761 lines
98 KiB
XML
761 lines
98 KiB
XML
<document id="57B1E8AEE7D1581C0E674EC6E7FCA6EC" ID-CLB-Dataset="304675" ID-DOI="10.1093/zoolinnean/zlae090" ID-GBIF-Dataset="7b00e980-780d-4e4c-ad80-ce055675d5e0" ID-ISSN="0024-4082" ID-Zenodo-Dep="13774241" IM.bibliography_approvedBy="valdenar" IM.illustrations_approvedBy="juliana" IM.materialsCitations_approvedBy="juliana" IM.metadata_approvedBy="valdenar" IM.tables_approvedBy="juliana" IM.taxonomicNames_approvedBy="juliana" IM.treatments_approvedBy="juliana" checkinTime="1726225572511" checkinUser="plazi" docAuthor="Rauhut, Oliver W. M., Bakirov, Aizek A., Wings, Oliver, Fernandes, Alexandra E. & Hübner, Tom R." docDate="2024" docId="03F2A447F151E025FF4A84815F7C1554" docLanguage="en" docName="zlae090.pdf" docOrigin="Zoological Journal of the Linnean Society 201 (4)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlae090" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.6:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleVersion="6" docTitle="Alpkarakush" docType="treatment" docVersion="3" lastPageNumber="44" masterDocId="FFCBDC3FF176E00EFFBB87755E621141" masterDocTitle="A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan" masterLastPageNumber="51" masterPageNumber="1" pageNumber="40" updateTime="1729533217363" updateUser="ExternalLinkService">
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<mods:title id="F31222AD24A3AEDEB3DDE8DF6DEF9AEC">A new theropod dinosaur from the Callovian Balabansai Formation of Kyrgyzstan</mods:title>
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<mods:namePart id="30ED8EA90D5C4175309B3470ADA3F83C">Rauhut, Oliver W. M.</mods:namePart>
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<mods:affiliation id="95BACC2C639CDD5AAF1B71FE06C8CFA8">SNSB - Bayerische Staatssammlung für Paläontologie und Geologie, Munich, Germany & Department für Geo- und Umweltwissenschaften, Ludwig-Maximilians-Universität, Munich, Germany & GeoBioCenter, Ludwig-Maximilians-Universität, Munich, Germany</mods:affiliation>
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<mods:nameIdentifier id="F53696B59DE6CD13862631778C546F3E" type="email">rauhut@snsb.de</mods:nameIdentifier>
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<mods:roleTerm id="6E1EAE5975783046A97F6BBC64378E1F">Author</mods:roleTerm>
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<mods:namePart id="88E5C9E24E1CE3F03D385221D426A309">Bakirov, Aizek A.</mods:namePart>
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<mods:affiliation id="41F49DC1F20BDAF284C68D3161019001">Institute of Geology n. a. M. M. Adyshev, National Academy of Sciences of Kyrgyz Republic, Bishkek, Kyrgyzstan</mods:affiliation>
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<mods:namePart id="FD46AD6ACF582A216390056E3DD4054D">Wings, Oliver</mods:namePart>
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<mods:affiliation id="0E6BA5DDEE6ABB1550A311CB737FCC70">SNSB - Naturkundemuseum Bamberg, Bamberg, Germany</mods:affiliation>
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<mods:namePart id="F60E8FCEA0D11F2DE41FA4289FA30A3E">Fernandes, Alexandra E.</mods:namePart>
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<mods:affiliation id="FF79945CB91C94B18F251B18FD793E28">SNSB - Bayerische Staatssammlung für Paläontologie und Geologie, Munich, Germany & Department für Geo- und Umweltwissenschaften, Ludwig-Maximilians-Universität, Munich, Germany</mods:affiliation>
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<mods:namePart id="35C4A0F5C8D744E7B65778630BE68BEC">Hübner, Tom R.</mods:namePart>
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<mods:affiliation id="F966F53244EBAEF23BB66A5EE626B8BB">Friedenstein Stiftung Gotha, Gotha, Germany</mods:affiliation>
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<mods:date id="7B0D6BFA7839578DFB1F148BC7F834BC">2024</mods:date>
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<treatment id="03F2A447F151E025FF4A84815F7C1554" ID-DOI="http://doi.org/10.5281/zenodo.13963634" ID-Zenodo-Dep="13963634" LSID="urn:lsid:plazi:treatment:03F2A447F151E025FF4A84815F7C1554" httpUri="http://treatment.plazi.org/id/03F2A447F151E025FF4A84815F7C1554" lastPageId="43" lastPageNumber="44" pageId="39" pageNumber="40">
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<subSubSection id="C34146DAF151E029FF4A84815C18154E" box="[241,634,1012,1039]" pageId="39" pageNumber="40" type="nomenclature">
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<paragraph id="8BE41551F151E029FF4A84815C18154E" blockId="39.[113,764,1012,1952]" box="[241,634,1012,1039]" pageId="39" pageNumber="40">
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<emphasis id="B92FC943F151E029FF4A84815C18154E" bold="true" box="[241,634,1012,1039]" pageId="39" pageNumber="40">
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Phylogenetic position of
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<taxonomicName id="4C5B6ED2F151E029FE4384815C18154E" authority="(Currie and Zhao 1993)" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[504,634,1012,1039]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
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<emphasis id="B92FC943F151E029FE4384815C18154E" bold="true" box="[504,634,1012,1039]" italics="true" pageId="39" pageNumber="40">Alpkarakush</emphasis>
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</taxonomicName>
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</emphasis>
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</paragraph>
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<subSubSection id="C34146DAF151E025FFCA836E5F7C1554" lastPageId="43" lastPageNumber="44" pageId="39" pageNumber="40" type="description">
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<paragraph id="8BE41551F151E029FFCA836E5CCC14EB" blockId="39.[113,764,1012,1952]" pageId="39" pageNumber="40">
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The phylogenetic analysis using equal weights resulted in the recovery of 12 154 equally parsimonious trees with a length of 4792 steps. The strict consensus tree of these trees (Supporting information,
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<figureCitation id="136009D4F151E029FEBB830C5F2C15D0" box="[256,334,1145,1169]" captionStart="Figure 1" captionStartId="2.[133,198,1925,1949]" captionTargetBox="[134,1471,487,1894]" captionTargetId="figure-200@2.[131,1475,483,1897]" captionTargetPageId="2" captionText="Figure 1.Geographic location of the locality FTU-1 within Kyrgyzstan (A, B) and general geological map of the area around the city of Tashkumyr (C)." figureDoi="http://doi.org/10.5281/zenodo.13774243" httpUri="https://zenodo.org/record/13774243/files/figure.png" pageId="39" pageNumber="40">Fig. S1</figureCitation>
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) shows a reasonably good resolution, but with a major polytomy at the base of Tetanurae, in which most basal tetanuran taxa were unresolved. Reduced consensus methods removed numerous taxa at the base of Tetanurae, and the following description was based on the reduced consensus tree. As our matrix was mainly designed to resolve the phylogenetic position of
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<taxonomicName id="4C5B6ED2F151E029FEF282405FA4140C" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[329,454,1333,1357]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
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<emphasis id="B92FC943F151E029FEF282405FA4140C" box="[329,454,1333,1357]" italics="true" pageId="39" pageNumber="40">Alpkarakush</emphasis>
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</taxonomicName>
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within basal tetanurans, we will focus on this part of the tree; unexpected results, e.g. in the Coelurosauria, might at least partially be caused by incomplete taxon and/or character sampling for this part of the tree.
