCFAAC4EFDAE2287" blockId="3.[840,1479,669,1956]" lastBlockId="4.[155,794,1012,1717]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
|
|||
|
|
Thirty-two skeletons were collected from the U3 unit, four of which were collected prior to the Hell Creek Project by the Museum of the Rockies (
|
|||
|
|
<materialsCitation id="CD95A7093B26345AFBD7AC06FB5E20E6" ID-GBIF-Occurrence="3336525301" box="[1101,1209,1654,1676]" collectionCode="MOR" country="United States" formation="Hell Creek Formation" pageId="3" pageNumber="4" specimenCode="MOR 009" stateProvince="Montana">MOR 009</materialsCitation>
|
|||
|
|
,
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFB53AC06FAA320E1" box="[1225,1348,1654,1675]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFB53AC06FAA320E1" box="[1225,1348,1654,1675]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
; MOR 004,
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFCD2ACE2FC4320CD" authorityName="Marsh" authorityYear="1889" box="[840,932,1682,1703]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFCD2ACE2FC4320CD" box="[840,932,1682,1703]" italics="true" pageId="3" pageNumber="4">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
;
|
|||
|
|
<materialsCitation id="CD95A7093B26345AFC28ACE2FBFD20C2" ID-GBIF-Occurrence="3336525307" box="[946,1050,1682,1704]" collectionCode="MOR" country="United States" formation="Hell Creek Formation" pageId="3" pageNumber="4" specimenCode="MOR 555" stateProvince="Montana">MOR 555</materialsCitation>
|
|||
|
|
,
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFBBFACE2FB4720CD" box="[1061,1184,1682,1703]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFBBFACE2FB4720CD" box="[1061,1184,1682,1703]" italics="true" pageId="3" pageNumber="4">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
; MOR 622,
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFABBACE2FA9A20CD" authorityName="Marsh" authorityYear="1889" box="[1313,1405,1682,1703]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFABBACE2FA9A20CD" box="[1313,1405,1682,1703]" italics="true" pageId="3" pageNumber="4">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
; MOR 007,
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFCE7ACDEFC1D20A9" authorityName="Lambe" authorityYear="1917" box="[893,1018,1710,1731]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFCE7ACDEFC1D20A9" box="[893,1018,1710,1731]" italics="true" pageId="3" pageNumber="4">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
). These were included in the census because they were found in the study area with documented stratigraphic and locality information, and they are cataloged into MOR.
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFCD2AD72FC43217D" authorityName="Marsh" authorityYear="1889" box="[840,932,1794,1815]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFCD2AD72FC43217D" box="[840,932,1794,1815]" italics="true" pageId="3" pageNumber="4">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
skeletons (n = 22) greatly outnumbered other taxa (
|
|||
|
|
<figureCitation id="E5C6B1D13B26345AFCD4AD6EFC43215E" box="[846,932,1822,1844]" captionStart="Figure 2" captionStartId="2.[155,220,1221,1240]" captionTargetBox="[160,1012,169,1180]" captionTargetId="figure@2.[155,1017,162,1187]" captionTargetPageId="2" captionText="Figure 2. Stratigraphic divisions of the Hell Creek Formation with stratigraphic details of the upper third (U3) and lower third (L3) sequences referenced in the text and the associated pie chart showing dinosaur abundance. Pie charts I – IV reflect the relative abundance of dinosaur genera based on skeletons (charts I and IV) and individual bones (charts II and III). Tyrannosaurus skeletons are as abundant as the hadrosaurid Edmontosaurus, an herbivore, in the upper Hell Creek Formation and nearly twice as common as Edmontosaurus in the lower Hell Creek Formation. Individual bones of Tyrannosaurus and Edmontosaurus are found approximately in the same order of magnitude within the lag deposits. Key: Pie chart I, dinosaur skeletons in the upper Hell Creek Formation (U3); Pie chart II, dinosaur bones from the ‘‘Doldrum’s’’ lag deposit at the base of the Apex Sandstone (AS); Pie chart III, dinosaur bones from the ‘‘3B-1’’ lag deposit at the base of the Jen-rex Sand; and Pie chart IV, dinosaur skeletons from the lower Hell Creek Formation (L3). Abbreviations: L3lBS, lower basal sandstone; L3.mBS, middle basal sandstone; L3.uBS, upper basal sandstone; L3.lMS, lower mudstone; L3.mMS, middle mudstone; L3.uMS, upper mudstone; U3.AS, Apex sandstone; U3.ibMS, interbedded mudstone; U3.10mS, 10 meter sandstone; U3.uMS, upper mudstone; U3.CZ, coal zone. doi:10.1371/journal.pone.0016574.g002" figureDoi="http://doi.org/10.5281/zenodo.3744920" httpUri="https://zenodo.org/record/3744920/files/figure.png" pageId="3" pageNumber="4">Figure 2</figureCitation>
|
|||
|
|
,
|
|||
|
|
<tableCitation id="307F98EF3B26345AFC2BAD6EFC1B215E" box="[945,1020,1822,1844]" captionStart="Table 1" captionStartId="4.[171,235,174,195]" captionTargetBox="[171,1414,252,856]" captionTargetId="graphics@4.[163,1478,151,962]" captionTargetPageId="4" captionText="Table 1. Hell Creek Formation dinosaur census." httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="3" pageNumber="4" tableUuid="2982FDDC3B21345DFF31AADEFD7F26A9">Table 1</tableCitation>
|
|||
|
|
) and contribute to 69% of the total dinosaur skeletal fauna in U3. Specimens of
|
|||
|
|
<emphasis id="4F8971463B26345AFB25AD4AFA272125" box="[1215,1472,1850,1871]" italics="true" pageId="3" pageNumber="4">
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFB25AD4AFAD12125" authorityName="Marsh" authorityYear="1890" box="[1215,1334,1850,1871]" class="Reptilia" family="Ornithomimidae" genus="Ornithomimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Ornithomimus</taxonomicName>
|
|||
|
|
,
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFADCAD4AFA272125" authorityName="Gilmore" authorityYear="1913" box="[1350,1472,1850,1871]" class="Reptilia" family="Thescelosauridae" genus="Thescelosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Thescelosaurus</taxonomicName>
|
|||
|
|
</emphasis>
|
|||
|
|
,
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFCD2AD26FC532101" authorityName="Brown" authorityYear="1908" box="[840,948,1878,1899]" class="Reptilia" family="Ankylosauridae" genus="Ankylosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFCD2AD26FC532101" box="[840,948,1878,1899]" italics="true" pageId="3" pageNumber="4">Ankylosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
or
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFC68AD26FB742101" authorityName="Brown & Schlaikjer" authorityYear="1943" box="[1010,1171,1878,1899]" class="Reptilia" family="Pachycephalosauridae" genus="Pachycephalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFC68AD26FB742101" box="[1010,1171,1878,1899]" italics="true" pageId="3" pageNumber="4">Pachycephalosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
were conspicuously absent, although isolated bones of
|
|||
|
|
<emphasis id="4F8971463B26345AFBE5AD02FA6F21ED" box="[1151,1416,1906,1927]" italics="true" pageId="3" pageNumber="4">
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFBE5AD02FB1B21ED" authorityName="Gilmore" authorityYear="1913" box="[1151,1276,1906,1927]" class="Reptilia" family="Thescelosauridae" genus="Thescelosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Thescelosaurus</taxonomicName>
|
|||
|
|
,
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFA89AD02FA6F21ED" authorityName="Marsh" authorityYear="1890" box="[1299,1416,1906,1927]" class="Reptilia" family="Ornithomimidae" genus="Ornithomimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Ornithomimus</taxonomicName>
|
|||
|
|
</emphasis>
|
|||
|
|
, and
|
|||
|
|
<taxonomicName id="BAFDD6D73B26345AFCD2ADFEFC0E21C9" authorityName="Brown & Schlaikjer" authorityYear="1943" box="[840,1001,1934,1955]" class="Reptilia" family="Pachycephalosauridae" genus="Pachycephalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B26345AFCD2ADFEFC0E21C9" box="[840,1001,1934,1955]" italics="true" pageId="3" pageNumber="4">Pachycephalosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
were present in the Doldrum’s Lag deposit (MOR loc. HC-530) at the base of the Apex sandstone.
