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<document ID-DOI="10.7717/peerj.4417" ID-GBIF-Dataset="58039a1f-b9d0-4c93-b627-410c6decc440" ID-PMC="PMC5822849" ID-PubMed="29479504" ID-Zenodo-Dep="3097200" checkinTime="1519205475471" checkinUser="plazi" docAuthor="Jonathan P. Tennant, Alfio Alessandro Chiarenza &amp; Matthew Baron" docDate="2018" docId="03FE9007FFA08311FE21FE127944D531" docLanguage="en" docName="peerj-4417.pdf" docOrigin="PeerJ 4417" docStyle="DocumentStyle{}" docTitle="Theropoda" docType="treatment" docVersion="10" lastPageNumber="28" masterDocId="FFC7E87FFFBA830AFFD3FFB17D18D113" masterDocTitle="How has our knowledge of dinosaur diversity through geologic time changed through research history?" masterLastPageNumber="42" masterPageNumber="1" pageNumber="27" updateTime="1668138782670" updateUser="ExternalLinkService">
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<mods:title>How has our knowledge of dinosaur diversity through geologic time changed through research history?</mods:title>
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<mods:namePart>Jonathan P. Tennant</mods:namePart>
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<mods:namePart>Alfio Alessandro Chiarenza</mods:namePart>
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<mods:namePart>Matthew Baron</mods:namePart>
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<emphasis bold="true" box="[498,647,419,447]" italics="true" pageId="26" pageNumber="27">Theropods</emphasis>
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As already shown elsewhere (
<bibRefCitation author="Barrett PM &amp; McGowan AJ &amp; Page V." box="[846,1222,463,490]" journalOrPublisher="Proceedings of the RoYal SocietY Series B: Biological Sciences" pageId="26" pageNumber="27" pagination="2667 - 2674" part="1667" publicationUrl="10.1098/rspb.2009.0352." refId="ref14577" refString="Barrett PM, McGowan AJ, Page V. 2009. Dinosaur diversity and the rock record. Proceedings of the RoYal SocietY Series B: Biological Sciences 276 (1667): 2667 - 2674 DOI 10.1098 / rspb. 2009.0352." title="Dinosaur diversity and the rock record" type="journal article" year="2009">
<emphasis box="[846,1222,463,490]" italics="true" pageId="26" pageNumber="27">Barrett, McGowan &amp; Page, 2009</emphasis>
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;
<bibRefCitation author="Brusatte SL &amp; Butler RJ &amp; Prieto-Marquez A &amp; Norell MA." box="[1238,1469,463,490]" journalOrPublisher="Nature Communications" pageId="26" pageNumber="27" pagination="804" part="3" publicationUrl="10.1038/ncomms1815." refId="ref15369" refString="Brusatte SL, Butler RJ, Prieto-Marquez A, Norell MA. 2012. Dinosaur morphological diversity and the end-Cretaceous extinction. Nature Communications 3: 804 DOI 10.1038 / ncomms 1815." title="Dinosaur morphological diversity and the end-Cretaceous extinction" type="journal article" year="2012">
<emphasis box="[1238,1469,463,490]" italics="true" pageId="26" pageNumber="27">Brusatte et al., 2012</emphasis>
</bibRefCitation>
), global theropod diversity trends are overall very similar to that of
<taxonomicName box="[1300,1450,503,530]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="26" pageNumber="27" phylum="Chordata" rank="order">Ornithischia</taxonomicName>
, with subsampled diversity increases during the Late Jurassic (Oxfordian and Tithonian peaks punctuated by a Kimmeridgian decline), late Early Cretaceous (Aptian), early Late Cretaceous (Cenomanian) and latest Cretaceous. Moderately high Middle and Late Jurassic diversity subsampled levels represent the radiation of major avetheropodan clades, and a wealth of new discoveries in recent years, particularly from Asia (