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</paragraph>
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<paragraph id="8BE41551F151E029FF3682C75B571515" blockId="39.[113,764,1012,1952]" lastBlockId="39.[809,1461,144,1954]" pageId="39" pageNumber="40">
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The results of our analysis (Supporting information,
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||
<figureCitation id="136009D4F151E029FD6F82C75EEE14A8" captionStart="Figure 2" captionStartId="3.[113,178,1836,1860]" captionTargetBox="[179,1392,144,1808]" captionTargetId="figure-6@3.[179,1392,144,1808]" captionTargetPageId="3" captionText="Figure 2. Excavation site FTU-1. A, position of the site within the Uurusai Valley west of Tashkumyr (red arrow). B, overview of the site, with one of the authors (A.E.F.) for scale.C, section of the excavation site. Black line indicates approximate dip of the sediments.D, quadratojugal of the new theropod as discovered in the field.E, metatarsal III of the new theropod during the excavation." figureDoi="http://doi.org/10.5281/zenodo.13774245" httpUri="https://zenodo.org/record/13774245/files/figure.png" pageId="39" pageNumber="40">Fig. S2</figureCitation>
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||
;
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||
<figureCitation id="136009D4F151E029FF2582A45E8A14A8" box="[158,232,1489,1513]" captionStart="Figure 25" captionStartId="38.[128,193,1776,1800]" captionTargetBox="[131,1471,146,1745]" captionTargetId="figure-88@38.[129,1473,144,1748]" captionTargetPageId="38" captionText="Figure 25. Phylogenetic position of A. kyrgyzicus. Simplified reduced consensus tree resulting from the equally weighted parsimony analysis. For full results see Supporting information, Fig. S2." figureDoi="http://doi.org/10.5281/zenodo.13774293" httpUri="https://zenodo.org/record/13774293/files/figure.png" pageId="39" pageNumber="40">Fig. 25</figureCitation>
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) mainly conformed to other recent phylogenetic analyses of theropod dinosaurs, with a monophyletic clade Coelophysoidea at the base of Neotheropoda, followed by monophyletic Ceratosauria and Tetanurae.
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<taxonomicName id="4C5B6ED2F151E029FD91815A5CAD1706" authorityName="Hammer & Hickerson" authorityYear="1994" box="[554,719,1583,1607]" class="Reptilia" family="Hadrosauridae" genus="Cryolophosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
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<emphasis id="B92FC943F151E029FD91815A5CAD1706" box="[554,719,1583,1607]" italics="true" pageId="39" pageNumber="40">Cryolophosaurus</emphasis>
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</taxonomicName>
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and ‘
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<taxonomicName id="4C5B6ED2F151E029FFCC813A5F671726" authorityName="Welles" authorityYear="1970" box="[119,261,1615,1639]" class="Reptilia" family="Troodontidae" genus="Dilophosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
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<emphasis id="B92FC943F151E029FFCC813A5F671726" box="[119,261,1615,1639]" italics="true" pageId="39" pageNumber="40">Dilophosaurus</emphasis>
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</taxonomicName>
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’
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<emphasis id="B92FC943F151E029FEA8813A5F391727" box="[275,347,1615,1638]" italics="true" pageId="39" pageNumber="40">sinensis</emphasis>
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(considered to be a junior synonym of
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<taxonomicName id="4C5B6ED2F151E029FFCA811B5F5A17C7" authorityName="Young" authorityYear="1940" box="[113,312,1646,1670]" family="Herrerasauridae" genus="Sinosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="39" pageNumber="40" phylum="Chordata" rank="species" species="triassicus">
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<emphasis id="B92FC943F151E029FFCA811B5F5A17C7" box="[113,312,1646,1670]" italics="true" pageId="39" pageNumber="40">Sinosaurus triassicus</emphasis>
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</taxonomicName>
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by some authors; see
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<bibRefCitation id="EFCA68A0F151E029FDA9811B5CCC17C7" author="Xing L & Paulina Carabajal A & Currie PJ" box="[530,686,1646,1670]" pageId="39" pageNumber="40" pagination="1653 - 61" refId="ref40604" refString="Xing L, Paulina Carabajal A, Currie PJ et al. Braincase anatomy of the basal theropod Sinosaurus from the Early Jurassic of China. Acta Geological Sinica 2014; 88: 1653 - 61." type="journal article" year="2014">
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Xing
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<emphasis id="B92FC943F151E029FDF3811A5C1217C7" box="[584,624,1646,1670]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
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. 2014
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</bibRefCitation>
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||
,
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<bibRefCitation id="EFCA68A0F151E029FD03811B5EBA17E4" author="Zhang Z-C & Wang T & You H-L" pageId="39" pageNumber="40" pagination="857 - 71" refId="ref41274" refString="Zhang Z-C, Wang T, You H-L. A new specimen of Sinosaurus triassicus (Dinosauria: Theropoda) from the Early Jurassic of Lufeng, Yunnan, China. Historical Biology 2023; 36: 857 - 71. https: // doi. org / 10.1080 / 08912963.2023. 2190760" type="journal article" year="2023">
|
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Zhang
|
||
<emphasis id="B92FC943F151E029FFCA81FB5EFB17E4" box="[113,153,1677,1701]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. 2023
|
||
</bibRefCitation>
|
||
), which were sometimes regarded as early Tetanurans (e.g.
|
||
<bibRefCitation id="EFCA68A0F151E029FF1D81D85F171784" author="Carrano MT & Benson RBJ & Sampson SD" box="[166,373,1709,1733]" pageId="39" pageNumber="40" refId="ref35875" refString="Carrano MT, Benson RBJ, Sampson SD. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology 2012; 10: 599. https: // doi. org / 10.1080 / 14772019.2012.713753" type="journal volume" year="2012">
|
||
Carrano
|
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<emphasis id="B92FC943F151E029FEBE81D85F531784" box="[261,305,1709,1733]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. 2012
|
||
</bibRefCitation>
|
||
) or, in the case of ‘
|
||
<emphasis id="B92FC943F151E029FDF581D85C031785" box="[590,609,1709,1732]" italics="true" pageId="39" pageNumber="40">D</emphasis>
|
||
’.