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFF01AE60FEFF224C" authorityName="Lambe" authorityYear="1917" box="[155,280,1040,1062]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFF01AE60FEFF224C" box="[155,280,1040,1062]" italics="true" pageId="4" pageNumber="5">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
and
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFEC8AE60FE2A224C" box="[338,461,1040,1062]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFEC8AE60FE2A224C" box="[338,461,1040,1062]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
skeletons were equal in number (n = 5) in U3 and comprise 16% each of the large dinosaur taxa. The pie charts in
|
|||
|
|
<figureCitation id="E5C6B1D13B21345DFED9AE39FE752235" box="[323,402,1097,1119]" captionStart="Figure 2" captionStartId="2.[155,220,1221,1240]" captionTargetBox="[160,1012,169,1180]" captionTargetId="figure@2.[155,1017,162,1187]" captionTargetPageId="2" captionText="Figure 2. Stratigraphic divisions of the Hell Creek Formation with stratigraphic details of the upper third (U3) and lower third (L3) sequences referenced in the text and the associated pie chart showing dinosaur abundance. Pie charts I – IV reflect the relative abundance of dinosaur genera based on skeletons (charts I and IV) and individual bones (charts II and III). Tyrannosaurus skeletons are as abundant as the hadrosaurid Edmontosaurus, an herbivore, in the upper Hell Creek Formation and nearly twice as common as Edmontosaurus in the lower Hell Creek Formation. Individual bones of Tyrannosaurus and Edmontosaurus are found approximately in the same order of magnitude within the lag deposits. Key: Pie chart I, dinosaur skeletons in the upper Hell Creek Formation (U3); Pie chart II, dinosaur bones from the ‘‘Doldrum’s’’ lag deposit at the base of the Apex Sandstone (AS); Pie chart III, dinosaur bones from the ‘‘3B-1’’ lag deposit at the base of the Jen-rex Sand; and Pie chart IV, dinosaur skeletons from the lower Hell Creek Formation (L3). Abbreviations: L3lBS, lower basal sandstone; L3.mBS, middle basal sandstone; L3.uBS, upper basal sandstone; L3.lMS, lower mudstone; L3.mMS, middle mudstone; L3.uMS, upper mudstone; U3.AS, Apex sandstone; U3.ibMS, interbedded mudstone; U3.10mS, 10 meter sandstone; U3.uMS, upper mudstone; U3.CZ, coal zone. doi:10.1371/journal.pone.0016574.g002" figureDoi="http://doi.org/10.5281/zenodo.3744920" httpUri="https://zenodo.org/record/3744920/files/figure.png" pageId="4" pageNumber="5">Figure 2</figureCitation>
|
|||
|
|
illustrate the similarity in overall percent composition between the large dinosaur fauna recorded in L3 and the two overlying lag deposits. The greatest contrast occurs within the upper Hell Creek (U3) record of dinosaur skeletons where
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFF01AECBFF1022BB" authorityName="Marsh" authorityYear="1889" box="[155,247,1211,1233]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFF01AECBFF1022BB" box="[155,247,1211,1233]" italics="true" pageId="4" pageNumber="5">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
dominates (69%; n = 22), followed by
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFD04AECBFCFE22BB" box="[670,793,1211,1233]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFD04AECBFCFE22BB" box="[670,793,1211,1233]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
(16%; n = 5) and
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFEDCAEA7FE242287" authorityName="Lambe" authorityYear="1917" box="[326,451,1239,1261]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFEDCAEA7FE242287" box="[326,451,1239,1261]" italics="true" pageId="4" pageNumber="5">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
(16%; n = 5).
|
|||
|
|
</paragraph>
|
|||
|
|
<caption id="2982FDDC3B21345DFF31AADEFD7F26A9" ID-Table-UUID="2982FDDC3B21345DFF31AADEFD7F26A9" box="[171,664,174,195]" httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="4" pageNumber="5" startId="4.[171,235,174,195]" targetBox="[171,1414,252,856]" targetIsTable="true" targetPageId="4">
|
|||
|
|
<paragraph id="7D42AD543B21345DFF31AADEFD7F26A9" box="[171,664,174,195]" pageId="4" pageNumber="5">
|
|||
|
|
<emphasis id="4F8971463B21345DFF31AADEFEE026A9" bold="true" box="[171,263,174,195]" pageId="4" pageNumber="5">Table 1.</emphasis>
|
|||
|
|
Hell Creek Formation dinosaur census.
|
|||
|
|
</paragraph>
|
|||
|
|
</caption>
|
|||
|
|
<paragraph id="7D42AD543B21345DFC52AA8CFA91252B" pageId="4" pageNumber="5">
|
|||
|
|
<table id="0FFD5FF43B21CBA6FF31AA8CFA612532" box="[171,1414,252,856]" gridcols="9" gridrows="11" pageId="4" pageNumber="5">
|
|||
|
|
<emphasis id="4F8971463B21345DFC52AA8CFA612744" bold="true" pageId="4" pageNumber="5">
|
|||
|
|
<tr id="C3CDAF163B21CBA6FF31AA8CFA612767" box="[171,1414,252,269]" gridrow="0" pageId="4" pageNumber="5">
|
|||
|
|
<th id="801CC66A3B21CBA6FF31AA8CFA612767" box="[171,1414,252,269]" colspan="9" colspanRight="8" gridcol="0" gridrow="0" pageId="4" pageNumber="5">
|
|||
|
|
<emphasis id="4F8971463B21345DFC52AA8CFC182767" bold="true" box="[968,1023,252,269]" pageId="4" pageNumber="5">Taxon</emphasis>
|
|||
|
|
</th>
|
|||
|
|
</tr>
|
|||
|
|
<tr id="C3CDAF163B21CBA6FF31AB6DFA612744" box="[171,1414,285,302]" gridrow="1" pageId="4" pageNumber="5" rowspan-1="1">
|
|||
|
|
<th id="801CC66A3B21CBA6FF31AB6DFE2A2744" box="[171,461,285,302]" gridcol="0" gridrow="1" pageId="4" pageNumber="5">
|
|||
|
|
<emphasis id="4F8971463B21345DFF31AB6DFEB72744" bold="true" box="[171,336,285,302]" pageId="4" pageNumber="5">Stratigraphic level</emphasis>
|
|||
|
|
</th>
|
|||
|
|
<td id="801CC66A3B21CBA6FD4AAB6DFD162744" box="[720,753,285,302]" gridcol="2" gridrow="1" pageId="4" pageNumber="5">
|
|||
|
|
<emphasis id="4F8971463B21345DFD4AAB6DFD0A2744" bold="true" box="[720,749,285,302]" pageId="4" pageNumber="5">Tric</emphasis>
|
|||
|
|
</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FCB1AB6DFCA82744" box="[811,847,285,302]" gridcol="3" gridrow="1" pageId="4" pageNumber="5">
|
|||
|
|
<emphasis id="4F8971463B21345DFCB1AB6DFCA82744" bold="true" box="[811,847,285,302]" pageId="4" pageNumber="5">Tyrn</emphasis>
|
|||
|
|
</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FC0EAB6DFC242744" box="[916,963,285,302]" gridcol="4" gridrow="1" pageId="4" pageNumber="5">
|
|||
|
|
<emphasis id="4F8971463B21345DFC0EAB6DFC242744" bold="true" box="[916,963,285,302]" pageId="4" pageNumber="5">Edmn</emphasis>
|
|||
|
|
</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FB93AB6DFBC82744" box="[1033,1071,285,302]" gridcol="5" gridrow="1" pageId="4" pageNumber="5">
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFB93AB6DFBC82744" box="[1033,1071,285,302]" class="Insecta" family="Latridiidae" genus="Thes" higherTaxonomySource="GBIF" kingdom="Animalia" order="Coleoptera" pageId="4" pageNumber="5" phylum="Arthropoda" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFB93AB6DFBC82744" bold="true" box="[1033,1071,285,302]" pageId="4" pageNumber="5">Thes</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FBE8AB6DFB722744" box="[1138,1173,285,302]" gridcol="6" gridrow="1" pageId="4" pageNumber="5">
|
|||
|
|
<emphasis id="4F8971463B21345DFBE8AB6DFB722744" bold="true" box="[1138,1173,285,302]" pageId="4" pageNumber="5">Orni</emphasis>
|
|||
|
|
</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FB41AB6DFAEC2744" box="[1243,1291,285,302]" gridcol="7" gridrow="1" pageId="4" pageNumber="5">
|
|||
|
|
<emphasis id="4F8971463B21345DFB41AB6DFAEC2744" bold="true" box="[1243,1291,285,302]" pageId="4" pageNumber="5">Pachy</emphasis>
|
|||
|
|
</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FAC7AB6DFA612744" box="[1373,1414,285,302]" gridcol="8" gridrow="1" pageId="4" pageNumber="5">
|
|||
|
|
<emphasis id="4F8971463B21345DFAC7AB6DFA612744" bold="true" box="[1373,1414,285,302]" pageId="4" pageNumber="5">Anky</emphasis>
|
|||
|
|
</td>
|
|||
|
|
</tr>
|
|||
|
|
</emphasis>
|
|||
|
|
<tr id="C3CDAF163B21CBA6FF31AB35FA612706" box="[171,1414,325,364]" gridrow="2" pageId="4" pageNumber="5" rowspan-5="1" rowspan-6="1" rowspan-7="1" rowspan-8="1">
|
|||
|
|
<th id="801CC66A3B21CBA6FF31AB35FE2A2706" box="[171,461,325,364]" gridcol="0" gridrow="2" pageId="4" pageNumber="5">Upper Hell Creek Fm (U3) skeletons</th>
|
|||
|
|