<bibRefCitation author="Butler RJ &amp; Benson RBJ &amp; Carrano MT &amp; Mannion PD &amp; Upchurch P." journalOrPublisher="Proceedings of the RoYal SocietY of London Series B: Biological Sciences" pageId="26" pageNumber="27" pagination="1165 - 1170" part="278" publicationUrl="10.1098/rspb.2010.1754." refId="ref15579" refString="Butler RJ, Benson RBJ, Carrano MT, Mannion PD, Upchurch P. 2011. Sea level, dinosaur diversity and sampling biases: investigating the ' common cause' hypothesis in the terrestrial realm. Proceedings of the RoYal SocietY of London Series B: Biological Sciences 278 (1709): 1165 - 1170 DOI 10.1098 / rspb. 2010.1754." title="Sea level, dinosaur diversity and sampling biases: investigating the ' common cause' hypothesis in the terrestrial realm" type="journal article" year="2011">
<emphasis italics="true" pageId="26" pageNumber="27">Upchurch et al., 2011</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Xu X &amp; You H &amp; Du K &amp; Han F." box="[642,811,742,769]" journalOrPublisher="Nature" pageId="26" pageNumber="27" pagination="465 - 470" part="475" publicationUrl="10.1038/nature10288." refId="ref20297" refString="Xu X, You H, Du K, Han F. 2011. An ArchaeopterYx - like theropod from China and the origin of Avialae. Nature 475 (7357): 465 - 470 DOI 10.1038 / nature 10288." title="An ArchaeopterYx - like theropod from China and the origin of Avialae" type="journal article" year="2011">
<emphasis box="[642,811,742,769]" italics="true" pageId="26" pageNumber="27">Xu et al., 2011</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Carrano MT &amp; Benson RB &amp; Sampson SD." box="[827,1231,742,769]" journalOrPublisher="Journal of SYstematic PalaeontologY" pageId="26" pageNumber="27" pagination="211 - 300" part="10" publicationUrl="10.1080/14772019.2011.630927." refId="ref15786" refString="Carrano MT, Benson RB, Sampson SD. 2012. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of SYstematic PalaeontologY 10 (2): 211 - 300 DOI 10.1080 / 14772019.2011.630927." title="The phylogeny of Tetanurae (Dinosauria: Theropoda)" type="journal article" year="2012">
<emphasis box="[827,1231,742,769]" italics="true" pageId="26" pageNumber="27">Carrano, Benson &amp; Sampson, 2012</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Benson RB &amp; Campione NE &amp; Carrano MT &amp; Mannion PD &amp; Sullivan C &amp; Upchurch P &amp; Evans DC." box="[1247,1465,742,769]" journalOrPublisher="PLOS BiologY" pageId="26" pageNumber="27" pagination="5" part="12" publicationUrl="10.1371/journal.pbio.1001853." refId="ref14823" refString="Benson RB, Campione NE, Carrano MT, Mannion PD, Sullivan C, Upchurch P, Evans DC. 2014. Rates of dinosaur body mass evolution indicate 170 million years of sustained ecological innovation on the avian stem lineage. PLOS BiologY 12 (5): e 1001853 DOI 10.1371 / journal. pbio. 1001853." title="Rates of dinosaur body mass evolution indicate 170 million years of sustained ecological innovation on the avian stem lineage" type="journal article" year="2014">
<emphasis box="[1247,1465,742,769]" italics="true" pageId="26" pageNumber="27">Benson et al., 2014</emphasis>
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;
<bibRefCitation author="Tennant JP &amp; Mannion PD &amp; Upchurch P." box="[498,948,782,809]" journalOrPublisher="Nature Communications" pageId="26" pageNumber="27" pagination="12737" part="7" publicationUrl="10.1038/ncomms12737." refId="ref19700" refString="Tennant JP, Mannion PD, Upchurch P. 2016 b. Sea level regulated tetrapod diversity dynamics through the Jurassic / Cretaceous interval. Nature Communications 7: 12737 DOI 10.1038 / ncomms 12737." title="Sea level regulated tetrapod diversity dynamics through the Jurassic / Cretaceous interval" type="journal article" year="2016">
<emphasis box="[498,948,782,809]" italics="true" pageId="26" pageNumber="27">Tennant, Mannion &amp; Upchurch, 2016b</emphasis>
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).