|
||
<emphasis id="B92FC943F151E029FDC981D85CD81785" box="[626,698,1709,1732]" italics="true" pageId="39" pageNumber="40">sinensis</emphasis>
|
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, early ceratosaurs (
|
||
<bibRefCitation id="EFCA68A0F151E029FF4181B95FD117A5" author="Wang S & Stiegler J & Amiot R" box="[250,435,1740,1764]" pageId="39" pageNumber="40" pagination="144 - 8" refId="ref40468" refString="Wang S, Stiegler J, Amiot R et al. Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology 2017; 27: 144 - 8." type="journal article" year="2017">
|
||
Wang
|
||
<emphasis id="B92FC943F151E029FEFB81B85F0C17A5" box="[320,366,1740,1764]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. 2017
|
||
</bibRefCitation>
|
||
), were here found to be outside Averostra, as subsequent sister taxa to this clade. Within Tetanurae, the poorly known ‘
|
||
<taxonomicName id="4C5B6ED2F151E029FE00807E5C031662" authorityName="Young" authorityYear="1942" box="[443,609,1803,1827]" class="Reptilia" family="Allosauridae" genus="Szechuanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FE00807E5C031662" box="[443,609,1803,1827]" italics="true" pageId="39" pageNumber="40">Szechuanosaurus</emphasis>
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||
</taxonomicName>
|
||
’
|
||
<emphasis id="B92FC943F151E029FDC8807E5CBD1663" box="[627,735,1803,1826]" italics="true" pageId="39" pageNumber="40">zigongensis</emphasis>
|
||
is found as the earliest branching taxon, followed by a dichotomy of two major clades, the Coelurosauria and Carnosauria (
|
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<emphasis id="B92FC943F151E029FD7C803F5C981620" box="[711,762,1866,1889]" italics="true" pageId="39" pageNumber="40">sensu</emphasis>
|
||
<bibRefCitation id="EFCA68A0F151E029FFCA801C5F2A16C0" author="Rauhut OWM & Pol D" box="[113,328,1897,1921]" pageId="39" pageNumber="40" refId="ref39277" refString="Rauhut OWM, Pol D. Probable basal allosauroid from the early Middle Jurassic Canadon Asfalto Formation of Argentina highlights phylogenetic uncertainty in tetanuran theropod dinosaurs. Scientific Reports 2019; 9: 18826. https: // doi. org / 10.1038 / s 41598 - 019 - 53672 - 7" type="journal volume" year="2019">Rauhut and Pol 2019</bibRefCitation>
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), the latter including Megalosauroidea and Allosauroidea. This result thus resembles that found by
|
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<bibRefCitation id="EFCA68A0F151E029FD1480FD5D1611E9" author="Rauhut OWM" box="[687,884,144,1952]" pageId="39" pageNumber="40" pagination="1 - 213" refId="ref39053" refString="Rauhut OWM. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology 2003; 69: 1 - 213." type="journal article" year="2003">Rauhut (2003)</bibRefCitation>
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,
|
||
<bibRefCitation id="EFCA68A0F151E029FCC487E55D9A11E9" author="Cau A" box="[895,1016,144,168]" pageId="39" pageNumber="40" pagination="1 - 25" refId="ref36065" refString="Cau A. The assembly of the avian body plan: a 160 - million-year long process. Bollettino della Societa Paleontologica Italiana 2018; 57: 1 - 25." type="journal article" year="2018">Cau (2018)</bibRefCitation>
|
||
, and
|
||
<bibRefCitation id="EFCA68A0F151E029FB9787E55B7A11E9" author="Rauhut OWM & Pol D" box="[1068,1304,144,168]" pageId="39" pageNumber="40" refId="ref39277" refString="Rauhut OWM, Pol D. Probable basal allosauroid from the early Middle Jurassic Canadon Asfalto Formation of Argentina highlights phylogenetic uncertainty in tetanuran theropod dinosaurs. Scientific Reports 2019; 9: 18826. https: // doi. org / 10.1038 / s 41598 - 019 - 53672 - 7" type="journal volume" year="2019">Rauhut and Pol (2019)</bibRefCitation>
|
||
, whereas many recent phylogenetic analyses found the Megalosauroidea to be sister taxon of a clade uniting Allosauroidea and Coelurosauria, in a clade termed Neotetanurae or
|
||
<taxonomicName id="4C5B6ED2F151E029FB07879B5B2F1047" box="[1212,1357,238,262]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Avetheropoda" pageId="39" pageNumber="40" phylum="Chordata" rank="order">Avetheropoda</taxonomicName>
|
||
(see e.g.
|
||
<bibRefCitation id="EFCA68A0F151E029FC9186785D851064" author="Carrano MT & Benson RBJ & Sampson SD" box="[810,999,269,293]" pageId="39" pageNumber="40" refId="ref35875" refString="Carrano MT, Benson RBJ, Sampson SD. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology 2012; 10: 599. https: // doi. org / 10.1080 / 14772019.2012.713753" type="journal volume" year="2012">
|
||
Carrano
|
||
<emphasis id="B92FC943F151E029FC39867B5DCB1064" box="[898,937,269,293]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. 2012
|
||
</bibRefCitation>
|
||
, Rauhut
|
||
<emphasis id="B92FC943F151E029FBFB867B5A051064" box="[1088,1127,269,293]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. 2016,
|
||
<bibRefCitation id="EFCA68A0F151E029FB1586785B581064" author="Dai H & Benson RBJ & Hu X" box="[1198,1338,269,293]" pageId="39" pageNumber="40" refId="ref36710" refString="Dai H, Benson RBJ, Hu X et al. A new possible megalosauroid theropod from the Middle Jurassic Xintiangou Formation of Chongqing, People's Republic of China and its implication for early tetanuran evolution. Scientific Reports 2020; 10: 139. https: // doi. org / 10.1038 / s 41598 - 019 - 56959 - x" type="journal volume" year="2020">
|
||
Dai
|
||
<emphasis id="B92FC943F151E029FB6D867B5A9F1064" box="[1238,1277,269,293]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. 2020
|
||
</bibRefCitation>
|
||
), and
|
||
<bibRefCitation id="EFCA68A0F151E029FACC867B5DD21004" author="Wang S & Stiegler J & Amiot R" pageId="39" pageNumber="40" pagination="144 - 8" refId="ref40468" refString="Wang S, Stiegler J, Amiot R et al. Extreme ontogenetic changes in a ceratosaurian theropod. Current Biology 2017; 27: 144 - 8." type="journal article" year="2017">
|
||
Wang
|
||
<emphasis id="B92FC943F151E029FC9186585D351004" box="[810,855,301,325]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. (2017)
|
||
</bibRefCitation>
|
||
even found the Allosauroidea of the above analyses to be paraphyletic, with the
|
||
<taxonomicName id="4C5B6ED2F151E029FBC086395B391025" authorityName="Paul" authorityYear="1988" box="[1147,1371,332,356]" class="Reptilia" family="Metriacanthosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="39" pageNumber="40" phylum="Chordata" rank="family">Metriacanthosauridae</taxonomicName>
|
||
forming the sister taxon to an Allosauria-Coelurosauria clade. Within Carnosauria, the taxa usually grouped in the
|
||
<taxonomicName id="4C5B6ED2F151E029FB4C86FE5BD110E2" box="[1271,1459,395,419]" family="Piatnitzkysauridae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="39" pageNumber="40" phylum="Chordata" rank="family">Piatnitzkysauridae</taxonomicName>
|
||
(
|
||
<taxonomicName id="4C5B6ED2F151E029FC8E86DF5DDD1083" authorityName="Madsen" authorityYear="1976" box="[821,959,426,450]" class="Reptilia" family="Piatnitzkysauridae" genus="Marshosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC8E86DF5DDD1083" box="[821,959,426,450]" italics="true" pageId="39" pageNumber="40">Marshosaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F151E029FC6386DF5A1E1083" authorityName="Bonaparte" authorityYear="1979" box="[984,1148,426,450]" class="Reptilia" family="Allosauridae" genus="Piatnitzkysaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC6386DF5A1E1083" box="[984,1148,426,450]" italics="true" pageId="39" pageNumber="40">Piatnitzkysaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F151E029FB2E86DF5B7E1083" baseAuthorityName="Rauhut" baseAuthorityYear="2005" box="[1173,1308,426,450]" class="Reptilia" family="Piatnitzkysauridae" genus="Condorraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FB2E86DF5B7E1083" box="[1173,1308,426,450]" italics="true" pageId="39" pageNumber="40">Condorraptor</emphasis>
|
||
</taxonomicName>
|
||
, and, sometimes,
|
||
<taxonomicName id="4C5B6ED2F151E029FCD686BC5A7010A0" box="[877,1042,457,481]" class="Reptilia" family="Piatnitzkysauridae" genus="Xuanhanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FCD686BC5A7010A0" box="[877,1042,457,481]" italics="true" pageId="39" pageNumber="40">Xuanhanosaurus</emphasis>
|
||
</taxonomicName>
|
||
) were all removed by reduced consensus methods, as was the supposed early allosauroid
|
||
<taxonomicName id="4C5B6ED2F151E029FA9B869C5BCF1340" authorityName="Rauhut & Pol" authorityYear="2019" box="[1312,1453,489,513]" genus="Asfaltovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FA9B869C5BCF1340" box="[1312,1453,489,513]" italics="true" pageId="39" pageNumber="40">Asfaltovenator</emphasis>
|
||
</taxonomicName>
|
||
. The remaining carnosaurs were thus placed either in the Megalosauroidea or the Allosauroidea, which form sister taxa. The taxonomic composition of the two clades largely conformed to that found in other recent analysis, with a few notable exceptions.
