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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="03FE9007FFA18317FE21FBF97FBBD248" docLanguage="en" docName="peerj-4417.pdf" docOrigin="PeerJ 4417" docStyle="DocumentStyle{}" docTitle="Sauropodomorpha" docType="treatment" docVersion="10" lastPageNumber="30" masterDocId="FFC7E87FFFBA830AFFD3FFB17D18D113" masterDocTitle="How has our knowledge of dinosaur diversity through geologic time changed through research history?" masterLastPageNumber="42" masterPageNumber="1" pageNumber="28" 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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<taxonomicName box="[498,757,1096,1124]" class="Reptilia" kingdom="Animalia" order="Saurischia" pageId="27" pageNumber="28" phylum="Chordata" rank="subOrder" subOrder="Sauropodomorpha">
<emphasis bold="true" box="[498,757,1096,1124]" italics="true" pageId="27" pageNumber="28">Sauropodomorphs</emphasis>
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Subsampled diversity patterns of sauropodomorphs share some characteristics of those of theropods and ornithischians, despite having a different fossil record due to taphonomic differences (i.e. larger, more robust skeletons being preferentially preserved in different environmental settings) (
<bibRefCitation author="Mannion PD &amp; Upchurch P." box="[906,1232,1260,1287]" journalOrPublisher="PaleobiologY" pageId="27" pageNumber="28" pagination="283 - 302" part="36" publicationUrl="10.1666/09008.1." refId="ref17506" refString="Mannion PD, Upchurch P. 2010. Completeness metrics and the quality of the sauropodomorph fossil record through geological and historical time. PaleobiologY 36 (2): 283 - 302 DOI 10.1666 / 09008.1." title="Completeness metrics and the quality of the sauropodomorph fossil record through geological and historical time" type="journal article" year="2010">
<emphasis box="[906,1232,1260,1287]" italics="true" pageId="27" pageNumber="28">Mannion &amp; Upchurch, 2010</emphasis>
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,
<bibRefCitation author="Butler RJ &amp; Benson RBJ &amp; Carrano MT &amp; Mannion PD &amp; Upchurch P." box="[1248,1303,1260,1287]" journalOrPublisher="Proceedings of the RoYal SocietY of London Series B: Biological Sciences" pageId="27" pageNumber="28" 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="[1248,1303,1260,1287]" italics="true" pageId="27" pageNumber="28">2011</emphasis>
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;
<bibRefCitation author="Dean CD &amp; Mannion PD &amp; Butler RJ." journalOrPublisher="PalaeontologY" pageId="27" pageNumber="28" pagination="225 - 247" part="59" publicationUrl="10.1111/pala.12225." refId="ref16136" refString="Dean CD, Mannion PD, Butler RJ. 2016. Preservational bias controls the fossil record of pterosaurs. PalaeontologY 59 (2): 225 - 247 DOI 10.1111 / pala. 12225." title="Preservational bias controls the fossil record of pterosaurs" type="journal article" year="2016">
<emphasis italics="true" pageId="27" pageNumber="28">Dean, Mannion &amp; Butler, 2016</emphasis>
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). This is compounded by a difficulty in assigning a large number of taxa to specific stage bins, which unfortunately excludes many of them from our analyses (Supplemental Information 1). Differences in diversity patterns between sauropodomorphs and ornithischians have classically been interpreted as being due to exclusive competition between the two main herbivorous dinosaurian subtaxa (