</paragraph>
<paragraph blockId="26.[498,1542,419,1925]" pageId="26" pageNumber="27">
European subsampled theropod diversity is more constant than in other regions, with a Bajocian peak followed by a BathonianOxfordian trough, and a Kimmeridgian peak followed by a Tithonian to Valanginian drop. This can, at least in part, be explained by an abundance of well-sampled Late Jurassic formations from across Western Europe (
<bibRefCitation author="Butler RJ &amp; Benson RBJ &amp; Carrano MT &amp; Mannion PD &amp; Upchurch P." box="[509,756,981,1008]" journalOrPublisher="Proceedings of the RoYal SocietY of London Series B: Biological Sciences" pageId="26" pageNumber="27" pagination="1165 - 1170" part="278" publicationUrl="10.1098/rspb.2010.1754." refId="ref15579" refString="Butler RJ, Benson RBJ, Carrano MT, Mannion PD, Upchurch P. 2011. Sea level, dinosaur diversity and sampling biases: investigating the ' common cause' hypothesis in the terrestrial realm. Proceedings of the RoYal SocietY of London Series B: Biological Sciences 278 (1709): 1165 - 1170 DOI 10.1098 / rspb. 2010.1754." title="Sea level, dinosaur diversity and sampling biases: investigating the ' common cause' hypothesis in the terrestrial realm" type="journal article" year="2011">
<emphasis box="[509,756,981,1008]" italics="true" pageId="26" pageNumber="27">Upchurch et al., 2011</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Benson RBJ &amp; Mannion PD &amp; Butler RJ &amp; Upchurch P &amp; Goswami A &amp; Evans SE." box="[771,989,981,1008]" journalOrPublisher="PalaeogeographY, PalaeoclimatologY, PalaeoecologY" pageId="26" pageNumber="27" pagination="88 - 107" part="372" publicationUrl="10.1016/j.palaeo.2012.10.028." refId="ref14884" refString="Benson RBJ, Mannion PD, Butler RJ, Upchurch P, Goswami A, Evans SE. 2013. Cretaceous tetrapod fossil record sampling and faunal turnover: implications for biogeography and the rise of modern clades. PalaeogeographY, PalaeoclimatologY, PalaeoecologY 372: 88 - 107 DOI 10.1016 / j. palaeo. 2012.10.028." title="Cretaceous tetrapod fossil record sampling and faunal turnover: implications for biogeography and the rise of modern clades" type="journal article" year="2013">
<emphasis box="[771,989,981,1008]" italics="true" pageId="26" pageNumber="27">Benson et al., 2013</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Tennant JP &amp; Mannion PD &amp; Upchurch P." box="[1005,1236,981,1008]" journalOrPublisher="Proceedings of the RoYal SocietY B: Biological Sciences" pageId="26" pageNumber="27" pagination="20152840" part="283" publicationUrl="10.1098/rspb.2015.2840." refId="ref19654" refString="Tennant JP, Mannion PD, Upchurch P. 2016 a. Environmental drivers of crocodyliform extinction across the Jurassic / Cretaceous transition. Proceedings of the RoYal SocietY B: Biological Sciences 283 (1826): 20152840 DOI 10.1098 / rspb. 2015.2840." title="Environmental drivers of crocodyliform extinction across the Jurassic / Cretaceous transition" type="journal article" year="2016">
<emphasis box="[1005,1236,981,1008]" italics="true" pageId="26" pageNumber="27">Tennant et al., 2016</emphasis>
</bibRefCitation>
). Barremian diversity is increasing rapidly through publication history, and is now as high as calculated for the Kimmeridgian. As with the Late Jurassic, at least part of this signal represents the influence of a Lagerstätten effect (e.g. Las Hoyas, Spain) (
<bibRefCitation author="Buscalioni A &amp; Fregenal M &amp; Bravo A &amp; Poyato-Ariza F &amp; Sanchiz B &amp; Baez A &amp; Moo OC &amp; Closas CM &amp; Evans S &amp; LobOn JM." box="[1068,1322,1101,1128]" journalOrPublisher="Cretaceous Research" pageId="26" pageNumber="27" pagination="687 - 710" part="29" publicationUrl="10.1016/j.cretres.2008.02.004." refId="ref15402" refString="Buscalioni A, Fregenal M, Bravo A, Poyato-Ariza F, Sanchiz B, Baez A, Moo OC, Closas CM, Evans S, LobOn JM. 2008. The vertebrate assemblage of Buenache de la Sierra (Upper Barremian of Serrania de Cuenca, Spain) with insights into its taphonomy and palaeoecology. Cretaceous Research 29 (4): 687 - 710 DOI 10.1016 / j. cretres. 2008.02.004." title="The vertebrate assemblage of Buenache de la Sierra (Upper Barremian of Serrania de Cuenca, Spain) with insights into its taphonomy and palaeoecology" type="journal article" year="2008">