|
||
<taxonomicName id="4C5B6ED2F151E029FC6E85F05AE013DC" baseAuthorityName="Zhao and Currie" baseAuthorityYear="1993" box="[981,1154,645,669]" class="Reptilia" genus="Monolophosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC6E85F05AE013DC" box="[981,1154,645,669]" italics="true" pageId="39" pageNumber="40">Monolophosaurus</emphasis>
|
||
</taxonomicName>
|
||
, which has been recovered in varying positions within basal tetanurans in recent years, was placed in the Megalosauroidea. In respect to Allosauroidea, the megaraptorans, which were found to be derived allosauroids within the
|
||
<taxonomicName id="4C5B6ED2F151E029FC1984765A27125A" box="[930,1093,771,795]" class="Reptilia" family="Neovenatoridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="family">Neovenatoridae</taxonomicName>
|
||
by
|
||
<bibRefCitation id="EFCA68A0F151E029FBC984765B33125A" author="Benson RBJ & Carrano MT & Brusatte SL" box="[1138,1361,771,795]" pageId="39" pageNumber="40" pagination="71 - 8" refId="ref35034" refString="Benson RBJ, Carrano MT, Brusatte SL. A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the Latest Mesozoic. Naturwissenschaften 2010; 97: 71 - 8. https: // doi. org / 10.1007 / s 00114 - 009 - 0614 - x" type="journal article" year="2010">
|
||
Benson
|
||
<emphasis id="B92FC943F151E029FB7284765A95125A" box="[1225,1271,771,795]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. (2010)
|
||
</bibRefCitation>
|
||
and several subsequent analyses (e.g.
|
||
<bibRefCitation id="EFCA68A0F151E029FBDD84575B50127B" author="Carrano MT & Benson RBJ & Sampson SD" box="[1126,1330,802,826]" pageId="39" pageNumber="40" refId="ref35875" refString="Carrano MT, Benson RBJ, Sampson SD. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology 2012; 10: 599. https: // doi. org / 10.1080 / 14772019.2012.713753" type="journal volume" year="2012">
|
||
Carrano
|
||
<emphasis id="B92FC943F151E029FB7884565A8D127B" box="[1219,1263,802,826]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. 2012
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFCA68A0F151E029FAFB84575DB51218" author="Zanno LE & Makovicky PJ" pageId="39" pageNumber="40" refId="ref41194" refString="Zanno LE, Makovicky PJ. Neovenatorid theropods are apex predators in the Late Cretaceous of North America. Nature Communications 2013; 4: 2827. https: // doi. org / 10.1038 / ncomms 3827" type="journal volume" year="2013">Zanno and Makovicky 2013</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFCA68A0F151E029FC5F84345AD21218" author="Malafaia E & Mocho P & Escaso F" box="[996,1200,833,857]" pageId="39" pageNumber="40" refId="ref38378" refString="Malafaia E, Mocho P, Escaso F et al. A new carcharodontosaurian theropod from the Lusitanian Basin: evidence of allosauroid sympatry in the European Late Jurassic. Journal of Vertebrate Paleontology 2020; 40: e 1768106. https: // doi. org / 10.1080 / 02724634.2020.1768 106" type="journal volume" year="2020">
|
||
Malafaia
|
||
<emphasis id="B92FC943F151E029FBF884375A0C1218" box="[1091,1134,833,857]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. 2020
|
||
</bibRefCitation>
|
||
) were placed as an early branching clade within Tyrannosauroidea, as argued by
|
||
<bibRefCitation id="EFCA68A0F151E029FAC984145DC812D9" author="Novas FE & Agnolin FL & Ezcurra MD" pageId="39" pageNumber="40" pagination="174 - 215" refId="ref38768" refString="Novas FE, Agnolin FL, Ezcurra MD et al. Evolution of the carnivorous dinosaurs during the Cretaceous: the evidence from Patagonia. Cretaceous Research 2013; 45: 174 - 215. https: // doi. org / 10.1016 / j. cretres. 2013.04.001" type="journal article" year="2013">
|
||
Novas
|
||
<emphasis id="B92FC943F151E029FC9184F45D3112D9" box="[810,851,896,920]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. (2013)
|
||
</bibRefCitation>
|
||
and subsequent authors. Likewise,
|
||
<taxonomicName id="4C5B6ED2F151E029FAAC84F55BE912D9" box="[1303,1419,896,920]" class="Reptilia" family="Carcharodontosauridae" genus="Shaochilong" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FAAC84F55BE912D9" box="[1303,1419,896,920]" italics="true" pageId="39" pageNumber="40">Shaochilong</emphasis>
|
||
</taxonomicName>
|
||
, regarded as a carcharodontosaurid in recent years (
|
||
<bibRefCitation id="EFCA68A0F151E029FA9C84D55D3C1296" author="Brusatte SL & Benson RBJ & Chure DJ" pageId="39" pageNumber="40" pagination="1051 - 8" refId="ref35429" refString="Brusatte SL, Benson RBJ, Chure DJ et al. The first definitive carcharodontosaurid (Dinosauria: Theropoda) from Asia and the delayed ascent of tyrannosaurids. Naturwissenschaften 2009; 96: 1051 - 8. https: // doi. org / 10.1007 / s 00114 - 009 - 0565 - 2" type="journal article" year="2009">
|
||
Brusatte
|
||
<emphasis id="B92FC943F151E029FA3884D55BCF12F6" box="[1411,1453,927,951]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. 2009
|
||
</bibRefCitation>
|
||
, 2010,
|
||
<bibRefCitation id="EFCA68A0F151E029FC1384CA5A081296" author="Carrano MT & Benson RBJ & Sampson SD" box="[936,1130,959,983]" pageId="39" pageNumber="40" refId="ref35875" refString="Carrano MT, Benson RBJ, Sampson SD. The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology 2012; 10: 599. https: // doi. org / 10.1080 / 14772019.2012.713753" type="journal volume" year="2012">
|
||
Carrano
|
||
<emphasis id="B92FC943F151E029FBB984CA5A491296" box="[1026,1067,959,983]" italics="true" pageId="39" pageNumber="40">et al</emphasis>
|
||
. 2012
|
||
</bibRefCitation>
|
||
), was found to be a coelurosaur, as in some previous analyses (e.g.