<td id="801CC66A3B21CBA6FD14AB35FD402706" box="[654,679,325,364]" gridcol="1" gridrow="2" pageId="4" pageNumber="5">n = %</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FD4AAB35FD162706" box="[720,753,325,364]" gridcol="2" gridrow="2" pageId="4" pageNumber="5">23 69%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FCB1AB35FCA82706" box="[811,847,325,364]" gridcol="3" gridrow="2" pageId="4" pageNumber="5">5 16%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FC0EAB35FC242706" box="[916,963,325,364]" gridcol="4" gridrow="2" pageId="4" pageNumber="5">5 16%</td>
|
|||
|
|
</tr>
|
|||
|
|
<tr id="C3CDAF163B21CBA6FF31AB0DFA6127CE" box="[171,1414,381,420]" gridrow="3" pageId="4" pageNumber="5">
|
|||
|
|
<th id="801CC66A3B21CBA6FF31AB0DFA6127CE" box="[171,1414,381,420]" colspan="9" colspanRight="8" gridcol="0" gridrow="3" pageId="4" pageNumber="5">Pie chart Iı Fig. 2 Table S4</th>
|
|||
|
|
</tr>
|
|||
|
|
<tr id="C3CDAF163B21CBA6FF31ABC4FA612799" box="[171,1414,436,499]" gridrow="4" pageId="4" pageNumber="5" rowspan-8="1">
|
|||
|
|
<th id="801CC66A3B21CBA6FF31ABC4FE2A2799" box="[171,461,436,499]" gridcol="0" gridrow="4" pageId="4" pageNumber="5">"Doldrum’s" lag deposit at base of Apex sandstone (MOR locality HC-530)</th>
|
|||
|
|
<td id="801CC66A3B21CBA6FD14ABC4FD402799" box="[654,679,436,499]" gridcol="1" gridrow="4" pageId="4" pageNumber="5">n = %</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FD4AABC4FD162799" box="[720,753,436,499]" gridcol="2" gridrow="4" pageId="4" pageNumber="5">16 41%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FCB1ABC4FCA82799" box="[811,847,436,499]" gridcol="3" gridrow="4" pageId="4" pageNumber="5">9 23%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FC0EABC4FC242799" box="[916,963,436,499]" gridcol="4" gridrow="4" pageId="4" pageNumber="5">7 18%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FB93ABC4FBC82799" box="[1033,1071,436,499]" gridcol="5" gridrow="4" pageId="4" pageNumber="5">4 10%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FBE8ABC4FB722799" box="[1138,1173,436,499]" gridcol="6" gridrow="4" pageId="4" pageNumber="5">2 5%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FB41ABC4FAEC2799" box="[1243,1291,436,499]" gridcol="7" gridrow="4" pageId="4" pageNumber="5">1 3%</td>
|
|||
|
|
</tr>
|
|||
|
|
<tr id="C3CDAF163B21CBA6FF31A873FA612440" box="[171,1414,515,554]" gridrow="5" pageId="4" pageNumber="5">
|
|||
|
|
<th id="801CC66A3B21CBA6FF31A873FA612440" box="[171,1414,515,554]" colspan="9" colspanRight="8" gridcol="0" gridrow="5" pageId="4" pageNumber="5">Pie chart IIı Fig. 2 Table S6</th>
|
|||
|
|
</tr>
|
|||
|
|
<tr id="C3CDAF163B21CBA6FF31A84BFA612408" box="[171,1414,571,610]" gridrow="6" pageId="4" pageNumber="5" rowspan-8="1">
|
|||
|
|
<th id="801CC66A3B21CBA6FF31A84BFE2A2408" box="[171,461,571,610]" gridcol="0" gridrow="6" pageId="4" pageNumber="5">"3B-1" lag deposit at base of Jen-rex sand (MOR locality HC-312)</th>
|
|||
|
|
<td id="801CC66A3B21CBA6FD14A84BFD402408" box="[654,679,571,610]" gridcol="1" gridrow="6" pageId="4" pageNumber="5">n = %</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FD4AA84BFD162408" box="[720,753,571,610]" gridcol="2" gridrow="6" pageId="4" pageNumber="5">23 33%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FCB1A84BFCA82408" box="[811,847,571,610]" gridcol="3" gridrow="6" pageId="4" pageNumber="5">19 27%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FC0EA84BFC242408" box="[916,963,571,610]" gridcol="4" gridrow="6" pageId="4" pageNumber="5">18 26%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FB93A84BFBC82408" box="[1033,1071,571,610]" gridcol="5" gridrow="6" pageId="4" pageNumber="5">7 10%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FBE8A84BFB722408" box="[1138,1173,571,610]" gridcol="6" gridrow="6" pageId="4" pageNumber="5">2 3%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FB41A84BFAEC2408" box="[1243,1291,571,610]" gridcol="7" gridrow="6" pageId="4" pageNumber="5">1 1%</td>
|
|||
|
|
</tr>
|
|||
|
|
<tr id="C3CDAF163B21CBA6FF31A802FA6124F0" box="[171,1414,626,666]" gridrow="7" pageId="4" pageNumber="5">
|
|||
|
|
<th id="801CC66A3B21CBA6FF31A802FA6124F0" box="[171,1414,626,666]" colspan="9" colspanRight="8" gridcol="0" gridrow="7" pageId="4" pageNumber="5">Pie chart IIIı Fig. 2 Table S3</th>
|
|||
|
|
</tr>
|
|||
|
|
<tr id="C3CDAF163B21CBA6FF31A8DAFA6124B8" box="[171,1414,682,722]" gridrow="8" pageId="4" pageNumber="5" rowspan-7="1">
|
|||
|
|
<th id="801CC66A3B21CBA6FF31A8DAFE2A24B8" box="[171,461,682,722]" gridcol="0" gridrow="8" pageId="4" pageNumber="5">Lower Hell Creek Fm (L3) skeletons</th>
|
|||
|
|
<td id="801CC66A3B21CBA6FD14A8DAFD4024B8" box="[654,679,682,722]" gridcol="1" gridrow="8" pageId="4" pageNumber="5">n = %</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FD4AA8DAFD1624B8" box="[720,753,682,722]" gridcol="2" gridrow="8" pageId="4" pageNumber="5">11 28%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FCB1A8DAFCA824B8" box="[811,847,682,722]" gridcol="3" gridrow="8" pageId="4" pageNumber="5">11 28%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FC0EA8DAFC2424B8" box="[916,963,682,722]" gridcol="4" gridrow="8" pageId="4" pageNumber="5">6 15%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FB93A8DAFBC824B8" box="[1033,1071,682,722]" gridcol="5" gridrow="8" pageId="4" pageNumber="5">4 10%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FBE8A8DAFB7224B8" box="[1138,1173,682,722]" gridcol="6" gridrow="8" pageId="4" pageNumber="5">5 13%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FAC7A8DAFA6124B8" box="[1373,1414,682,722]" gridcol="8" gridrow="8" pageId="4" pageNumber="5">2 5%</td>
|
|||
|
|
</tr>
|
|||
|
|
<tr id="C3CDAF163B21CBA6FF31A892FA612563" box="[171,1414,738,777]" gridrow="9" pageId="4" pageNumber="5">
|
|||
|
|
<th id="801CC66A3B21CBA6FF31A892FA612563" box="[171,1414,738,777]" colspan="9" colspanRight="8" gridcol="0" gridrow="9" pageId="4" pageNumber="5">Pie chart IVı Fig. 2 Table S1</th>
|
|||
|
|
</tr>
|
|||
|
|
<tr id="C3CDAF163B21CBA6FF31A96AFA612532" box="[171,1414,794,856]" gridrow="10" pageId="4" pageNumber="5">
|
|||
|
|
<th id="801CC66A3B21CBA6FF31A96AFE2A2532" box="[171,461,794,856]" gridcol="0" gridrow="10" pageId="4" pageNumber="5">Totals for the entire Hell Creek Formation (see Figure 4)</th>
|
|||
|
|
<td id="801CC66A3B21CBA6FD14A96AFD402532" box="[654,679,794,856]" gridcol="1" gridrow="10" pageId="4" pageNumber="5">n = %</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FD4AA96AFD162532" box="[720,753,794,856]" gridcol="2" gridrow="10" pageId="4" pageNumber="5">73 40%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FCB1A96AFCA82532" box="[811,847,794,856]" gridcol="3" gridrow="10" pageId="4" pageNumber="5">44 24%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FC0EA96AFC242532" box="[916,963,794,856]" gridcol="4" gridrow="10" pageId="4" pageNumber="5">36 20%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FB93A96AFBC82532" box="[1033,1071,794,856]" gridcol="5" gridrow="10" pageId="4" pageNumber="5">15 8%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FBE8A96AFB722532" box="[1138,1173,794,856]" gridcol="6" gridrow="10" pageId="4" pageNumber="5">9 5%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FB41A96AFAEC2532" box="[1243,1291,794,856]" gridcol="7" gridrow="10" pageId="4" pageNumber="5">2 1%</td>
|
|||
|
|
<td id="801CC66A3B21CBA6FAC7A96AFA612532" box="[1373,1414,794,856]" gridcol="8" gridrow="10" pageId="4" pageNumber="5">2 1%</td>
|
|||
|
|
</tr>
|
|||
|
|
</table>
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B21345DFF31A901FE3C25AC" pageId="4" pageNumber="5">
|
|||
|
|
<tableNote id="801BACDA3B21345DFF31A901FE3C25AC" pageId="4" pageNumber="5" targetBox="[171,1414,252,856]" targetPageId="4">
|
|||
|
|
Values determined from the dinosaur census tables (Tables S1ı S2ı S3ı S4ı S5ı S6). Empty cell indicates no record for that taxon. See Figure 2 for detailed stratigraphic section of the Hell Creek Formation and corresponding pie chart showing relative abundance of dinosaur genera. Abbreviations:
|
|||
|
|
<emphasis id="4F8971463B21345DFB3DA9F8FBF825DA" italics="true" pageId="4" pageNumber="5">
|
|||
|
|
Tricı
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFB51A9F8FAFA25F3" authorityName="Marsh" authorityYear="1889" box="[1227,1309,904,921]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">Triceratops</taxonomicName>