<bibRefCitation author="Butler RJ &amp; Barrett PM &amp; Kenrick P &amp; Penn MG." box="[509,716,1499,1526]" journalOrPublisher="Biological Reviews" pageId="27" pageNumber="28" pagination="73 - 89" part="84" publicationUrl="10.1111/j.1469-185x.2008.00065.x." refId="ref15479" refString="Butler RJ, Barrett PM, Kenrick P, Penn MG. 2009. Testing co-evolutionary hypotheses over geological timescales: interactions between Mesozoic non-avian dinosaurs and cycads. Biological Reviews 84 (1): 73 - 89 DOI 10.1111 / j. 1469 - 185 x. 2008.00065. x." title="Testing co-evolutionary hypotheses over geological timescales: interactions between Mesozoic non-avian dinosaurs and cycads" type="journal article" year="2009">
<emphasis box="[509,716,1499,1526]" italics="true" pageId="27" pageNumber="28">Butler et al., 2009</emphasis>
</bibRefCitation>
), with an explosive radiation in ornithischians during the Early Cretaceous resulting from the apparent decline in diversity of sauropodomorphs. In fact, the J/K transition represents a major extinction event for sauropodomorphs, reflecting the decline of non-neosauropods, diplodocoids and basal macronarians (
<bibRefCitation author="Benson RBJ &amp; Mannion PD &amp; Butler RJ &amp; Upchurch P &amp; Goswami A &amp; Evans SE." journalOrPublisher="PalaeogeographY, PalaeoclimatologY, PalaeoecologY" pageId="27" pageNumber="28" 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 italics="true" pageId="27" pageNumber="28">Mannion et al., 2013</emphasis>
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;
<bibRefCitation author="Tennant JP &amp; Mannion PD &amp; Upchurch P." box="[569,1018,1658,1685]" journalOrPublisher="Nature Communications" pageId="27" pageNumber="28" 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="[569,1018,1658,1685]" italics="true" pageId="27" pageNumber="28">Tennant, Mannion &amp; Upchurch, 2016b</emphasis>
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). Sauropodomorph faunas have a low subsampled diversity in the earliest Cretaceous, coupled with a generally poor fossil record (
<bibRefCitation author="Mannion PD &amp; Upchurch P." box="[509,835,1738,1765]" journalOrPublisher="PaleobiologY" pageId="27" pageNumber="28" pagination="283 - 302" part="36" publicationUrl="10.1666/09008.1." refId="ref17506" refString="Mannion PD, Upchurch P. 2010. Completeness metrics and the quality of the sauropodomorph fossil record through geological and historical time. PaleobiologY 36 (2): 283 - 302 DOI 10.1666 / 09008.1." title="Completeness metrics and the quality of the sauropodomorph fossil record through geological and historical time" type="journal article" year="2010">
<emphasis box="[509,835,1738,1765]" italics="true" pageId="27" pageNumber="28">Mannion &amp; Upchurch, 2010</emphasis>
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), but at a time when we otherwise see rapid increases in theropod and ornithischian diversity and a prolonged phase of faunal turnover (
<bibRefCitation author="Brocklehurst N &amp; Upchurch P &amp; Mannion PD &amp; O'Connor J." journalOrPublisher="PLOS ONE" pageId="27" pageNumber="28" pagination="6" part="7" publicationUrl="10.1371/journal.pone.0039056." refId="ref15275" refString="Brocklehurst N, Upchurch P, Mannion PD, O'Connor J. 2012. The completeness of the fossil record of Mesozoic birds: implications for early avian evolution. PLOS ONE 7 (6): e 39056 DOI 10.1371 / journal. pone. 0039056." title="The completeness of the fossil record of Mesozoic birds: implications for early avian evolution" type="journal article" year="2012">
<emphasis italics="true" pageId="27" pageNumber="28">Upchurch &amp; Mannion, 2012</emphasis>
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;
<bibRefCitation author="Tennant JP &amp; Mannion PD &amp; Upchurch P." box="[716,1160,1818,1845]" journalOrPublisher="Nature Communications" pageId="27" pageNumber="28" 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="[716,1160,1818,1845]" italics="true" pageId="27" pageNumber="28">Tennant, Mannion &amp; Upchurch, 2016b</emphasis>