|
||
<bibRefCitation id="EFCA68A0F151E029FB3784AB5B7312B7" author="Rauhut OWM" box="[1164,1297,990,1014]" pageId="39" pageNumber="40" pagination="1 - 213" refId="ref39053" refString="Rauhut OWM. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology 2003; 69: 1 - 213." type="journal article" year="2003">Rauhut 2003</bibRefCitation>
|
||
). Furthermore, the poorly known Middle Jurassic French taxon
|
||
<taxonomicName id="4C5B6ED2F151E029FA9884885BD11554" box="[1315,1459,1021,1045]" class="Reptilia" family="Megalosauridae" genus="Poekilopleuron" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FA9884885BD11554" box="[1315,1459,1021,1045]" italics="true" pageId="39" pageNumber="40">Poekilopleuron</emphasis>
|
||
</taxonomicName>
|
||
was found to be an early carcharodontosaurid, rather than a megalosaurid, as is the case in most recent analyses.
|
||
</paragraph>
|
||
<paragraph id="8BE41551F151E029FCFE832E5B32174A" blockId="39.[809,1461,144,1954]" pageId="39" pageNumber="40">
|
||
Within megalosauroids, two subclades were recovered, the
|
||
<taxonomicName id="4C5B6ED2F151E029FCEC830E5D8A15D2" authorityName="Stromer" authorityYear="1915" box="[855,1000,1147,1171]" class="Reptilia" family="Spinosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="family">Spinosauridae</taxonomicName>
|
||
and the
|
||
<taxonomicName id="4C5B6ED2F151E029FBE9830E5A9415D2" box="[1106,1270,1147,1171]" class="Reptilia" family="Megalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="family">Megalosauridae</taxonomicName>
|
||
.
|
||
<taxonomicName id="4C5B6ED2F151E029FABE830E5BD115D2" baseAuthorityName="Zhao and Currie" baseAuthorityYear="1993" box="[1285,1459,1147,1171]" class="Reptilia" genus="Monolophosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FABE830E5BD115D2" box="[1285,1459,1147,1171]" italics="true" pageId="39" pageNumber="40">Monolophosaurus</emphasis>
|
||
</taxonomicName>
|
||
was found as the earliest branching spinosaurid, as in the equal weights analysis of
|
||
<bibRefCitation id="EFCA68A0F151E029FB8C83CC5B6F1590" author="Schade M & Rauhut OWM & Foth C" box="[1079,1293,1209,1233]" pageId="39" pageNumber="40" refId="ref39872" refString="Schade M, Rauhut OWM, Foth C et al. A reappraisal of the cranial osteology of the spinosaurid Irritator challengeri (Dinosauria: Theropoda). Palaeontologia Electronica 2023; 26: a 17." type="journal volume" year="2023">
|
||
Schade
|
||
<emphasis id="B92FC943F151E029FB3283CF5AD91590" box="[1161,1211,1209,1233]" italics="true" pageId="39" pageNumber="40">et al.</emphasis>
|
||
(2023)
|
||
</bibRefCitation>
|
||
. The remaining spinosaurids were subdivided into the Baryonichinae (
|
||
<taxonomicName id="4C5B6ED2F151E029FAF483AC5BCF15B1" authorityName="Charig & Milner" authorityYear="1986" box="[1359,1453,1241,1264]" class="Reptilia" family="Spinosauridae" genus="Baryonyx" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FAF483AC5BCF15B1" box="[1359,1453,1241,1264]" italics="true" pageId="39" pageNumber="40">Baryonyx</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F151E029FC91838D5DC71451" box="[810,933,1272,1296]" class="Reptilia" family="Spinosauridae" genus="Suchomimus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC91838D5DC71451" box="[810,933,1272,1296]" italics="true" pageId="39" pageNumber="40">Suchomimus</emphasis>
|
||
</taxonomicName>
|
||
), and the
|
||
<taxonomicName id="4C5B6ED2F151E029FBB5838D5AFD1451" box="[1038,1183,1272,1296]" class="Reptilia" family="Spinosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="subFamily" subFamily="Spinosaurinae">Spinosaurinae</taxonomicName>
|
||
(with a polytomy between
|
||
<taxonomicName id="4C5B6ED2F151E029FC91826D5D18146E" box="[810,890,1304,1327]" class="Reptilia" family="Spinosauridae" genus="Irritator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC91826D5D18146E" box="[810,890,1304,1327]" italics="true" pageId="39" pageNumber="40">Irritator</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F151E029FC3F82625A76146E" authorityName="Allain, Xaisanavong, Richir & Khentavong" authorityYear="2012" box="[900,1044,1303,1327]" family="Spinosauridae" genus="Ichthyovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC3F82625A76146E" box="[900,1044,1303,1327]" italics="true" pageId="39" pageNumber="40">Ichthyovenator</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C5B6ED2F151E029FBF9826D5ADA146E" box="[1090,1208,1304,1327]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FBF9826D5ADA146E" box="[1090,1208,1304,1327]" italics="true" pageId="39" pageNumber="40">Spinosaurus</emphasis>
|
||
</taxonomicName>
|
||
). Within megalosaurids, a polytomy was recovered between a
|
||
<emphasis id="B92FC943F151E029FAA182425DC0142F" italics="true" pageId="39" pageNumber="40">
|
||
<taxonomicName id="4C5B6ED2F151E029FAA182425BC9140E" box="[1306,1451,1335,1359]" family="Megalosaurinidae" genus="Wiehenvenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">Wiehenvenator</taxonomicName>
|
||
-
|
||
<taxonomicName id="4C5B6ED2F151E029FC9182225DC0142F" box="[810,930,1367,1390]" class="Reptilia" family="Megalosauridae" genus="Torvosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">Torvosaurus</taxonomicName>
|
||
</emphasis>
|
||
clade, a
|
||
<emphasis id="B92FC943F151E029FBAC82235B7B142F" box="[1047,1305,1366,1390]" italics="true" pageId="39" pageNumber="40">
|
||
<taxonomicName id="4C5B6ED2F151E029FBAC82235AFF142F" baseAuthorityName="Benson" baseAuthorityYear="2010" box="[1047,1181,1366,1390]" class="Reptilia" family="Megalosauridae" genus="Megalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">Megalosaurus</taxonomicName>
|
||
-
|
||
<taxonomicName id="4C5B6ED2F151E029FB1E82235B7B142F" box="[1189,1305,1366,1390]" class="Reptilia" family="Megalosauridae" genus="Afrovenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">Afrovenator</taxonomicName>
|
||
</emphasis>
|
||
clade, and a clade that recovered
|
||