|
|||
|
|
; Tyrnı
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFAC9A9F8FA2625F3" box="[1363,1473,904,921]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
|||
|
|
; Edmnı
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFF7AA9EFFEB325DA" authorityName="Lambe" authorityYear="1917" box="[224,340,927,944]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">Edmontosaurus</taxonomicName>
|
|||
|
|
; Thesı
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFE16A9EFFE1B25DA" authorityName="Gilmore" authorityYear="1913" box="[396,508,927,944]" class="Reptilia" family="Thescelosauridae" genus="Thescelosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">Thescelosaurus</taxonomicName>
|
|||
|
|
; Orniı
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFDA9A9EFFD7925DA" authorityName="Marsh" authorityYear="1890" box="[563,670,927,944]" class="Reptilia" family="Ornithomimidae" genus="Ornithomimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">Ornithomimus</taxonomicName>
|
|||
|
|
; Pachyı
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFD7BA9EFFC9D25DA" authorityName="Brown & Schlaikjer" authorityYear="1943" box="[737,890,927,944]" class="Reptilia" family="Pachycephalosauridae" genus="Pachycephalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">Pachycephalosaurus</taxonomicName>
|
|||
|
|
; Ankyı
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFC2FA9EFFBFC25DA" authorityName="Brown" authorityYear="1908" box="[949,1051,927,944]" class="Reptilia" family="Ankylosauridae" genus="Ankylosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">Ankylosaurus</taxonomicName>
|
|||
|
|
.
|
|||
|
|
</emphasis>
|
|||
|
|
doi:10.1371/journal.pone.0016574.t001
|
|||
|
|
</tableNote>
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B21345DFF29AE84FEBC2029" blockId="4.[155,794,1012,1717]" pageId="4" pageNumber="5">
|
|||
|
|
Teeth were not collected or annotated because of the difficulties in using them for ontogenetic assessment with the exception of two large
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFF49AF5DFEA92329" box="[211,334,1325,1347]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFF49AF5DFEA92329" box="[211,334,1325,1347]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
teeth from the ‘‘3B-1 Lag’’ at the base of the Jen-rex sand. Tooth size varies as much as 300% in a single jaw, particularly in hadrosaurids (MOR 1609, Becky’s Giant), ceratopsids (MOR 2574, Quittin Time) and tyrannosaurids (
|
|||
|
|
<materialsCitation id="CD95A7093B21345DFD44AFF3FEE823DF" ID-GBIF-Occurrence="3336525303" collectionCode="MOR" country="United States" formation="Hell Creek Formation" pageId="4" pageNumber="5" specimenCode="MOR 1125" stateProvince="Montana">MOR 1125, B-rex</materialsCitation>
|
|||
|
|
). This is one reason the assignment of some dinosaur teeth to ‘‘babies’’ [19] may be incorrect. These teeth are more accurately interpreted as being derived from the anterior or posterior portions of jaws from older individuals (
|
|||
|
|
<figureCitation id="E5C6B1D13B21345DFD18AF84FD342060" box="[642,723,1524,1546]" captionStart="Figure 3" captionStartId="5.[840,905,172,191]" captionTargetBox="[160,768,170,1904]" captionTargetId="figure@5.[155,776,162,1911]" captionTargetPageId="5" captionText="Figure 3. Tyrannosaurus (MOR 1125ı ‘‘B-rex’’) teeth from the lower jaw of this medium-sized skeleton illustrate the extreme range in overall tooth size within one individual. A. A smaller posterior tooth from position #14 from the front of the jaw. B. A larger tooth from position #4 in the same jaw. This demonstrates why shed dinosaur teeth are not a reliable indicator of relative skeletal size and ontogenetic age. doi:10.1371/journal.pone.0016574.g003" figureDoi="http://doi.org/10.5281/zenodo.3744922" httpUri="https://zenodo.org/record/3744922/files/figure.png" pageId="4" pageNumber="5">Figure 3</figureCitation>
|
|||
|
|
). Only the largest and most robust tyrannosaurid teeth are reliable indicators of adults.
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B21345DFF29AC3AFF1120DF" blockId="4.[155,794,1012,1717]" pageId="4" pageNumber="5">
|
|||
|
|
The
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFF70AC3AFEA12035" authorityName="Marsh" authorityYear="1889" box="[234,326,1610,1631]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFF70AC3AFEA12035" box="[234,326,1610,1631]" italics="true" pageId="4" pageNumber="5">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
specimens recorded in Table S6 represent specimens that were collected, but remain unprepared, uncataloged and consist of an unknown number of disarticulated elements.
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B21345DFF01ACAFFE9A209D" blockId="4.[155,794,1759,1955]" box="[155,381,1759,1783]" pageId="4" pageNumber="5">
|
|||
|
|
<heading id="260A1A383B21345DFF01ACAFFE9A209D" box="[155,381,1759,1783]" fontSize="10" level="2" pageId="4" pageNumber="5" reason="1">Ontogenetic Results</heading>
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B21345DFF29AC8EFB502360" blockId="4.[155,794,1759,1955]" lastBlockId="4.[840,1478,1012,1290]" pageId="4" pageNumber="5">
|
|||
|
|
In this census, growth stages at either end of the dinosaurian ontogenetic spectrum are least represented. Specimens of both the smallest and presumably youngest juveniles and the largest, and presumably oldest adults are the most rare dinosaurs recorded. The smallest specimen of
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFE03AD00FE1221EF" authorityName="Marsh" authorityYear="1889" box="[409,501,1904,1925]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFE03AD00FE1221EF" box="[409,501,1904,1925]" italics="true" pageId="4" pageNumber="5">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
found during this project is a partially complete skull that is half again the length of the smallest previously known skull [20]. None of the
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFB66A984FABF2260" authorityName="Marsh" authorityYear="1889" box="[1276,1368,1012,1034]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFB66A984FABF2260" box="[1276,1368,1012,1034]" italics="true" pageId="4" pageNumber="5">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
specimens found in the census area could be positively identified as ‘‘
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFCCEAE5CFC572228" authorityName="Marsh" authorityYear="1891" box="[852,944,1068,1090]" class="Reptilia" family="Ceratopsidae" genus="Torosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFCCEAE5CFC572228" box="[852,944,1068,1090]" italics="true" pageId="4" pageNumber="5">Torosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
’’ size, although the specimen collected from the ‘‘BAB’’ locality has elongated squamosals characteristic of the ‘‘
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFCCEAE15FC572211" authorityName="Marsh" authorityYear="1891" box="[852,944,1125,1147]" class="Reptilia" family="Ceratopsidae" genus="Torosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFCCEAE15FC572211" box="[852,944,1125,1147]" italics="true" pageId="4" pageNumber="5">Torosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
’’ morph. Two specimens of
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFB43AE15FAB12211" authorityName="Lambe" authorityYear="1917" box="[1241,1366,1125,1147]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFB43AE15FAB12211" box="[1241,1366,1125,1147]" italics="true" pageId="4" pageNumber="5">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
are in the ‘‘XL’’ size range: ‘‘Becky’s Giant’’ (MOR 1609) is a maxilla with a tooth-row length of 570 mm and the tail of ‘‘X-rex’’ (MOR 1142) is 7.5 meters in length from the posterior end of the sacrum. Both these specimens are indicative of greater size ranges then previously attributed to
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFBA9AE83FB572363" authorityName="Lambe" authorityYear="1917" box="[1075,1200,1267,1289]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFBA9AE83FB572363" box="[1075,1200,1267,1289]" italics="true" pageId="4" pageNumber="5">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
.