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). Sauropodomorph subsampled diversity levels fluctuate from the middle Cretaceous until the final latest Cretaceous radiation, with a possible small decline in the Maastrichtian. This finding is somewhat contrary to that of
<bibRefCitation author="Sakamoto M &amp; Benton MJ &amp; Venditti C." box="[725,1145,234,261]" journalOrPublisher="Proceedings of the National AcademY of Sciences of the United States of America" pageId="28" pageNumber="29" pagination="5036 - 5040" part="113" publicationUrl="10.1073/pnas.1521478113." refId="ref18970" refString="Sakamoto M, Benton MJ, Venditti C. 2016. Dinosaurs in decline tens of millions of years before their final extinction. Proceedings of the National AcademY of Sciences of the United States of America 113 (18): 5036 - 5040 DOI 10.1073 / pnas. 1521478113." title="Dinosaurs in decline tens of millions of years before their final extinction" type="journal article" year="2016">
<emphasis box="[725,1145,234,261]" italics="true" pageId="28" pageNumber="29">Sakamoto, Benton &amp; Venditti (2016)</emphasis>
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who found that their decline was initiated in the Early Cretaceous, and that the diversification of titanosaurs was at an insufficient rate to compensate for the overall loss of sauropodomorph lineages throughout the rest of the Cretaceous. This discrepancy could be due to the differences in datasets used, and that several recently named titanosaurs taxa have yet to be included in published phylogenies, or the distinction between estimation diversity levels against diversification rates. However, we find that sauropodomorphs are at their most diverse during the Albian (
<figureCitation box="[724,802,513,540]" captionStart="Figure 12" captionStartId="19.[116,180,1302,1324]" captionTargetBox="[92,1490,236,1272]" captionTargetId="figure@19.[90,1492,234,1274]" captionTargetPageId="19" captionText="Figure 12 Subsampled sauropodomorph 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-12" httpUri="https://zenodo.org/record/3097264/files/figure.png" pageId="28" pageNumber="29">Fig. 12</figureCitation>
). Sauropodomorphs appear to be overrepresented with respect to what we might expect for almost the entire duration of the Jurassic, whereas the opposite is true for the Cretaceous (
<bibRefCitation author="Butler RJ &amp; Benson RBJ &amp; Carrano MT &amp; Mannion PD &amp; Upchurch P." box="[925,1166,593,620]" journalOrPublisher="Proceedings of the RoYal SocietY of London Series B: Biological Sciences" pageId="28" pageNumber="29" 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="[925,1166,593,620]" italics="true" pageId="28" pageNumber="29">Mannion et al., 2011</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Butler RJ &amp; Benson RBJ &amp; Carrano MT &amp; Mannion PD &amp; Upchurch P." box="[1180,1425,593,620]" journalOrPublisher="Proceedings of the RoYal SocietY of London Series B: Biological Sciences" pageId="28" pageNumber="29" 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="[1180,1425,593,620]" italics="true" pageId="28" pageNumber="29">Upchurch et al., 2011</emphasis>
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;
<bibRefCitation author="Tennant JP &amp; Mannion PD &amp; Upchurch P." journalOrPublisher="Nature Communications" pageId="28" pageNumber="29" 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 italics="true" pageId="28" pageNumber="29">Tennant, Mannion &amp; Upchurch, 2016b</emphasis>
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). The general patterns of global subsampled diversity shows a steady increase from Middle to the end of Jurassic with a decline through J/K transition (