<taxonomicName id="4C5B6ED2F151E029FBB082005AD014CC" authorityName="Allain" authorityYear="2005" box="[1035,1202,1397,1421]" class="Reptilia" family="Megalosauridae" genus="Dubreuillosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FBB082005AD014CC" box="[1035,1202,1397,1421]" italics="true" pageId="39" pageNumber="40">Dubreuillosaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F151E029FB7982005B1E14CC" authorityName="Walker" authorityYear="1964" box="[1218,1404,1397,1421]" class="Reptilia" family="Megalosauridae" genus="Eustreptospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FB7982005B1E14CC" box="[1218,1404,1397,1421]" italics="true" pageId="39" pageNumber="40">Eustreptospondylus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C5B6ED2F151E029FC9182E05DAC14EC" authorityName="Meyer" authorityYear="1830" box="[810,974,1429,1453]" class="Reptilia" genus="Streptospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC9182E05DAC14EC" box="[810,974,1429,1453]" italics="true" pageId="39" pageNumber="40">Streptospondylus</emphasis>
|
||
</taxonomicName>
|
||
in a polytomy. Allosauroidea were subdivided into
|
||
<taxonomicName id="4C5B6ED2F151E029FC2A82C15A13148D" authorityName="Paul" authorityYear="1988" box="[913,1137,1460,1484]" class="Reptilia" family="Metriacanthosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="39" pageNumber="40" phylum="Chordata" rank="family">Metriacanthosauridae</taxonomicName>
|
||
and Allosauria, with the latter including
|
||
<taxonomicName id="4C5B6ED2F151E029FC2B82A65D9514AA" baseAuthorityName="Madsen" baseAuthorityYear="1976" box="[912,1015,1491,1515]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC2B82A65D9514AA" box="[912,1015,1491,1515]" italics="true" pageId="39" pageNumber="40">Allosaurus</emphasis>
|
||
</taxonomicName>
|
||
, the
|
||
<taxonomicName id="4C5B6ED2F151E029FB9182A65AAF14AA" box="[1066,1229,1491,1515]" class="Reptilia" family="Neovenatoridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="family">Neovenatoridae</taxonomicName>
|
||
(including
|
||
<taxonomicName id="4C5B6ED2F151E029FAFF82A15BD614AA" box="[1348,1460,1492,1515]" class="Reptilia" family="Carcharodontosauridae" genus="Neovenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FAFF82A15BD614AA" box="[1348,1460,1492,1515]" italics="true" pageId="39" pageNumber="40">Neovenator</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C5B6ED2F151E029FCED82865D93174A" authorityName="Hu" authorityYear="1964" box="[854,1009,1523,1547]" class="Reptilia" family="Allosauridae" genus="Chilantaisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FCED82865D93174A" box="[854,1009,1523,1547]" italics="true" pageId="39" pageNumber="40">Chilantaisaurus</emphasis>
|
||
</taxonomicName>
|
||
), and the
|
||
<taxonomicName id="4C5B6ED2F151E029FBE382865B2E174A" authorityName="Stromer" authorityYear="1931" box="[1112,1356,1523,1547]" class="Reptilia" family="Carcharodontosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="family">Carcharodontosauridae</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="8BE41551F151E029FCFE81675B1F16E0" blockId="39.[809,1461,144,1954]" pageId="39" pageNumber="40">
|
||
<taxonomicName id="4C5B6ED2F151E029FCFE81675DA0176B" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[837,962,1554,1578]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FCFE81675DA0176B" box="[837,962,1554,1578]" italics="true" pageId="39" pageNumber="40">Alpkarakush</emphasis>
|
||
</taxonomicName>
|
||
was found to be a metriacanthosaurid. In general, metriacanthosaurid interrelationships were poorly resolved, with only two distinct subclades within this clade, one consisting of the two species of the genus
|
||
<taxonomicName id="4C5B6ED2F151E029FB7181055B1C17C9" box="[1226,1406,1648,1672]" class="Reptilia" family="Ceratosauridae" genus="Yangchuanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FB7181055B1C17C9" box="[1226,1406,1648,1672]" italics="true" pageId="39" pageNumber="40">Yangchuanosaurus</emphasis>
|
||
</taxonomicName>
|
||
, and the other, conforming to the subfamily
|
||
<taxonomicName id="4C5B6ED2F151E029FB7581FA5BD217E6" box="[1230,1456,1679,1703]" class="Reptilia" family="Metriacanthosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="39" pageNumber="40" phylum="Chordata" rank="subFamily" subFamily="Metriacanthosaurinae">Metriacanthosaurinae</taxonomicName>
|
||
, of a polytomy including
|
||
<taxonomicName id="4C5B6ED2F151E029FBCB81DA5AAE1787" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[1136,1228,1711,1734]" class="Reptilia" family="Sinraptoridae" genus="Sinraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FBCB81DA5AAE1787" box="[1136,1228,1711,1734]" italics="true" pageId="39" pageNumber="40">Sinraptor</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F151E029FB5081DA5BCF1786" box="[1259,1453,1711,1735]" class="Reptilia" family="Megalosauridae" genus="Metriacanthosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FB5081DA5BCF1786" box="[1259,1453,1711,1735]" italics="true" pageId="39" pageNumber="40">Metriacanthosaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F151E029FC9181BB5DDF17A4" box="[810,957,1742,1765]" family="Metriacanthosauridae" genus="Siamotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC9181BB5DDF17A4" box="[810,957,1742,1765]" italics="true" pageId="39" pageNumber="40">Siamotyrannus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C5B6ED2F151E029FC4381BB5A1117A7" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[1016,1139,1742,1766]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC4381BB5A1117A7" box="[1016,1139,1742,1766]" italics="true" pageId="39" pageNumber="40">Alpkarakush</emphasis>
|
||
</taxonomicName>
|
||
. The other taxon found within
|
||
<taxonomicName id="4C5B6ED2F151E029FC9781985A6C1644" authorityName="Paul" authorityYear="1988" box="[812,1038,1773,1797]" class="Reptilia" family="Metriacanthosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="39" pageNumber="40" phylum="Chordata" rank="family">Metriacanthosauridae</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F151E029FBA281985AF51644" box="[1049,1175,1773,1797]" family="Metriacanthosauridae" genus="Shidaisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FBA281985AF51644" box="[1049,1175,1773,1797]" italics="true" pageId="39" pageNumber="40">Shidaisaurus</emphasis>
|
||
</taxonomicName>
|
||
, could attain various positions in relation to these clades and their subtaxa.