<emphasis box="[1068,1322,1101,1128]" italics="true" pageId="26" pageNumber="27">Buscalioni et al., 2008</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Butler RJ &amp; Benson RBJ &amp; Carrano MT &amp; Mannion PD &amp; Upchurch P." journalOrPublisher="Proceedings of the RoYal SocietY of London Series B: Biological Sciences" pageId="26" pageNumber="27" pagination="1165 - 1170" part="278" publicationUrl="10.1098/rspb.2010.1754." refId="ref15579" refString="Butler RJ, Benson RBJ, Carrano MT, Mannion PD, Upchurch P. 2011. Sea level, dinosaur diversity and sampling biases: investigating the ' common cause' hypothesis in the terrestrial realm. Proceedings of the RoYal SocietY of London Series B: Biological Sciences 278 (1709): 1165 - 1170 DOI 10.1098 / rspb. 2010.1754." title="Sea level, dinosaur diversity and sampling biases: investigating the ' common cause' hypothesis in the terrestrial realm" type="journal article" year="2011">
<emphasis italics="true" pageId="26" pageNumber="27">Upchurch et al., 2011</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Sanchez-Hernandez B &amp; Benton MJ." box="[569,987,1140,1167]" journalOrPublisher="Acta Palaeontologica Polonica" pageId="26" pageNumber="27" pagination="581 - 600" part="59" publicationUrl="10.4202/app.2011.0144." refId="ref19123" refString="Sanchez-Hernandez B, Benton MJ. 2012. Filling the ceratosaur gap: a new ceratosaurian theropod from the Early Cretaceous of Spain. Acta Palaeontologica Polonica 59: 581 - 600 DOI 10.4202 / app. 2011.0144." title="Filling the ceratosaur gap: a new ceratosaurian theropod from the Early Cretaceous of Spain" type="journal article" year="2012">
<emphasis box="[569,987,1140,1167]" italics="true" pageId="26" pageNumber="27">Sánchez-Hernández &amp; Benton, 2012</emphasis>
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), highlighting that single, well-sampled formations can have a profound historical effect on our understanding of regional diversity patterns, even when subsampling methods are applied. The European AptianAlbian record is increasing slower through time compared to the Campanian Maastrichtian. However, this might possibly change in the future, as the ichnological record in southern Europe is quite abundant for the AptianAlbian interval, and suggests a currently unrecognised dinosaurian diversity present there (
<bibRefCitation author="Dalla Vecchia F." box="[1232,1460,1379,1406]" journalOrPublisher="Memorie della Societa Geologica Italiana" pageId="26" pageNumber="27" pagination="89 - 100" part="57" refId="ref16034" refString="Dalla Vecchia F. 2002. Cretaceous dinosaurs in the Adriatic-Dinaric carbonate platform (Italy and Croatia): paleoenvironmental implications and paleogeographical hypotheses. Memorie della Societa Geologica Italiana 57: 89 - 100." title="Cretaceous dinosaurs in the Adriatic-Dinaric carbonate platform (Italy and Croatia): paleoenvironmental implications and paleogeographical hypotheses" type="journal article" year="2002">
<emphasis box="[1232,1460,1379,1406]" italics="true" pageId="26" pageNumber="27">Dalla Vecchia, 2002</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Meyer C &amp; Thuring B." box="[498,773,1419,1446]" journalOrPublisher="DOI" pageId="26" pageNumber="27" pagination="221 - 228" part="10" publicationUrl="10.1080/10420940390256186." refId="ref17841" refString="Meyer C, Thuring B. 2003. The first iguanodontid dinosaur tracks from the Swiss Alps (Schrattenkalk Formation, Aptian). Ichnos 10 (2 - 4): 221 - 228 DOI 10.1080 / 10420940390256186." title="The first iguanodontid dinosaur tracks from the Swiss Alps (Schrattenkalk Formation, Aptian). Ichnos" type="journal article" year="2003">
<emphasis box="[498,773,1419,1446]" italics="true" pageId="26" pageNumber="27">Meyer &amp; Thuring, 2003</emphasis>
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).
</paragraph>
<paragraph blockId="26.[498,1542,419,1925]" lastBlockId="27.[498,1542,234,1058]" lastPageId="27" lastPageNumber="28" pageId="26" pageNumber="27">
The North American theropod record is dwarfed by an oversampling of latest Cretaceous dinosaur-bearing formations (e.g. Dinosaur Provincial Park, Hell Creek Formation). An increasingly even representation of latitudinally diverse localities from the CenomanianCampanian of Utah, Colorado, New Mexico, and Mexico (e.g. Wahweap Formation), may increase the magnitude of the small subsampled diversity drop through the Maastrichtian. Subsampling highlights a latest Jurassic peak in diversity (due to the abundance of remains from the well-sampled Morrison Formation;