|
|||
|
|
</paragraph>
|
|||
|
|
</subSubSection>
|
|||
|
|
<subSubSection id="35E7FEDF3B21345FFCD2AF4CFC9022BE" lastPageId="6" lastPageNumber="7" pageId="4" pageNumber="5" type="discussion">
|
|||
|
|
<paragraph id="7D42AD543B21345DFCD2AF4CFC37233E" blockId="4.[840,976,1340,1364]" box="[840,976,1340,1364]" pageId="4" pageNumber="5">
|
|||
|
|
<heading id="260A1A383B21345DFCD2AF4CFC37233E" bold="true" box="[840,976,1340,1364]" fontSize="10" level="1" pageId="4" pageNumber="5" reason="0">
|
|||
|
|
<emphasis id="4F8971463B21345DFCD2AF4CFC37233E" bold="true" box="[840,976,1340,1364]" pageId="4" pageNumber="5">Discussion</emphasis>
|
|||
|
|
</heading>
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B21345DFCD2AF1DFC7023EF" blockId="4.[840,1478,1389,1954]" box="[840,919,1389,1413]" pageId="4" pageNumber="5">
|
|||
|
|
<heading id="260A1A383B21345DFCD2AF1DFC7023EF" box="[840,919,1389,1413]" fontSize="10" level="2" pageId="4" pageNumber="5" reason="1">Census</heading>
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B21345DFCFAAFFCFAA621C8" blockId="4.[840,1478,1389,1954]" pageId="4" pageNumber="5">
|
|||
|
|
The dinosaur collections made over the past decade during the Hell Creek Project yielded new information from an improved genus-level collecting schema and robust data set that revealed relative dinosaur abundances that were unexpected, and ontogenetic age classes previously considered rare. We recognize a much higher percentage of
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFB8CAC6AFB762045" box="[1046,1169,1562,1583]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFB8CAC6AFB762045" box="[1046,1169,1562,1583]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
(
|
|||
|
|
<tableCitation id="307F98EF3B21345DFB04AC6AFB01205A" box="[1182,1254,1562,1584]" captionStart="Table 1" captionStartId="4.[171,235,174,195]" captionTargetBox="[171,1414,252,856]" captionTargetId="graphics@4.[163,1478,151,962]" captionTargetPageId="4" captionText="Table 1. Hell Creek Formation dinosaur census." httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="4" pageNumber="5" tableUuid="2982FDDC3B21345DFF31AADEFD7F26A9">Table 1</tableCitation>
|
|||
|
|
) than previous surveys [3,4,21].
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFC33AC47FBC32026" box="[937,1060,1591,1612]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFC33AC47FBC32026" box="[937,1060,1591,1612]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
equals
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFBE7AC46FB1D2021" authorityName="Lambe" authorityYear="1917" box="[1149,1274,1590,1611]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFBE7AC46FB1D2021" box="[1149,1274,1590,1611]" italics="true" pageId="4" pageNumber="5">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
in U3 and in L3 comprises a greater percentage of the large dinosaur fauna as the second most abundant taxon after
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFB4FAC1FFAD620EE" authorityName="Marsh" authorityYear="1889" box="[1237,1329,1647,1668]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFB4FAC1FFAD620EE" box="[1237,1329,1647,1668]" italics="true" pageId="4" pageNumber="5">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
, followed by
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFCD2ACFBFC2220CA" authorityName="Lambe" authorityYear="1917" box="[840,965,1675,1696]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFCD2ACFBFC2220CA" box="[840,965,1675,1696]" italics="true" pageId="4" pageNumber="5">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
. This is surprisingly consistent in (1) the two major lag deposits (MOR loc. HC-530 and HC-312) in the Apex sandstone and Jen-rex sand (
|
|||
|
|
<figureCitation id="E5C6B1D13B21345DFBC0ACB5FB4F20B1" box="[1114,1192,1733,1755]" captionStart="Figure 2" captionStartId="2.[155,220,1221,1240]" captionTargetBox="[160,1012,169,1180]" captionTargetId="figure@2.[155,1017,162,1187]" captionTargetPageId="2" captionText="Figure 2. Stratigraphic divisions of the Hell Creek Formation with stratigraphic details of the upper third (U3) and lower third (L3) sequences referenced in the text and the associated pie chart showing dinosaur abundance. Pie charts I – IV reflect the relative abundance of dinosaur genera based on skeletons (charts I and IV) and individual bones (charts II and III). Tyrannosaurus skeletons are as abundant as the hadrosaurid Edmontosaurus, an herbivore, in the upper Hell Creek Formation and nearly twice as common as Edmontosaurus in the lower Hell Creek Formation. Individual bones of Tyrannosaurus and Edmontosaurus are found approximately in the same order of magnitude within the lag deposits. Key: Pie chart I, dinosaur skeletons in the upper Hell Creek Formation (U3); Pie chart II, dinosaur bones from the ‘‘Doldrum’s’’ lag deposit at the base of the Apex Sandstone (AS); Pie chart III, dinosaur bones from the ‘‘3B-1’’ lag deposit at the base of the Jen-rex Sand; and Pie chart IV, dinosaur skeletons from the lower Hell Creek Formation (L3). Abbreviations: L3lBS, lower basal sandstone; L3.mBS, middle basal sandstone; L3.uBS, upper basal sandstone; L3.lMS, lower mudstone; L3.mMS, middle mudstone; L3.uMS, upper mudstone; U3.AS, Apex sandstone; U3.ibMS, interbedded mudstone; U3.10mS, 10 meter sandstone; U3.uMS, upper mudstone; U3.CZ, coal zone. doi:10.1371/journal.pone.0016574.g002" figureDoi="http://doi.org/10.5281/zenodo.3744920" httpUri="https://zenodo.org/record/3744920/files/figure.png" pageId="4" pageNumber="5">Figure 2</figureCitation>
|
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|
|
) where individual bones were counted and (2) in two-thirds of the formation reflected in L3 and U3 records of dinosaur skeletons only. Measured throughout the entire formation,
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFC6BAD6AFBAA2145" authorityName="Marsh" authorityYear="1889" box="[1009,1101,1818,1839]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFC6BAD6AFBAA2145" box="[1009,1101,1818,1839]" italics="true" pageId="4" pageNumber="5">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
is by far the most common dinosaur at 40% (n = 72),
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFC55AD46FBAD2121" box="[975,1098,1846,1867]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFC55AD46FBAD2121" box="[975,1098,1846,1867]" italics="true" pageId="4" pageNumber="5">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
is second at 24% (n = 44),
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFAD2AD46FA222121" authorityName="Lambe" authorityYear="1917" box="[1352,1477,1846,1867]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
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|
|
<emphasis id="4F8971463B21345DFAD2AD46FA222121" box="[1352,1477,1846,1867]" italics="true" pageId="4" pageNumber="5">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
is third at 20% (n = 36), followed by
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFB29AD22FACA210D" authorityName="Gilmore" authorityYear="1913" box="[1203,1325,1874,1895]" class="Reptilia" family="Thescelosauridae" genus="Thescelosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFB29AD22FACA210D" box="[1203,1325,1874,1895]" italics="true" pageId="4" pageNumber="5">Thescelosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
at 8% (n = 15),
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFCD2AD1FFC5A21EE" authorityName="Marsh" authorityYear="1890" box="[840,957,1903,1924]" class="Reptilia" family="Ornithomimidae" genus="Ornithomimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFCD2AD1FFC5A21EE" box="[840,957,1903,1924]" italics="true" pageId="4" pageNumber="5">Ornithomimus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
at 5% (n = 9), and
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFB18AD1FFAC421EE" authorityName="Brown & Schlaikjer" authorityYear="1943" box="[1154,1315,1903,1924]" class="Reptilia" family="Pachycephalosauridae" genus="Pachycephalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFB18AD1FFAC421EE" box="[1154,1315,1903,1924]" italics="true" pageId="4" pageNumber="5">Pachycephalosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
and
|
|||
|
|
<taxonomicName id="BAFDD6D73B21345DFAC3AD1FFA2221EE" authorityName="Brown" authorityYear="1908" box="[1369,1477,1903,1924]" class="Reptilia" family="Ankylosauridae" genus="Ankylosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="5" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B21345DFAC3AD1FFA2221EE" box="[1369,1477,1903,1924]" italics="true" pageId="4" pageNumber="5">Ankylosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
both at 1% (n = 2) are relatively rare (see
|
|||
|
|
<figureCitation id="E5C6B1D13B21345DFB78ADFCFAD221C8" box="[1250,1333,1932,1954]" captionStart="Figure 4" captionStartId="6.[155,220,997,1016]" captionTargetBox="[161,724,167,959]" captionTargetId="figure@6.[155,730,162,964]" captionTargetPageId="6" captionText="Figure 4. Pie chart of the time averaged census for largebodied dinosaurs from the entire Hell Creek Formation in the study area. Triceratops is the most common dinosaur at 40% (n = 72); Tyrannosaurus is second at 24% (n = 44); Edmontosaurus is third at 20% (n = 36) followed by Thescelosaurus at 8% (n = 15)ı Ornithomimus at 5% (n = 9)ı and Pachycephalosaurus and Ankylosaurus at 1% (n = 2). doi:10.1371/journal.pone.0016574.g004" figureDoi="http://doi.org/10.5281/zenodo.3744924" httpUri="https://zenodo.org/record/3744924/files/figure.png" pageId="4" pageNumber="5">Figure 4</figureCitation>
|
|||
|
|
).