|
||
<taxonomicName id="4C5B6ED2F151E029FA8C80785BD71664" box="[1335,1461,1805,1829]" family="Metriacanthosauridae" genus="Shidaisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FA8C80785BD71664" box="[1335,1461,1805,1829]" italics="true" pageId="39" pageNumber="40">Shidaisaurus</emphasis>
|
||
</taxonomicName>
|
||
was either found as an early branching metriacanthosaurid, outside the
|
||
<taxonomicName id="4C5B6ED2F151E029FC3A803E5A561622" box="[897,1076,1867,1891]" class="Reptilia" family="Ceratosauridae" genus="Yangchuanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC3A803E5A561622" box="[897,1076,1867,1891]" italics="true" pageId="39" pageNumber="40">Yangchuanosaurus</emphasis>
|
||
</taxonomicName>
|
||
-
|
||
<taxonomicName id="4C5B6ED2F151E029FB80803E5B7E1622" box="[1083,1308,1867,1891]" class="Reptilia" family="Metriacanthosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="39" pageNumber="40" phylum="Chordata" rank="subFamily" subFamily="Metriacanthosaurinae">Metriacanthosaurinae</taxonomicName>
|
||
clade, as sister taxon to the
|
||
<taxonomicName id="4C5B6ED2F151E029FC67801E5A5A16C3" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[988,1080,1899,1922]" class="Reptilia" family="Sinraptoridae" genus="Sinraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FC67801E5A5A16C3" box="[988,1080,1899,1922]" italics="true" pageId="39" pageNumber="40">Sinraptor</emphasis>
|
||
</taxonomicName>
|
||
/
|
||
<taxonomicName id="4C5B6ED2F151E029FBFE801E5B6516C2" box="[1093,1287,1899,1923]" class="Reptilia" family="Megalosauridae" genus="Metriacanthosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FBFE801E5B6516C2" box="[1093,1287,1899,1923]" italics="true" pageId="39" pageNumber="40">Metriacanthosaurus</emphasis>
|
||
</taxonomicName>
|
||
/
|
||
<emphasis id="B92FC943F151E029FAAF801E5DC516E3" italics="true" pageId="39" pageNumber="40">
|
||
<taxonomicName id="4C5B6ED2F151E029FAAF801E5BC516C3" box="[1300,1447,1899,1922]" family="Metriacanthosauridae" genus="Siamotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">Siamotyrannus</taxonomicName>
|
||
/
|
||
<taxonomicName id="4C5B6ED2F151E029FC9180FF5DC516E3" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[810,935,1930,1954]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">Alpkarakush</taxonomicName>
|
||
</emphasis>
|
||
clade, or as sister taxon to
|
||
<taxonomicName id="4C5B6ED2F151E029FB0F80FF5B1416E3" box="[1204,1398,1930,1954]" class="Reptilia" family="Megalosauridae" genus="Metriacanthosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="39" pageNumber="40" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F151E029FB0F80FF5B1416E3" box="[1204,1398,1930,1954]" italics="true" pageId="39" pageNumber="40">Metriacanthosaurus</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<caption id="DF2445D9F15EE026FF3A805E5CCB161F" ID-DOI="http://doi.org/10.5281/zenodo.13774295" ID-Zenodo-Dep="13774295" httpUri="https://zenodo.org/record/13774295/files/figure.png" pageId="40" pageNumber="41" startId="40.[129,194,1835,1859]" targetBox="[129,1473,144,1807]" targetPageId="40" targetType="figure">
|
||
<paragraph id="8BE41551F15EE026FF3A805E5CCB161F" blockId="40.[129,1464,1835,1887]" pageId="40" pageNumber="41">
|
||
<emphasis id="B92FC943F15EE026FF3A805E5E871602" bold="true" box="[129,229,1835,1859]" pageId="40" pageNumber="41">Figure 26.</emphasis>
|
||
Phylogenetic position of
|
||
<taxonomicName id="4C5B6ED2F15EE026FE6E805E5C2F1602" box="[469,589,1835,1859]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="40" pageNumber="41" phylum="Chordata" rank="species" species="kyrgyzicus">
|
||
<emphasis id="B92FC943F15EE026FE6E805E5C2F1602" box="[469,589,1835,1859]" italics="true" pageId="40" pageNumber="41">A. kyrgyzicus</emphasis>
|
||
</taxonomicName>
|
||
. Simplified strict consensus tree resulting from the implied weighing parsimony analysis with k = 12. For full results see Supporting information, Fig. S3.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BE41551F15FE027FF3687E55F4B14CC" blockId="41.[111,764,144,1954]" pageId="41" pageNumber="42">
|
||
The analyses using implied weights found few most parsimonious trees, with scores of 208.21175 (k = 12; 3 Most Parsimonious Trees [MPT]), 251.68722 (k = 9; 1 MPT), 319.46481 (k = 6; 5 MPT), and 443.57498 (k = 3; 30 MPT). The strict consensus tree resulting from the analysis with k = 12 (Supporting information,
|
||
<figureCitation id="136009D4F15FE027FDA986585C3C1004" box="[530,606,301,325]" captionStart="Figure 3" captionStartId="6.[130,195,1952,1976]" captionTargetBox="[165,1437,1530,1921]" captionTargetId="figure-591@6.[160,1443,1525,1924]" captionTargetPageId="6" captionText="Figure 3. Outline reconstruction of Alpkarakush kyrgyzicus, with recovered elements indicated. Scale bar is 1 m." figureDoi="http://doi.org/10.5281/zenodo.13774247" httpUri="https://zenodo.org/record/13774247/files/figure.png" pageId="41" pageNumber="42">Fig. S3</figureCitation>
|
||
;
|
||
<figureCitation id="136009D4F15FE027FDCA86585CDF1004" box="[625,701,301,325]" captionStart="Figure 26" captionStartId="40.[129,194,1835,1859]" captionTargetBox="[129,1473,144,1807]" captionTargetId="figure-8@40.[129,1473,144,1807]" captionTargetPageId="40" captionText="Figure 26. Phylogenetic position of A. kyrgyzicus. Simplified strict consensus tree resulting from the implied weighing parsimony analysis with k = 12. For full results see Supporting information, Fig. S3." figureDoi="http://doi.org/10.5281/zenodo.13774295" httpUri="https://zenodo.org/record/13774295/files/figure.png" pageId="41" pageNumber="42">Fig. 26</figureCitation>
|
||
) was in general agreement with the reduced consensus tree of the equally weighted analysis, but showed considerably better resolution. At higher weighing strengths, changes in topology were mainly found within Megalosauroidea, but also between some metriacanthosaurids and a few other taxa (Supporting information,
|
||
<figureCitation id="136009D4F15FE027FF72869C5F501340" box="[201,306,489,513]" captionStart="Figure 4" captionStartId="8.[129,194,1836,1860]" captionTargetBox="[180,1423,144,1808]" captionTargetId="figure-7@8.[180,1423,144,1808]" captionTargetPageId="8" captionText="Figure 4. Postorbital of A. kyrgyzicus. A–D, left postorbital IGB 2-1 in lateral (A, stereophotographs), posterior (B), medial (C), and dorsal (D, stereophotographs) views. E, F, right postorbital IGB 2-2 in lateral (E) and medial (F) views. Abbreviations:g, groove; jf, jugal facet; ls, laterosphenoid contact; ob, orbital brow; pp, posterior process; sf, supratemporal fossa; sq, squamosal facet; st, step. Scale bar is 5 cm." figureDoi="http://doi.org/10.5281/zenodo.13774249" httpUri="https://zenodo.org/record/13774249/files/figure.png" pageId="41" pageNumber="42">Figs S4–S</figureCitation>
|
||
<figureCitation id="136009D4F15FE027FE89869C5F5D1340" box="[306,319,489,513]" captionStart="Figure 6" captionStartId="11.[115,180,1629,1653]" captionTargetBox="[258,1314,144,1601]" captionTargetId="figure-178@11.[258,1314,144,1601]" captionTargetPageId="11" captionText="Figure 6. Teeth referred to of A. kyrgyzicus. A–C, possible premaxillary tooth IGB 2-6 in mesial (A), labial or lingual (B), and distal (C) views. D, E, lateral tooth IGB 2-3 in (?)lingual (D) and distal (E) views. Scale bar is 1 cm." figureDoi="http://doi.org/10.5281/zenodo.13774253" httpUri="https://zenodo.org/record/13774253/files/figure.png" pageId="41" pageNumber="42">6</figureCitation>