<bibRefCitation author="Foster JR." box="[1306,1471,1698,1725]" journalOrPublisher="Albuquerque: New Mexico Museum of Natural History and Science" pageId="26" pageNumber="27" pagination="23" refId="ref16259" refString="Foster JR. 2003. Paleoecological AnalYsis of the Vertebrate Fauna of the Morrison Formation (Upper Jurassic), RockY Mountain Region, USA: Bulletin 23. Albuquerque: New Mexico Museum of Natural History and Science." title="Paleoecological AnalYsis of the Vertebrate Fauna of the Morrison Formation (Upper Jurassic), RockY Mountain Region, USA: Bulletin" type="book chapter" year="2003">
<emphasis box="[1306,1459,1698,1725]" italics="true" pageId="26" pageNumber="27">Foster (2003)</emphasis>
)
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, although Jurassic subsampled diversity never attains that of the Cretaceous highs during the Aptian and Campanian. In contrast to
<bibRefCitation author="Brusatte SL &amp; Butler RJ &amp; Barrett PM &amp; Carrano MT &amp; Evans DC &amp; Lloyd GT &amp; Mannion PD &amp; Norell MA &amp; Peppe DJ &amp; Upchurch P." box="[1096,1342,1778,1805]" journalOrPublisher="Biological Reviews" pageId="26" pageNumber="27" pagination="628 - 642" part="90" refId="ref15323" refString="Brusatte SL, Butler RJ, Barrett PM, Carrano MT, Evans DC, Lloyd GT, Mannion PD, Norell MA, Peppe DJ, Upchurch P. 2015. The extinction of the dinosaurs. Biological Reviews 90: 628 - 642." title="The extinction of the dinosaurs" type="journal article" year="2015">
<emphasis box="[1096,1342,1778,1805]" italics="true" pageId="26" pageNumber="27">Brusatte et al. (2015)</emphasis>
</bibRefCitation>
, who found no evidence for a progressive Campanian-Maastrichtian decline in North American theropod faunas using similar SQS analyses (implemented in R; see (
<bibRefCitation author="Tennant JP &amp; Mannion PD &amp; Upchurch P." journalOrPublisher="Proceedings of the RoYal SocietY B: Biological Sciences" pageId="26" pageNumber="27" pagination="20152840" part="283" publicationUrl="10.1098/rspb.2015.2840." refId="ref19654" refString="Tennant JP, Mannion PD, Upchurch P. 2016 a. Environmental drivers of crocodyliform extinction across the Jurassic / Cretaceous transition. Proceedings of the RoYal SocietY B: Biological Sciences 283 (1826): 20152840 DOI 10.1098 / rspb. 2015.2840." title="Environmental drivers of crocodyliform extinction across the Jurassic / Cretaceous transition" type="journal article" year="2016">
<emphasis italics="true" pageId="26" pageNumber="27">Tennant, Mannion &amp; Upchurch, 2016a</emphasis>
</bibRefCitation>
,
<bibRefCitation author="Tennant JP &amp; Mannion PD &amp; Upchurch P." box="[580,648,1898,1925]" journalOrPublisher="Nature Communications" pageId="26" pageNumber="27" pagination="12737" part="7" publicationUrl="10.1038/ncomms12737." refId="ref19700" refString="Tennant JP, Mannion PD, Upchurch P. 2016 b. Sea level regulated tetrapod diversity dynamics through the Jurassic / Cretaceous interval. Nature Communications 7: 12737 DOI 10.1038 / ncomms 12737." title="Sea level regulated tetrapod diversity dynamics through the Jurassic / Cretaceous interval" type="journal article" year="2016">
<emphasis box="[580,648,1898,1925]" italics="true" pageId="26" pageNumber="27">2016b</emphasis>
</bibRefCitation>
) and (
<bibRefCitation author="Alroy J." box="[727,869,1898,1925]" editor="Alroy J" journalOrPublisher="Boulder: The Paleontological Society" pageId="26" pageNumber="27" pagination="55 - 80" refId="ref14263" refString="Alroy J. 2010 a. Fair sampling of taxonomic richness and unbiased estimation of origination and extinction rates. In: Alroy J, Hunt G, eds. Quantitative Methods in PaleobiologY. Boulder: The Paleontological Society, 55 - 80." title="Fair sampling of taxonomic richness and unbiased estimation of origination and extinction rates" type="book chapter" volumeTitle="Quantitative Methods in PaleobiologY" year="2010">
<emphasis box="[727,869,1898,1925]" italics="true" pageId="26" pageNumber="27">Alroy, 2010a</emphasis>
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,
<bibRefCitation author="Alroy J." box="[882,947,1898,1925]" journalOrPublisher="Science" pageId="26" pageNumber="27" pagination="1191 - 1194" part="329" publicationUrl="10.1126/science.1189910." refId="ref14346" refString="Alroy J. 2010 c. The shifting balance of diversity among major marine animal groups. Science 329 (5996): 1191 - 1194 DOI 10.1126 / science. 1189910." title="The shifting balance of diversity among major marine animal groups" type="journal article" year="2010">