<bibRefCitation author="Brocklehurst N &amp; Upchurch P &amp; Mannion PD &amp; O'Connor J." box="[509,834,713,740]" journalOrPublisher="PLOS ONE" pageId="28" pageNumber="29" pagination="6" part="7" publicationUrl="10.1371/journal.pone.0039056." refId="ref15275" refString="Brocklehurst N, Upchurch P, Mannion PD, O'Connor J. 2012. The completeness of the fossil record of Mesozoic birds: implications for early avian evolution. PLOS ONE 7 (6): e 39056 DOI 10.1371 / journal. pone. 0039056." title="The completeness of the fossil record of Mesozoic birds: implications for early avian evolution" type="journal article" year="2012">
<emphasis box="[509,834,713,740]" italics="true" pageId="28" pageNumber="29">Upchurch &amp; Mannion, 2012</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Tennant JP &amp; Mannion PD &amp; Upchurch P." box="[848,1296,713,740]" journalOrPublisher="Nature Communications" pageId="28" pageNumber="29" 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="[848,1296,713,740]" italics="true" pageId="28" pageNumber="29">Tennant, Mannion &amp; Upchurch, 2016b</emphasis>
</bibRefCitation>
). The relatively high Late Cretaceous subsampled diversity levels can at least be partially explained by the constant discovery of new titanosaurian taxa, especially from Gondwanan continents (
<bibRefCitation author="Vieira WLS &amp; Vieira KS &amp; Nobrega RP &amp; Montenegro PFGP &amp; Pereira Filho GA &amp; Santana GG &amp; Alves RRN &amp; Almeida WO &amp; Vasconcellos A." box="[510,716,832,859]" journalOrPublisher="PLOS ONE" pageId="28" pageNumber="29" pagination="9" part="9" publicationUrl="10.1371/journal.pone.0108307." refId="ref20049" refString="Vieira WLS, Vieira KS, Nobrega RP, Montenegro PFGP, Pereira Filho GA, Santana GG, Alves RRN, Almeida WO, Vasconcellos A. 2014. Species richness and evidence of random patterns in assemblages of South American Titanosauria during the Late Cretaceous (Campanian - Maastrichtian). PLOS ONE 9 (9): e 108307 DOI 10.1371 / journal. pone. 0108307." title="Species richness and evidence of random patterns in assemblages of South American Titanosauria during the Late Cretaceous (Campanian - Maastrichtian)" type="journal article" year="2014">
<emphasis box="[510,716,832,859]" italics="true" pageId="28" pageNumber="29">Vieira et al., 2014</emphasis>
</bibRefCitation>
;
<emphasis box="[732,1027,832,859]" italics="true" pageId="28" pageNumber="29">
de
<bibRefCitation author="de Jesus Faria CC, Riga BG, dos Anjos Candeiro CR, da Silva Marinho T, David LO, Simbras FM, Castanho RB, Muniz FP, da Costa PVLG" box="[766,1027,832,859]" journalOrPublisher="Journal of South American Earth Sciences" pageId="28" pageNumber="29" pagination="154 - 163" part="61" publicationUrl="10.1016/j.jsames.2014.11.008." refId="ref16070" refString="de Jesus Faria CC, Riga BG, dos Anjos Candeiro CR, da Silva Marinho T, David LO, Simbras FM, Castanho RB, Muniz FP, da Costa PVLG. 2015. Cretaceous sauropod diversity and taxonomic succession in South America. Journal of South American Earth Sciences 61: 154 - 163 DOI 10.1016 / j. jsames. 2014.11.008." title="Cretaceous sauropod diversity and taxonomic succession in South America" type="journal article" year="2015">Jesus Faria et al., 2015</bibRefCitation>
</emphasis>
;
<bibRefCitation author="Bandeira KL &amp; Simbras FM &amp; Machado EB &amp; de Almeida Campos D &amp; Oliveira GR &amp; Kellner AW." box="[1042,1283,832,859]" journalOrPublisher="PLOS ONE" pageId="28" pageNumber="29" pagination="10" part="11" publicationUrl="10.1371/journal.pone.0163373." refId="ref14482" refString="Bandeira KL, Simbras FM, Machado EB, de Almeida Campos D, Oliveira GR, Kellner AW. 2016. A new giant Titanosauria (Dinosauria: Sauropoda) from the Late Cretaceous Bauru Group, Brazil. PLOS ONE 11 (10): e 0163373 DOI 10.1371 / journal. pone. 0163373." title="A new giant Titanosauria (Dinosauria: Sauropoda) from the Late Cretaceous Bauru Group, Brazil" type="journal article" year="2016">