|
|||
|
|
</paragraph>
|
|||
|
|
<caption id="2982FDDC3B20345CFCD2AADCFB4F270C" ID-DOI="http://doi.org/10.5281/zenodo.3744922" ID-Zenodo-Dep="3744922" httpUri="https://zenodo.org/record/3744922/files/figure.png" pageId="5" pageNumber="6" startId="5.[840,905,172,191]" targetBox="[160,768,170,1904]" targetPageId="5">
|
|||
|
|
<paragraph id="7D42AD543B20345CFCD2AADCFB4F270C" blockId="5.[840,1478,172,358]" pageId="5" pageNumber="6">
|
|||
|
|
<emphasis id="4F8971463B20345CFCD2AADCFA8E2684" bold="true" pageId="5" pageNumber="6">
|
|||
|
|
Figure 3.
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFC2BAADCFBDE26D5" box="[945,1081,172,191]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">Tyrannosaurus</taxonomicName>
|
|||
|
|
(
|
|||
|
|
<materialsCitation id="CD95A7093B20345CFBD0AADCFAF226D5" ID-GBIF-Occurrence="3336525304" box="[1098,1301,172,191]" collectionCode="MOR" country="United States" formation="Hell Creek Formation" pageId="5" pageNumber="6" specimenCode="MOR 1125" stateProvince="Montana">MOR 1125, ‘‘B-rex’</materialsCitation>
|
|||
|
|
’) teeth from the lower jaw of this medium-sized skeleton illustrate the extreme range in overall tooth size within one individual. A.
|
|||
|
|
</emphasis>
|
|||
|
|
A smaller posterior tooth from position #14 from the front of the jaw.
|
|||
|
|
<emphasis id="4F8971463B20345CFAFDAA83FA9C276C" bold="true" box="[1383,1403,243,262]" pageId="5" pageNumber="6">B.</emphasis>
|
|||
|
|
A larger tooth from position #4 in the same jaw. This demonstrates why shed dinosaur teeth are not a reliable indicator of relative skeletal size and ontogenetic age. doi:10.1371/journal.pone.0016574.g003
|
|||
|
|
</paragraph>
|
|||
|
|
</caption>
|
|||
|
|
<paragraph id="7D42AD543B20345CFCFAABF4FB7E2548" blockId="5.[840,1478,387,802]" pageId="5" pageNumber="6">
|
|||
|
|
Even though
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFC63ABF3FBB227F3" authorityName="Marsh" authorityYear="1889" box="[1017,1109,387,409]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B20345CFC63ABF3FBB227F3" box="[1017,1109,387,409]" italics="true" pageId="5" pageNumber="6">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
dominates this census, associated specimens of
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFC57ABEFFBCE27DF" authorityName="Marsh" authorityYear="1889" box="[973,1065,415,437]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B20345CFC57ABEFFBCE27DF" box="[973,1065,415,437]" italics="true" pageId="5" pageNumber="6">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
consisting of both cranial and postcranial elements remain relatively rare (see
|
|||
|
|
<tableCitation id="307F98EF3B20345CFB3FABCCFAE027B8" box="[1189,1287,444,466]" captionStart="Table 1" captionStartId="4.[171,235,174,195]" captionTargetBox="[171,1414,252,856]" captionTargetId="graphics@4.[163,1478,151,962]" captionTargetPageId="4" captionText="Table 1. Hell Creek Formation dinosaur census." httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="5" pageNumber="6" tableUuid="2982FDDC3B21345DFF31AADEFD7F26A9">Tables S1</tableCitation>
|
|||
|
|
, S2). This contrasts with the record of isolated skulls that contribute to a significant portion of this census. We propose that this inconsistency may be explained by a historical collecting bias influenced by taphonomic controls. This is documented in museum collections [18]. Alternatively, predation, scavenging, or some as yet unknown vital effect of rapid deterioration of
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFB33A813FAE22413" authorityName="Marsh" authorityYear="1889" box="[1193,1285,611,633]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B20345CFB33A813FAE22413" box="[1193,1285,611,633]" italics="true" pageId="5" pageNumber="6">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
limb elements may limit their preservation in the fossil record. We observed that postcranial elements are often located at some distance from the associated skull, particularly in the preservation of
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFAFEA8C7FA2724A7" authorityName="Marsh" authorityYear="1889" box="[1380,1472,695,717]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B20345CFAFEA8C7FA2724A7" box="[1380,1472,695,717]" italics="true" pageId="5" pageNumber="6">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
. Thus, the limited discovery of postcranial elements may, in some circumstances, simply depend on how extensive a quarry is expanded after a skull is collected.
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B20345CFCD2A936FBC12534" blockId="5.[840,1479,838,1956]" box="[840,1062,838,862]" pageId="5" pageNumber="6">
|
|||
|
|
<heading id="260A1A383B20345CFCD2A936FBC12534" box="[840,1062,838,862]" fontSize="10" level="2" pageId="5" pageNumber="6" reason="1">Ontogenetic Stages</heading>
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B20345CFCFAA915FB7B2375" blockId="5.[840,1479,838,1956]" pageId="5" pageNumber="6">
|
|||
|
|
When ontogenetic stages are considered, we observe a low number of both ‘‘A’’ and ‘‘F’’ class (see ontogeny column in
|
|||
|
|
<tableCitation id="307F98EF3B20345CFA1FA9F1FC8525D9" captionStart="Table 1" captionStartId="4.[171,235,174,195]" captionTargetBox="[171,1414,252,856]" captionTargetId="graphics@4.[163,1478,151,962]" captionTargetPageId="4" captionText="Table 1. Hell Creek Formation dinosaur census." httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="5" pageNumber="6" tableUuid="2982FDDC3B21345DFF31AADEFD7F26A9">Tables S1</tableCitation>
|
|||
|
|
, S3, S4, S6) of
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFB83A9EDFB9225D9" authorityName="Marsh" authorityYear="1889" box="[1049,1141,925,947]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B20345CFB83A9EDFB9225D9" box="[1049,1141,925,947]" italics="true" pageId="5" pageNumber="6">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
individuals and ‘‘S’’ and ‘‘XL’’ individuals of other taxa. Overall, the dinosaur assemblages represented in the Hell Creek Formation consist primarily of subadult or small adult size individuals (based on comparisons with the largest specimens of known taxa). Small juveniles and large adults are both extremely rare, whereas subadult individuals (M & L and D & E) are relatively common. The paucity of juveniles seen in the Hell Creek Formation and contemporaneous sediments puzzled earlier researchers [22]. This can likely be explained by a combination of: (1) extended parental care [23–25]; (2) rapid juvenile growth [26,27]; and (3) colonial nesting in select geographic environments [19,28]. This pattern likely reflects either a preservational (taphonomic) or life history consequence acting on the dinosaur population.