<emphasis box="[882,947,1898,1925]" italics="true" pageId="26" pageNumber="27">2010c</emphasis>
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) for comparative discussions), we find a very slight decline that remains constant through publication history, that likely relates to our usage of a slightly different subsampling approach. Aptian subsampled diversity is relatively high due to the more heavily sampled localities from Montana to Texas (
<bibRefCitation author="Kirkland JI &amp; Britt B &amp; Burge DL &amp; Carpenter K &amp; Cifelli R &amp; DeCourten F &amp; Eaton J &amp; Hasiotis S &amp; Lawton T." journalOrPublisher="Brigham Young UniversitY GeologY Studies" pageId="27" pageNumber="28" pagination="69 - 104" part="42" refId="ref17002" refString="Kirkland JI, Britt B, Burge DL, Carpenter K, Cifelli R, DeCourten F, Eaton J, Hasiotis S, Lawton T. 1997. Lower to middle Cretaceous dinosaur faunas of the central Colorado Plateau: a key to understanding 35 million years of tectonics, sedimentology, evolution, and biogeography. Brigham Young UniversitY GeologY Studies 42: 69 - 104." title="Lower to middle Cretaceous dinosaur faunas of the central Colorado Plateau: a key to understanding 35 million years of tectonics, sedimentology, evolution, and biogeography" type="journal article" year="1997">
<emphasis italics="true" pageId="27" pageNumber="28">Kirkland et al., 1997</emphasis>
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;
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<emphasis box="[569,774,354,381]" italics="true" pageId="27" pageNumber="28">Cifelli et al., 1999</emphasis>
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<bibRefCitation author="Kirkland JI &amp; Madsen SK." box="[789,1089,354,381]" editor="Lund WR" journalOrPublisher="Annual Meeting, Grand Junction Geological SocietY. Vol. 35. Utah: Geological Association Publication" pageId="27" pageNumber="28" pagination="1 - 108" refId="ref17071" refString="Kirkland JI, Madsen SK. 2007. The Lower Cretaceous Cedar Mountain Formation, eastern Utah: The view up an always interesting learning curve. In: Lund WR, ed. Field Guide to Geological Excursions in Southern Utah, Geological SocietY of America RockY Mountain Section 2007 Annual Meeting, Grand Junction Geological SocietY. Vol. 35. Utah: Geological Association Publication, 1 - 108." title="The Lower Cretaceous Cedar Mountain Formation, eastern Utah: The view up an always interesting learning curve" type="book chapter" volumeTitle="Field Guide to Geological Excursions in Southern Utah, Geological SocietY of America RockY Mountain" year="2007">
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).
</paragraph>
<paragraph blockId="27.[498,1542,234,1058]" pageId="27" pageNumber="28">
In Africa, there is a Cenomanian radiation (
<figureCitation box="[1055,1140,394,421]" captionStart="Figure 9" captionStartId="16.[116,180,1302,1324]" captionTargetBox="[92,1494,236,1272]" captionTargetId="figure@16.[90,1496,234,1274]" captionTargetPageId="16" captionText="Figure 9 Subsampled therOpOd diVersity at (A) glObal and (BF) regiOnal leVels (EurOpe, Africa, Asia, NOrth America, and SOuth America, respectiVely) based On Our published knOWledge in 1991 and 2015. Abbreviations as Fig. 4. Full-size DOI: 10.7717/peerj.4417/fig-9" httpUri="https://zenodo.org/record/3097248/files/figure.png" pageId="27" pageNumber="28">Fig. 9C</figureCitation>
) mainly due to the multitaxic theropod dominated Kem Kem beds and other AlbianCenomanian (middle Cretaceous) equivalents in Northern Africa, but this signal might have been altered by time averaging effects constraining a more temporally diluted diversity in a single unit (
<bibRefCitation author="Mannion PD &amp; Barrett PM." box="[509,805,553,580]" journalOrPublisher="Cretaceous Research" pageId="27" pageNumber="28" pagination="49 - 59" part="45" publicationUrl="10.1016/j.cretres.2013.07.007." refId="ref17350" refString="Mannion PD, Barrett PM. 2013. Additions to the sauropod dinosaur fauna of the Cenomanian (early Late Cretaceous) Kem Kem beds of Morocco: palaeobiogeographical implications of the mid-Cretaceous African sauropod fossil record. Cretaceous Research 45: 49 - 59 DOI 10.1016 / j. cretres. 2013.07.007." title="Additions to the sauropod dinosaur fauna of the Cenomanian (early Late Cretaceous) Kem Kem beds of Morocco: palaeobiogeographical implications of the mid-Cretaceous African sauropod fossil record" type="journal article" year="2013">