<emphasis box="[1042,1283,832,859]" italics="true" pageId="28" pageNumber="29">Bandeira et al., 2016</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Poropat SF &amp; Mannion PD &amp; Upchurch P &amp; Hocknull SA &amp; Kear BP &amp; Kundrat M &amp; Tischler TR &amp; Sloan T &amp; Sinapius GH &amp; Elliott JA." box="[1298,1524,832,859]" journalOrPublisher="Scientific Reports" pageId="28" pageNumber="29" pagination="34467" part="6" publicationUrl="10.1038/srep34467." refId="ref18638" refString="Poropat SF, Mannion PD, Upchurch P, Hocknull SA, Kear BP, Kundrat M, Tischler TR, Sloan T, Sinapius GH, Elliott JA. 2016. New Australian sauropods shed light on Cretaceous dinosaur palaeobiogeography. Scientific Reports 6: 34467 DOI 10.1038 / srep 34467." title="New Australian sauropods shed light on Cretaceous dinosaur palaeobiogeography" type="journal article" year="2016">
<emphasis box="[1298,1524,832,859]" italics="true" pageId="28" pageNumber="29">Poropat et al., 2016</emphasis>
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), and only recently a more appreciated diversity of diplodocoids (e.g. dicraeosaurids, rebbachisaurids) from relatively poorly sampled regions such as Africa (
<bibRefCitation author="Mannion PD &amp; Barrett PM." journalOrPublisher="Cretaceous Research" pageId="28" pageNumber="29" 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 italics="true" pageId="28" pageNumber="29">Mannion &amp; Barrett, 2013</emphasis>
</bibRefCitation>
;
<bibRefCitation author="Wilson JA &amp; Allain R." box="[665,923,952,979]" journalOrPublisher="Journal of Vertebrate PaleontologY" pageId="28" pageNumber="29" pagination="4" part="35" publicationUrl="10.1080/02724634.2014.1000701." refId="ref20246" refString="Wilson JA, Allain R. 2015. Osteology of Rebbachisaurus garasbae Lavocat, 1954, a diplodocoid (Dinosauria, Sauropoda) from the early Late Cretaceous-aged Kem Kem beds of southeastern Morocco. Journal of Vertebrate PaleontologY 35 (4): e 1000701 DOI 10.1080 / 02724634.2014.1000701." title="Osteology of Rebbachisaurus garasbae Lavocat, 1954, a diplodocoid (Dinosauria, Sauropoda) from the early Late Cretaceous-aged Kem Kem beds of southeastern Morocco" type="journal article" year="2015">
<emphasis box="[665,923,952,979]" italics="true" pageId="28" pageNumber="29">Wilson &amp; Allain, 2015</emphasis>
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;
<bibRefCitation author="Ibrahim N &amp; Dal Sasso C &amp; Maganuco S &amp; Fabbri M &amp; Martill DM &amp; Gorscak E &amp; Lamanna M." box="[939,1168,952,979]" editor="Khosla A" journalOrPublisher="Albuquerque: New Mexico Museum of Natural History &amp; Science" pageId="28" pageNumber="29" pagination="149 - 159" refId="ref16775" refString="Ibrahim N, Dal Sasso C, Maganuco S, Fabbri M, Martill DM, Gorscak E, Lamanna M. 2016. Evidence of a derived titanosaurian (Dinosauria, Sauropoda) in the &quot; Kem Kem beds &quot; of Morocco, with comments on sauropod paleoecology in the Cretaceous of Africa. In: Khosla A, Lucas SG, eds. Cretaceous Period: Biotic DiversitY and BiogeographY. Vol. 71. Albuquerque: New Mexico Museum of Natural History &amp; Science, 149 - 159." title="Evidence of a derived titanosaurian (Dinosauria, Sauropoda) in the &quot; Kem Kem beds &quot; of Morocco, with comments on sauropod paleoecology in the Cretaceous of Africa" type="book chapter" volumeTitle="Cretaceous Period: Biotic DiversitY and BiogeographY. Vol. 71" year="2016">
<emphasis box="[939,1168,952,979]" italics="true" pageId="28" pageNumber="29">Ibrahim et al., 2016</emphasis>
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).