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B20345CFCFAAF56FBD921CE" blockId="5.[840,1479,838,1956]" pageId="5" pageNumber="6">
|
|||
|
|
The uncommonness of apparently fully mature adults is more mysterious and not easily explained. What is now apparent, however, is this pattern contributed to an historical increase in the naming of new dinosaur species from the Hell Creek Formation. For example, over many decades it was presumed that the taxon ‘‘
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFCCEAFC2FC5723AD" authorityName="Marsh" authorityYear="1891" box="[852,944,1458,1479]" class="Reptilia" family="Ceratopsidae" genus="Torosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B20345CFCCEAFC2FC5723AD" box="[852,944,1458,1479]" italics="true" pageId="5" pageNumber="6">Torosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
’’ represented a horned dinosaur that reached enormous proportions, even though there were no reported juveniles in the literature. The relatively expanded and fenestrated parietosquamosal frill exhibited by ‘‘
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFBE1AC76FB302071" authorityName="Marsh" authorityYear="1891" box="[1147,1239,1542,1563]" class="Reptilia" family="Ceratopsidae" genus="Torosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B20345CFBE1AC76FB302071" box="[1147,1239,1542,1563]" italics="true" pageId="5" pageNumber="6">Torosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
’’ was among its most significant features [29]. With the advent of studies employing ontogenetic osteohistology, the alternative hypothesis that these giant dinosaurs were more likely mature individuals of existing taxa, rather than distinct taxa, became evident. This hypothesis is exemplified in recent studies of
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFB12ACE2FB0320CD" authorityName="Marsh" authorityYear="1889" box="[1160,1252,1682,1703]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B20345CFB12ACE2FB0320CD" box="[1160,1252,1682,1703]" italics="true" pageId="5" pageNumber="6">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
ontogeny [12,30] that reinterpret ‘‘
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFC5FACDEFBC620A9" authorityName="Marsh" authorityYear="1891" box="[965,1057,1710,1731]" class="Reptilia" family="Ceratopsidae" genus="Torosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B20345CFC5FACDEFBC620A9" box="[965,1057,1710,1731]" italics="true" pageId="5" pageNumber="6">Torosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
’’ as an adult
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFB2EACDEFAF720A9" authorityName="Marsh" authorityYear="1889" box="[1204,1296,1710,1731]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B20345CFB2EACDEFAF720A9" box="[1204,1296,1710,1731]" italics="true" pageId="5" pageNumber="6">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
. Nonetheless, this hypothesis fails to explain why these giant, mature individuals are so rare, or more explicitly, why most
|
|||
|
|
<taxonomicName id="BAFDD6D73B20345CFB49AC96FAC82091" authorityName="Marsh" authorityYear="1889" box="[1235,1327,1766,1787]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="5" pageNumber="6" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B20345CFB49AC96FAC82091" box="[1235,1327,1766,1787]" italics="true" pageId="5" pageNumber="6">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
specimens are subadult sized. We propose that mature individuals of at least some dinosaur taxa either lived in a separate geographic locale analogous to younger individuals inhabiting an upland fauna, or these taxa experienced high mortality rates before reaching terminal size where late stage and often extreme cranial morphology is expressed.
|
|||
|
|
</paragraph>
|
|||
|
|
<caption id="2982FDDC3B23345FFF01A995FE1D22E2" ID-DOI="http://doi.org/10.5281/zenodo.3744924" ID-Zenodo-Dep="3744924" httpUri="https://zenodo.org/record/3744924/files/figure.png" pageId="6" pageNumber="7" startId="6.[155,220,997,1016]" targetBox="[161,724,167,959]" targetPageId="6">
|
|||
|
|
<paragraph id="7D42AD543B23345FFF01A995FE1D22E2" blockId="6.[155,794,997,1160]" pageId="6" pageNumber="7">
|
|||
|
|
<emphasis id="4F8971463B23345FFF01A995FEEA2242" bold="true" pageId="6" pageNumber="7">Figure 4. Pie chart of the time averaged census for largebodied dinosaurs from the entire Hell Creek Formation in the study area.</emphasis>
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFE89AE65FE952242" authorityName="Marsh" authorityYear="1889" box="[275,370,1045,1064]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFE89AE65FE952242" box="[275,370,1045,1064]" italics="true" pageId="6" pageNumber="7">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
is the most common dinosaur at 40% (n = 72);
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFF01AE5DFEFD222A" box="[155,282,1069,1088]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFF01AE5DFEFD222A" box="[155,282,1069,1088]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
is second at 24% (n = 44);
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFD90AE5DFD77222A" authorityName="Lambe" authorityYear="1917" box="[522,656,1069,1088]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFD90AE5DFD77222A" box="[522,656,1069,1088]" italics="true" pageId="6" pageNumber="7">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
is third at 20% (n = 36) followed by
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFECEAE35FE322232" authorityName="Gilmore" authorityYear="1913" box="[340,469,1093,1112]" class="Reptilia" family="Thescelosauridae" genus="Thescelosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFECEAE35FE322232" box="[340,469,1093,1112]" italics="true" pageId="6" pageNumber="7">Thescelosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
at 8% (n = 15)ı
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFDF8AE35FD382232" authorityName="Marsh" authorityYear="1890" box="[610,735,1093,1112]" class="Reptilia" family="Ornithomimidae" genus="Ornithomimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFDF8AE35FD382232" box="[610,735,1093,1112]" italics="true" pageId="6" pageNumber="7">Ornithomimus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
at 5% (n = 9)ı and
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFE9FAE2DFE53221A" authorityName="Brown & Schlaikjer" authorityYear="1943" box="[261,436,1117,1136]" class="Reptilia" family="Pachycephalosauridae" genus="Pachycephalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFE9FAE2DFE53221A" box="[261,436,1117,1136]" italics="true" pageId="6" pageNumber="7">Pachycephalosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
and
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFE7FAE2DFDB0221A" authorityName="Brown" authorityYear="1908" box="[485,599,1117,1136]" class="Reptilia" family="Ankylosauridae" genus="Ankylosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFE7FAE2DFDB0221A" box="[485,599,1117,1136]" italics="true" pageId="6" pageNumber="7">Ankylosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
at 1% (n = 2). doi:10.1371/journal.pone.0016574.g004
|
|||
|
|
</paragraph>
|
|||
|
|
</caption>
|
|||
|
|
<paragraph id="7D42AD543B23345FFF29AEBBFEDF21CE" blockId="6.[155,793,1227,1956]" pageId="6" pageNumber="7">
|
|||
|
|
Reproductive maturity in some dinosaurs was achieved during subadulthood (e.g.,
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFEC1AE96FE302296" box="[347,471,1254,1276]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFEC1AE96FE302296" box="[347,471,1254,1276]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
,
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFE7FAE96FDDB2296" authorityName="Marsh" authorityYear="1877" box="[485,572,1254,1276]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFE7FAE96FDDB2296" box="[485,572,1254,1276]" italics="true" pageId="6" pageNumber="7">Allosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
and
|
|||
|
|
<emphasis id="4F8971463B23345FFDF5AE96FD002296" box="[623,743,1254,1276]" italics="true" pageId="6" pageNumber="7">Tenontosaurus</emphasis>
|
|||
|
|
) and this event led to high adult mortality [31]. Interestingly however, our census data indicate the highest mortality occurred when
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFF01AF4BFF1F233B" authorityName="Marsh" authorityYear="1889" box="[155,248,1339,1361]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFF01AF4BFF1F233B" box="[155,248,1339,1361]" italics="true" pageId="6" pageNumber="7">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
was about 2/3 grown (= skulls approximately 2.0 m in length compared to adults with 3.0 m long skulls) prior to the final ontogenetic stage of frill expansion and fenestration in
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFF01AFE1FF1F23CC" authorityName="Marsh" authorityYear="1889" box="[155,248,1425,1446]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFF01AFE1FF1F23CC" box="[155,248,1425,1446]" italics="true" pageId="6" pageNumber="7">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
(= ‘‘
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFEB4AFE1FE6C23CC" authorityName="Marsh" authorityYear="1891" box="[302,395,1425,1446]" class="Reptilia" family="Ceratopsidae" genus="Torosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFEB4AFE1FE6C23CC" box="[302,395,1425,1446]" italics="true" pageId="6" pageNumber="7">Torosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
’’).
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFE2AAFE0FDC823CF" authorityName="Lambe" authorityYear="1917" box="[432,559,1424,1445]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFE2AAFE0FDC823CF" box="[432,559,1424,1445]" italics="true" pageId="6" pageNumber="7">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
conforms to a similar scenario where the ‘‘XL’’ size individuals are the most rare, and the mid-size (‘‘M’’ and ‘‘L’’) individuals are the most common. This pattern is difficult to evaluate in
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFDB2AF96FD432391" box="[552,676,1510,1531]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFDB2AF96FD432391" box="[552,676,1510,1531]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
because of apparent variations in age relative to size [17]. Nonetheless, we predict a larger specimen of
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFE26AC6EFDDF2059" box="[444,568,1566,1587]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFE26AC6EFDDF2059" box="[444,568,1566,1587]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
than currently known will likely be discovered in future field studies. Although the lines of arrested growth (LAGs) observed in the largest yet known
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFF01AC03FEF020E2" box="[155,279,1651,1672]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFF01AC03FEF020E2" box="[155,279,1651,1672]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
specimens [17] suggest slowed growth, and therefore a presumed nearing of maturity, the cortex tissues of the femora and tibia of these individuals remain mostly primary. This contrasts with the femoral and tibial cortex tissues of the largest individuals of
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFEF7AC94FE2D2093" authorityName="Marsh" authorityYear="1889" box="[365,458,1764,1785]" class="Reptilia" family="Ceratopsidae" genus="Triceratops" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFEF7AC94FE2D2093" box="[365,458,1764,1785]" italics="true" pageId="6" pageNumber="7">Triceratops</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
and
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFE64AC94FD9A2093" authorityName="Lambe" authorityYear="1917" box="[510,637,1764,1785]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFE64AC94FD9A2093" box="[510,637,1764,1785]" italics="true" pageId="6" pageNumber="7">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
that are mostly secondary (dense Haversian), which is a much more mature form of cortical tissue. This suggests that
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFD93AD6DFD622158" box="[521,645,1821,1842]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFD93AD6DFD622158" box="[521,645,1821,1842]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
growth would have continued, resulting in a bulking-up of the skeleton by continued additions of periosteal bone tissues, possibly to the external fundamental system (EFS), which signifies maturity in other taxa [26].