<emphasis box="[509,805,553,580]" italics="true" pageId="27" pageNumber="28">Mannion &amp; Barrett, 2013</emphasis>
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;
<bibRefCitation author="Evers SW &amp; Rauhut OW &amp; Milner AC &amp; McFeeters B &amp; Allain R." box="[820,1014,553,580]" journalOrPublisher="PeerJ" pageId="27" pageNumber="28" pagination="1323" part="3" publicationUrl="10.7717/peerj.1323." refId="ref16208" refString="Evers SW, Rauhut OW, Milner AC, McFeeters B, Allain R. 2015. A reappraisal of the morphology and systematic position of the theropod dinosaur Sigilmassasaurus from the &quot; middle &quot; Cretaceous of Morocco. PeerJ 3: e 1323 DOI 10.7717 / peerj. 1323." title="A reappraisal of the morphology and systematic position of the theropod dinosaur Sigilmassasaurus from the &quot; middle &quot; Cretaceous of Morocco" type="journal article" year="2015">
<emphasis box="[820,1014,553,580]" italics="true" pageId="27" pageNumber="28">Evers et al., 2015</emphasis>
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;
<bibRefCitation author="Chiarenza AA &amp; Cau A." box="[1029,1303,553,580]" journalOrPublisher="PeerJ" pageId="27" pageNumber="28" pagination="1754" part="4" publicationUrl="10.7717/peerj.1754." refId="ref15861" refString="Chiarenza AA, Cau A. 2016. A large abelisaurid (Dinosauria, Theropoda) from Morocco and comments on the Cenomanian theropods from North Africa. PeerJ 4: e 1754 DOI 10.7717 / peerj. 1754." title="A large abelisaurid (Dinosauria, Theropoda) from Morocco and comments on the Cenomanian theropods from North Africa" type="journal article" year="2016">
<emphasis box="[1029,1303,553,580]" italics="true" pageId="27" pageNumber="28">Chiarenza &amp; Cau, 2016</emphasis>
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). Asian subsampled diversity peaks in the Aptian, Campanian, and Maastrichtian might be explained by a Lagerstätten bonanza effect, especially considering the high quality preservation deposits discovered and heavily sampled in the last 20 years (e.g. Liaoning) (
<emphasis box="[1306,1505,673,700]" italics="true" pageId="27" pageNumber="28">Lloyd et al., 2008</emphasis>
;
<bibRefCitation author="Zhou Z &amp; Wang Y." box="[498,731,713,740]" journalOrPublisher="Science China Earth Sciences" pageId="27" pageNumber="28" pagination="1894 - 1907" part="53" publicationUrl="10.1007/s11430-010-4094-9." refId="ref20339" refString="Zhou Z, Wang Y. 2010. Vertebrate diversity of the Jehol Biota as compared with other lagerstatten. Science China Earth Sciences 53 (12): 1894 - 1907 DOI 10.1007 / s 11430 - 010 - 4094 - 9." title="Vertebrate diversity of the Jehol Biota as compared with other lagerstatten" type="journal article" year="2010">
<emphasis box="[498,731,713,740]" italics="true" pageId="27" pageNumber="28">Zhou &amp; Wang, 2010</emphasis>
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;
<bibRefCitation author="Godefroit P &amp; Demuynck H &amp; Dyke G &amp; Hu D &amp; Escuillie F &amp; Claeys P." box="[746,986,713,740]" journalOrPublisher="Nature Communications" pageId="27" pageNumber="28" pagination="1394" part="4" publicationUrl="10.1038/ncomms2389." refId="ref16435" refString="Godefroit P, Demuynck H, Dyke G, Hu D, Escuillie F, Claeys P. 2013. Reduced plumage and flight ability of a new Jurassic paravian theropod from China. Nature Communications 4: 1394 DOI 10.1038 / ncomms 2389." title="Reduced plumage and flight ability of a new Jurassic paravian theropod from China" type="journal article" year="2013">
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;
<bibRefCitation author="O'Connor J &amp; Zhou Z." box="[1000,1281,713,740]" journalOrPublisher="DOI" pageId="27" pageNumber="28" pagination="333 - 342" part="156" publicationUrl="10.1007/s10336-015-1222-5." refId="ref18413" refString="O'Connor J, Zhou Z. 2015. Early evolution of the biological bird: perspectives from new fossil discoveries in China. Journal of OrnithologY 156 (S 1): 333 - 342 DOI 10.1007 / s 10336 - 015 - 1222 - 5." title="Early evolution of the biological bird: perspectives from new fossil discoveries in China. Journal of OrnithologY" type="journal article" year="2015">
<emphasis box="[1000,1281,713,740]" italics="true" pageId="27" pageNumber="28">OConnor &amp; Zhou, 2015</emphasis>
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;