</paragraph>
<paragraph blockId="28.[498,1542,234,1935]" pageId="28" pageNumber="29">
Large-bodied sauropodomorph diversity in the Tithonian is certainly influenced by the intense sampling history of the North American Morrison Formation, where there is an unusually high diversity and cranial disparity of megaherbivores within a relatively resource-poor environment (
<emphasis box="[843,1220,1111,1138]" italics="true" pageId="28" pageNumber="29">
Button,
<bibRefCitation author="Button DJ &amp; Rayfield EJ &amp; Barrett PM." box="[937,1220,1111,1138]" journalOrPublisher="Proceedings of the RoYal SocietY B: Biological Sciences" pageId="28" pageNumber="29" pagination="20142114" part="281" publicationUrl="10.1098/rspb.2014.2114." refId="ref15643" refString="Button DJ, Rayfield EJ, Barrett PM. 2014. Cranial biomechanics underpins high sauropod diversity in resource-poor environments. Proceedings of the RoYal SocietY B: Biological Sciences 281 (1795): 20142114 DOI 10.1098 / rspb. 2014.2114." title="Cranial biomechanics underpins high sauropod diversity in resource-poor environments" type="journal article" year="2014">Rayfield &amp; Barrett, 2014</bibRefCitation>
</emphasis>
). Here, high diversity remains in spite of our accounting for large collection biases associated with Konzentrat- Lagerstätten (
<bibRefCitation author="Alroy J." box="[661,805,1191,1218]" editor="Alroy J" journalOrPublisher="Boulder: The Paleontological Society" pageId="28" pageNumber="29" 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="[661,805,1191,1218]" italics="true" pageId="28" pageNumber="29">Alroy, 2010a</emphasis>
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,
<bibRefCitation author="Alroy J." box="[820,886,1191,1218]" journalOrPublisher="Science" pageId="28" pageNumber="29" 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="[820,886,1191,1218]" italics="true" pageId="28" pageNumber="29">2010c</emphasis>
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), implying that sauropodomorphs reached their zenith in diversity during the Late Jurassic. Sauropodomorphs appear to be better sampled than theropods and ornithischians in Africa (
<figureCitation box="[976,1075,1271,1298]" captionStart="Figure 13" captionStartId="20.[116,180,1302,1324]" captionTargetBox="[92,1477,236,1272]" captionTargetId="figure@20.[90,1479,234,1274]" captionTargetPageId="20" captionText="Figure 13 GOOds u estimates fOr saurOpOdOmOrphs at a (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-13" httpUri="https://zenodo.org/record/3097266/files/figure.png" pageId="28" pageNumber="29">Fig. 13C</figureCitation>
), although their records remain largely too inconsistent and patchy record to reveal any major patterns. Asian subsampled diversity is constantly low until the Maastrichtian, where it increases moderately due to a series of recent discoveries from Pakistan and China (
<bibRefCitation author="Malkani MS." box="[1113,1278,1390,1417]" journalOrPublisher="Sindh UniversitY Research Journal-SURJ (Science Series)" pageId="28" pageNumber="29" pagination="39 - 64" part="42" refId="ref17307" refString="Malkani MS. 2010. New Pakisaurus (Pakisauridae, Titanosauria, Sauropoda) remains, and Cretaceous Tertiary (KT) boundary from Pakistan. Sindh UniversitY Research Journal-SURJ (Science Series) 42 (1): 39 - 64." title="New Pakisaurus (Pakisauridae, Titanosauria, Sauropoda) remains, and Cretaceous Tertiary (KT) boundary from Pakistan" type="journal article" year="2010">
<emphasis box="[1113,1278,1390,1417]" italics="true" pageId="28" pageNumber="29">Malkani, 2010</emphasis>
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;