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B23345FFCD2AAD6FB9626D4" blockId="6.[840,1479,166,1236]" box="[840,1137,166,190]" pageId="6" pageNumber="7">
|
|||
|
|
<heading id="260A1A383B23345FFCD2AAD6FB9626D4" box="[840,1137,166,190]" fontSize="10" level="2" pageId="6" pageNumber="7" reason="1">
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFCD2AAD6FC0026D4" authority="Abundance" box="[840,999,166,190]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFCD2AAD6FC0026D4" box="[840,999,166,190]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
Abundance
|
|||
|
|
</heading>
|
|||
|
|
</paragraph>
|
|||
|
|
<paragraph id="7D42AD543B23345FFCFAAAB5FC9022BE" blockId="6.[840,1479,166,1236]" pageId="6" pageNumber="7">
|
|||
|
|
The abundance of
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFBBCAAB5FB4526B1" box="[1062,1186,197,219]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFBBCAAB5FB4526B1" box="[1062,1186,197,219]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
specimens both as skeletons and as isolated elements in the LAG deposits contradicts hypotheses concerning predator-prey ratios expected for large, predatory terrestrial animals such as tyrannosaurids [32,33]. Although constant ratios are suspect in modern ecosystems [34,35], there are always at least 75% more non-predators than predators, and in mammal populations the ratio is>90% [32 and references therein]. What is particularly interesting in this census is the indication that
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFB85ABD7FB7C27D7" box="[1055,1179,423,445]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFB85ABD7FB7C27D7" box="[1055,1179,423,445]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
is at least as abundant in the upper Hell Creek Formation as
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFB32ABB3FAC027B3" authorityName="Lambe" authorityYear="1917" box="[1192,1319,451,473]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFB32ABB3FAC027B3" box="[1192,1319,451,473]" italics="true" pageId="6" pageNumber="7">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
, an herbivore, previously suggested to be the primary food source of
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFAEDABAFFC9A247B" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFAEDABAFFC9A247B" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
[36] (
|
|||
|
|
<figureCitation id="E5C6B1D13B23345FFC54AB8CFBCC2478" box="[974,1067,508,530]" captionStart="Figure 2" captionStartId="2.[155,220,1221,1240]" captionTargetBox="[160,1012,169,1180]" captionTargetId="figure@2.[155,1017,162,1187]" captionTargetPageId="2" captionText="Figure 2. Stratigraphic divisions of the Hell Creek Formation with stratigraphic details of the upper third (U3) and lower third (L3) sequences referenced in the text and the associated pie chart showing dinosaur abundance. Pie charts I – IV reflect the relative abundance of dinosaur genera based on skeletons (charts I and IV) and individual bones (charts II and III). Tyrannosaurus skeletons are as abundant as the hadrosaurid Edmontosaurus, an herbivore, in the upper Hell Creek Formation and nearly twice as common as Edmontosaurus in the lower Hell Creek Formation. Individual bones of Tyrannosaurus and Edmontosaurus are found approximately in the same order of magnitude within the lag deposits. Key: Pie chart I, dinosaur skeletons in the upper Hell Creek Formation (U3); Pie chart II, dinosaur bones from the ‘‘Doldrum’s’’ lag deposit at the base of the Apex Sandstone (AS); Pie chart III, dinosaur bones from the ‘‘3B-1’’ lag deposit at the base of the Jen-rex Sand; and Pie chart IV, dinosaur skeletons from the lower Hell Creek Formation (L3). Abbreviations: L3lBS, lower basal sandstone; L3.mBS, middle basal sandstone; L3.uBS, upper basal sandstone; L3.lMS, lower mudstone; L3.mMS, middle mudstone; L3.uMS, upper mudstone; U3.AS, Apex sandstone; U3.ibMS, interbedded mudstone; U3.10mS, 10 meter sandstone; U3.uMS, upper mudstone; U3.CZ, coal zone. doi:10.1371/journal.pone.0016574.g002" figureDoi="http://doi.org/10.5281/zenodo.3744920" httpUri="https://zenodo.org/record/3744920/files/figure.png" pageId="6" pageNumber="7">Figure 2</figureCitation>
|
|||
|
|
). In the remaining two-thirds of the formation,
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFC24A867FBDD2447" box="[958,1082,535,557]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFC24A867FBDD2447" box="[958,1082,535,557]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
is more plentiful than
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFADDA867FA212447" authorityName="Lambe" authorityYear="1917" box="[1351,1478,535,557]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFADDA867FA212447" box="[1351,1478,535,557]" italics="true" pageId="6" pageNumber="7">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
(
|
|||
|
|
<tableCitation id="307F98EF3B23345FFCD5A844FC7C2420" box="[847,923,564,586]" captionStart="Table 1" captionStartId="4.[171,235,174,195]" captionTargetBox="[171,1414,252,856]" captionTargetId="graphics@4.[163,1478,151,962]" captionTargetPageId="4" captionText="Table 1. Hell Creek Formation dinosaur census." httpUri="http://table.plazi.org/id/2982FDDC3B21345DFF31AADEFD7F26A9" pageId="6" pageNumber="7" tableUuid="2982FDDC3B21345DFF31AADEFD7F26A9">Table 1</tableCitation>
|
|||
|
|
). Because the smaller, predatory dinosaur taxa
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFA1AA844FA212420" authorityName="Leidy" authorityYear="1856" box="[1408,1478,564,586]" class="Reptilia" family="Troodontidae" genus="Troodon" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFA1AA844FA212420" box="[1408,1478,564,586]" italics="true" pageId="6" pageNumber="7">Troödon</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
and dromaeosaurids (known from teeth found in microsites) are extremely rare (no skeletons or identifiable lag specimens), it stands to reason that
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFBB6A8F9FB4F24F5" box="[1068,1192,649,671]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFBB6A8F9FB4F24F5" box="[1068,1192,649,671]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
was not a typical predator [37]. In fact, the large numbers of
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFB3AA8D5FAFB24D1" box="[1184,1308,677,699]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
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|
|
<emphasis id="4F8971463B23345FFB3AA8D5FAFB24D1" box="[1184,1308,677,699]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
compared to the smaller theropods suggest that
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFBECA8B1FB1524BD" box="[1142,1266,705,727]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFBECA8B1FB1524BD" box="[1142,1266,705,727]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
benefited from much wider food choice opportunities than exclusively live prey and specific taxa such as
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFB88A88AFB76257A" authorityName="Lambe" authorityYear="1917" box="[1042,1169,762,784]" class="Reptilia" family="Hadrosauridae" genus="Edmontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFB88A88AFB76257A" box="[1042,1169,762,784]" italics="true" pageId="6" pageNumber="7">Edmontosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
[36]. A similar comparison can be made with mammal census numbers from the Serengeti plains where the hyena population is twice that of the combined population of lion, leopard and cheetah [38,39].
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFAD0A93EFA21250E" box="[1354,1478,846,868]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFAD0A93EFA21250E" box="[1354,1478,846,868]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
may have acquired a larger percentage of meat from carrion sources than did smaller theropods, therefore filling the role of a more generalized, carnivorous opportunist such as a hyena. Based on energetic arguments [40], a Serengeti type ecosystem would have provided ample carrion to feed a
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFA88A9ACFA692598" box="[1298,1422,988,1010]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFA88A9ACFA692598" box="[1298,1422,988,1010]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
sized scavenger, particularly if
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFBDDA988FB242264" box="[1095,1219,1016,1038]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFBDDA988FB242264" box="[1095,1219,1016,1038]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
did not have to compete with avian scavengers. In addition,
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFB36AE64FACF2240" box="[1196,1320,1044,1066]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFB36AE64FACF2240" box="[1196,1320,1044,1066]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
adults may not have competed with
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFBA8AE40FB49222C" box="[1074,1198,1072,1094]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFBA8AE40FB49222C" box="[1074,1198,1072,1094]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
juveniles if the potential proclivity for carrion increased with size during ontogeny [41,42]. Such a situation might well explain why
|
|||
|
|
<taxonomicName id="BAFDD6D73B23345FFAD0AE19FA212215" box="[1354,1478,1129,1151]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
|
|||
|
|
<emphasis id="4F8971463B23345FFAD0AE19FA212215" box="[1354,1478,1129,1151]" italics="true" pageId="6" pageNumber="7">Tyrannosaurus</emphasis>
|
|||
|
|
</taxonomicName>
|
|||
|
|
teeth increase in overall robustness while the total number of teeth in the lower jaws decrease during late stages of ontogeny [15].
|
|||
|
|
</paragraph>
|
|||
|
|
</subSubSection>
|
|||
|
|
</treatment>
|
|||
|
|
</document>
|