<bibRefCitation author="Tennant JP &amp; Mannion PD &amp; Upchurch P." box="[1296,1524,713,740]" journalOrPublisher="Proceedings of the RoYal SocietY B: Biological Sciences" pageId="27" pageNumber="28" pagination="20152840" part="283" publicationUrl="10.1098/rspb.2015.2840." refId="ref19654" refString="Tennant JP, Mannion PD, Upchurch P. 2016 a. Environmental drivers of crocodyliform extinction across the Jurassic / Cretaceous transition. Proceedings of the RoYal SocietY B: Biological Sciences 283 (1826): 20152840 DOI 10.1098 / rspb. 2015.2840." title="Environmental drivers of crocodyliform extinction across the Jurassic / Cretaceous transition" type="journal article" year="2016">
<emphasis box="[1296,1524,713,740]" italics="true" pageId="27" pageNumber="28">Tennant et al., 2016</emphasis>
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), although coverage remains only moderate (around 0.5) in each of these intervals (
<figureCitation box="[1446,1524,752,779]" captionStart="Figure 10" captionStartId="17.[116,180,1302,1324]" captionTargetBox="[92,1496,236,1272]" captionTargetId="figure@17.[90,1498,234,1274]" captionTargetPageId="17" captionText="Figure 10 GOOdsu estimates fOrtherOpOds at (A) glObal and (BF)regiOnal leVels(EurOpe, Africa, Asia, NOrth America, and SOuthAmerica, respectiVely) based On Our published knOWledge in 1991 and 2015. Abbreviations as Fig. 4. Full-size DOI: 10.7717/peerj.4417/fig-10" httpUri="https://zenodo.org/record/3097250/files/figure.png" pageId="27" pageNumber="28">Fig. 10</figureCitation>
). Similarly to the pattern in Africa and Asia, South American theropod subsampled diversity stands out compared to other North America and Europe, remaining relatively signal deficient. Despite an increasing rate of discovery of new taxa, which often alter our knowledge of dinosaur phylogeny and biogeography from the middle Cretaceous of Patagonia and Brazil (
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<emphasis box="[762,983,952,979]" italics="true" pageId="27" pageNumber="28">Novas et al., 2005a</emphasis>
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,
<bibRefCitation author="Novas FE &amp; Agnolin FL &amp; Ezcurra MD &amp; Porfiri J &amp; Canale JI." box="[998,1053,952,979]" journalOrPublisher="Cretaceous Research" pageId="27" pageNumber="28" pagination="174 - 215" part="45" publicationUrl="10.1016/j.cretres.2013.04.001." refId="ref18232" refString="Novas FE, Agnolin FL, Ezcurra MD, Porfiri J, Canale JI. 2013. Evolution of the carnivorous dinosaurs during the Cretaceous: the evidence from Patagonia. Cretaceous Research 45: 174 - 215 DOI 10.1016 / j. cretres. 2013.04.001." title="Evolution of the carnivorous dinosaurs during the Cretaceous: the evidence from Patagonia" type="journal article" year="2013">
<emphasis box="[998,1053,952,979]" italics="true" pageId="27" pageNumber="28">2013</emphasis>
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;
<bibRefCitation author="Novas FE &amp; Pol D." box="[1069,1285,952,979]" journalOrPublisher="Nature" pageId="27" pageNumber="28" pagination="858 - 861" part="433" publicationUrl="10.1038/nature03285." refId="ref18331" refString="Novas FE, Pol D. 2005. New evidence on deinonychosaurian dinosaurs from the Late Cretaceous of Patagonia. Nature 433 (7028): 858 - 861 DOI 10.1038 / nature 03285." title="New evidence on deinonychosaurian dinosaurs from the Late Cretaceous of Patagonia" type="journal article" year="2005">
<emphasis box="[1069,1285,952,979]" italics="true" pageId="27" pageNumber="28">Novas &amp; Pol, 2005</emphasis>
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;
<bibRefCitation author="Canale JI &amp; Scanferla CA &amp; Agnolin FL &amp; Novas FE." box="[1300,1517,952,979]" journalOrPublisher="Naturwissenschaften" pageId="27" pageNumber="28" pagination="409 - 414" part="96" publicationUrl="10.1007/s00114-008-0487-4." refId="ref15734" refString="Canale JI, Scanferla CA, Agnolin FL, Novas FE. 2009. New carnivorous dinosaur from the Late Cretaceous of NW Patagonia and the evolution of abelisaurid theropods. Naturwissenschaften 96 (3): 409 - 414 DOI 10.1007 / s 00114 - 008 - 0487 - 4." title="New carnivorous dinosaur from the Late Cretaceous of NW Patagonia and the evolution of abelisaurid theropods" type="journal article" year="2009">
<emphasis box="[1300,1517,952,979]" italics="true" pageId="27" pageNumber="28">Canale et al., 2009</emphasis>
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), coverage remains poor at the stage level, emphasising the need for greater stratigraphic resolution of the theropod-bearing formations here.
</paragraph>
</subSubSection>
</treatment>
</document>