<bibRefCitation author="Junchang L &amp; Laiping Y &amp; Hui Z &amp; Xuefang W." box="[1291,1524,1390,1417]" journalOrPublisher="Jiangxi Province of southern China. Acta Geologica Sinica (English Edition) 87" pageId="28" pageNumber="29" pagination="678 - 685" part="3" publicationUrl="10.1111/1755-6724.12079." refId="ref16908" refString="Junchang L, Laiping Y, Hui Z, Xuefang W. 2013. A new somphospondylan sauropod (Dinosauria, Titanosauriformes) from the Late Cretaceous of Ganzhou, Jiangxi Province of southern China. Acta Geologica Sinica (English Edition) 87 (3): 678 - 685 DOI 10.1111 / 1755 - 6724.12079." title="A new somphospondylan sauropod (Dinosauria, Titanosauriformes) from the Late Cretaceous of Ganzhou" type="journal article" year="2013">
<emphasis box="[1291,1524,1390,1417]" italics="true" pageId="28" pageNumber="29">Junchang et al., 2013</emphasis>
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). However, the Asian Cretaceous sauropodomorph record is otherwise very poorly sampled, especially compared to ornithischians and theropods. This phenomenon could be explained by a taphonomic size bias discriminating against the preservation of largerbodied animals in pre-Late Cretaceous Konservat-Lagerstätten, while they are more present although more rare in the dense bone assemblages from the latest Cretaceous of Mongolia, China, and India (
<bibRefCitation author="Kidwell SM." box="[848,1005,1629,1656]" editor="Briggs DEG" journalOrPublisher="Oxford: Blackwell" pageId="28" pageNumber="29" pagination="299 - 305" refId="ref16966" refString="Kidwell SM. 2001. Major biases in the fossil record. In: Briggs DEG, Crowther PR, eds. PaleobiologY II, A SYnthesis. Oxford: Blackwell, 299 - 305." title="Major biases in the fossil record" type="book chapter" volumeTitle="PaleobiologY II, A SYnthesis" year="2001">
<emphasis box="[848,1005,1629,1656]" italics="true" pageId="28" pageNumber="29">Kidwell, 2001</emphasis>
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).
</paragraph>
<paragraph blockId="28.[498,1542,234,1935]" lastBlockId="29.[498,1542,234,859]" lastPageId="29" lastPageNumber="30" pageId="28" pageNumber="29">
There is a notable subsampled diversity decline in European sauropodomorphs through the J/K transition, as with other dinosaurian groups (
<bibRefCitation author="Brocklehurst N &amp; Upchurch P &amp; Mannion PD &amp; O'Connor J." box="[1216,1535,1709,1736]" journalOrPublisher="PLOS ONE" pageId="28" pageNumber="29" pagination="6" part="7" publicationUrl="10.1371/journal.pone.0039056." refId="ref15275" refString="Brocklehurst N, Upchurch P, Mannion PD, O'Connor J. 2012. The completeness of the fossil record of Mesozoic birds: implications for early avian evolution. PLOS ONE 7 (6): e 39056 DOI 10.1371 / journal. pone. 0039056." title="The completeness of the fossil record of Mesozoic birds: implications for early avian evolution" type="journal article" year="2012">
<emphasis box="[1216,1535,1709,1736]" italics="true" pageId="28" pageNumber="29">Upchurch &amp; Mannion, 2012</emphasis>
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;
<bibRefCitation author="Tennant JP &amp; Mannion PD &amp; Upchurch P." box="[498,946,1749,1776]" journalOrPublisher="Nature Communications" pageId="28" pageNumber="29" 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="[498,946,1749,1776]" italics="true" pageId="28" pageNumber="29">Tennant, Mannion &amp; Upchurch, 2016b</emphasis>
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). This is distinct from results obtained with other methods (e.g. TRiPS, True Richness estimated using a Poisson Sampling) which do not find any evidence for such a decline (
<bibRefCitation author="Starrfelt J &amp; Liow LH." box="[927,1177,1828,1855]" journalOrPublisher="Philosophical Transactions of the RoYal SocietY B: Biological Sciences" pageId="28" pageNumber="29" pagination="20150219" part="371" publicationUrl="10.1098/rstb.2015.0219." refId="ref19486" refString="Starrfelt J, Liow LH. 2016. How many dinosaur species were there? Fossil bias and true richness estimated using a Poisson sampling model. Philosophical Transactions of the RoYal SocietY B: Biological Sciences 371 (1691): 20150219 DOI 10.1098 / rstb. 2015.0219." title="How many dinosaur species were there? Fossil bias and true richness estimated using a Poisson sampling model" type="journal article" year="2016">
<emphasis box="[927,1177,1828,1855]" italics="true" pageId="28" pageNumber="29">Starrfelt &amp; Liow, 2016</emphasis>
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). Subsampling also reveals that sauropodomorph diversity in the latest Cretaceous of Europe was relatively flat. The sauropodomorph record in South America is poor and mostly confined to the Late Cretaceous, with diversity levels rising and resolution improving through publication time as coverage increases and as new taxa get identified from emerging Patagonian and Brazilian deposits (
<bibRefCitation author="Novas F &amp; Salgado L &amp; Calvo J &amp; Agnolin F." box="[727,947,314,341]" journalOrPublisher="Revista del Museo Argentino de Ciencias Naturales Nueva Serie" pageId="29" pageNumber="30" pagination="31 - 36" part="7" refId="ref18366" refString="Novas F, Salgado L, Calvo J, Agnolin F. 2005 b. Giant titanosaur (Dinosauria, Sauropoda) from the Late Cretaceous of Patagonia. Revista del Museo Argentino de Ciencias Naturales Nueva Serie 7 (1): 31 - 36." title="Giant titanosaur (Dinosauria, Sauropoda) from the Late Cretaceous of Patagonia" type="journal article" year="2005">
<emphasis box="[727,947,314,341]" italics="true" pageId="29" pageNumber="30">Novas et al., 2005b</emphasis>
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;
<bibRefCitation author="Canale JI &amp; Scanferla CA &amp; Agnolin FL &amp; Novas FE." box="[963,1103,314,341]" journalOrPublisher="Naturwissenschaften" pageId="29" pageNumber="30" 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="[963,1103,314,341]" italics="true" pageId="29" pageNumber="30">Novas, 2009</emphasis>
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).
</paragraph>
<paragraph blockId="29.[498,1542,234,859]" pageId="29" pageNumber="30">
Here, it is worth noting the distinction between global and regional sauropodomorph records. On a global level, our results provide strong evidence for a substantial sauropod subsampled diversity decline from the Campanian to Maastrichtian. However, this decline is not represented in any of the regional sauropodomorph diversity signals. Instead, the global signal in the Maastrichtian is comprised of a medley of regional records, which are only continuous with the Campanian record in Europe and North America. Therefore, the global extinction of sauropods in the latest Cretaceous is actually due to regionally heterogeneous sampling signals that are summed into a misleading global curve. A similar case can be made for the apparently global radiation in the Albian, which is primarily a reflection of a well-sampled North American Albian sauropodomorph record (
<figureCitation box="[806,884,752,779]" captionStart="Figure 12" captionStartId="19.[116,180,1302,1324]" captionTargetBox="[92,1490,236,1272]" captionTargetId="figure@19.[90,1492,234,1274]" captionTargetPageId="19" captionText="Figure 12 Subsampled sauropodomorph 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-12" httpUri="https://zenodo.org/record/3097264/files/figure.png" pageId="29" pageNumber="30">Fig. 12</figureCitation>
). Thus, when looking at diversity signals, interpretation of global patterns without considering structural changes on a regional level is not recommended.
</paragraph>
</subSubSection>
</treatment>
</document>