|
||
). The most severe reordering of taxa was found at k = 3, in which the
|
||
<taxonomicName id="4C5B6ED2F15FE027FE21857D5C5E1361" box="[410,572,520,544]" class="Reptilia" family="Megalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Megalosauridae</taxonomicName>
|
||
were found as the earliest branching tetanurans, followed by metriacanthosaurids and Allosauria, whereas the
|
||
<taxonomicName id="4C5B6ED2F15FE027FE3585325C7D131E" authorityName="Stromer" authorityYear="1915" box="[398,543,583,607]" class="Reptilia" family="Spinosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Spinosauridae</taxonomicName>
|
||
were recovered as the earliest branching coelurosaurs (Supporting information,
|
||
<figureCitation id="136009D4F15FE027FD6F85135EEE13DC" captionStart="Figure 6" captionStartId="11.[115,180,1629,1653]" captionTargetBox="[258,1314,144,1601]" captionTargetId="figure-178@11.[258,1314,144,1601]" captionTargetPageId="11" captionText="Figure 6. Teeth referred to of A. kyrgyzicus. A–C, possible premaxillary tooth IGB 2-6 in mesial (A), labial or lingual (B), and distal (C) views. D, E, lateral tooth IGB 2-3 in (?)lingual (D) and distal (E) views. Scale bar is 1 cm." figureDoi="http://doi.org/10.5281/zenodo.13774253" httpUri="https://zenodo.org/record/13774253/files/figure.png" pageId="41" pageNumber="42">Fig. S6</figureCitation>
|
||
). Most of these changes seem to reflect the fragmentary nature of the unstable taxa, for which changing weights in a single character might then have severe consequences for their phylogenetic position. This might especially be the reason for the unstable relationships within Megalosauroidea; of the 23 taxa included in this clade in the k = 12 analysis, only two have more than 50% coded characters (
|
||
<taxonomicName id="4C5B6ED2F15FE027FE8384345F901218" authorityName="Walker" authorityYear="1964" box="[312,498,833,857]" class="Reptilia" family="Megalosauridae" genus="Eustreptospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FE8384345F901218" box="[312,498,833,857]" italics="true" pageId="41" pageNumber="42">Eustreptospondylus</emphasis>
|
||
</taxonomicName>
|
||
: 55%;
|
||
<taxonomicName id="4C5B6ED2F15FE027FDFE84345C911218" baseAuthorityName="Zhao and Currie" baseAuthorityYear="1993" box="[581,755,833,857]" class="Reptilia" genus="Monolophosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FDFE84345C911218" box="[581,755,833,857]" italics="true" pageId="41" pageNumber="42">Monolophosaurus</emphasis>
|
||
</taxonomicName>
|
||
: 51%), whereas for six taxa (
|
||
<taxonomicName id="4C5B6ED2F15FE027FE6F84145C351239" authorityName="Benson" authorityYear="2008" box="[468,599,865,888]" class="Reptilia" family="Megalosauridae" genus="Duriavenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FE6F84145C351239" box="[468,599,865,888]" italics="true" pageId="41" pageNumber="42">Duriavenator</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FDCA84145C971239" authorityName="von Huene" authorityYear="1932" box="[625,757,865,888]" class="Reptilia" family="Megalosauridae" genus="Magnosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FDCA84145C971239" box="[625,757,865,888]" italics="true" pageId="41" pageNumber="42">Magnosaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FFCA84F55F7712D9" authorityName="Meyer" authorityYear="1830" box="[113,277,896,920]" class="Reptilia" genus="Streptospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FFCA84F55F7712D9" box="[113,277,896,920]" italics="true" pageId="41" pageNumber="42">Streptospondylus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FE8684F45C4A12D9" authorityName="Hendrickx & Mateus" authorityYear="2014" box="[317,552,897,920]" class="Reptilia" family="Megalosauridae" genus="Torvosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="species" species="guerneyi">
|
||
<emphasis id="B92FC943F15FE027FE8684F45C4A12D9" box="[317,552,897,920]" italics="true" pageId="41" pageNumber="42">Torvosaurus guerneyi</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FDF484F55F7812F6" authority=", Yungyangosaurus" authorityName="Yungyangosaurus" class="Reptilia" family="Piatnitzkysauridae" genus="Xuanhanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FDF484F55C9612D9" box="[591,756,896,920]" italics="true" pageId="41" pageNumber="42">Xuanhanosaurus</emphasis>
|
||
,
|
||
<emphasis id="B92FC943F15FE027FFCA84D55F7812F6" box="[113,282,928,951]" italics="true" pageId="41" pageNumber="42">Yungyangosaurus</emphasis>
|
||
</taxonomicName>
|
||
) only 10% or less of the characters could be coded. Within metriacanthosaurids, it was mainly the very fragmentary
|
||
<taxonomicName id="4C5B6ED2F15FE027FF4384AB5FD812B7" box="[248,442,990,1014]" class="Reptilia" family="Megalosauridae" genus="Metriacanthosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FF4384AB5FD812B7" box="[248,442,990,1014]" italics="true" pageId="41" pageNumber="42">Metriacanthosaurus</emphasis>
|
||
</taxonomicName>
|
||
(6.5% coded characters) that changed position (being found as sister taxon to
|
||
<taxonomicName id="4C5B6ED2F15FE027FDDF84885C801554" box="[612,738,1021,1045]" family="Metriacanthosauridae" genus="Shidaisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FDDF84885C801554" box="[612,738,1021,1045]" italics="true" pageId="41" pageNumber="42">Shidaisaurus</emphasis>
|
||
</taxonomicName>
|
||
at the base of the clade in weighing concavities below 10), although
|
||
<taxonomicName id="4C5B6ED2F15FE027FFCB83495E8F1515" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[112,237,1084,1108]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FFCB83495E8F1515" box="[112,237,1084,1108]" italics="true" pageId="41" pageNumber="42">Alpkarakush</emphasis>
|
||
</taxonomicName>
|
||
(25% coded characters) also changed position and was found to be sister taxon of
|
||
<taxonomicName id="4C5B6ED2F15FE027FE1383295C591532" box="[424,571,1116,1139]" family="Metriacanthosauridae" genus="Siamotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FE1383295C591532" box="[424,571,1116,1139]" italics="true" pageId="41" pageNumber="42">Siamotyrannus</emphasis>
|
||
</taxonomicName>
|
||
(6.8% coded characters) in these analyses. However, as the implied weighing analysis with k = 12 was closest to the equally weighted analysis, just with improved resolution, and
|
||
<bibRefCitation id="EFCA68A0F15FE027FE6C83CC5CDA1590" author="Goloboff PA & Torres A & Arias JS" box="[471,696,1209,1233]" pageId="41" pageNumber="42" pagination="407 - 37" refId="ref37377" refString="Goloboff PA, Torres A, Arias JS. Weighted parsimony outperforms other methods of phylogenetic inference under models appropriate for morphology. Cladistics 2018; 34: 407 - 37. https: // doi. org / 10.1111 / cla. 12205" type="journal article" year="2018">
|
||
Goloboff
|
||
<emphasis id="B92FC943F15FE027FD8183CF5C0B1590" box="[570,617,1209,1233]" italics="true" pageId="41" pageNumber="42">et al.</emphasis>
|
||
(2018)
|
||
</bibRefCitation>
|
||
found the weighted analysis to perform best at this concavity (see also
|
||
<bibRefCitation id="EFCA68A0F15FE027FFC9838D5E981451" author="Ezcurra MD" box="[114,250,1272,1296]" pageId="41" pageNumber="42" pagination="242 - 81" refId="ref36927" refString="Ezcurra MD. Exploring the effects of weighting against homoplasy in genealogies of palaeontological phylogenetic matrices. Cladistics 2024; 40: 242 - 81. https: // doi. org / 10.1111 / cla. 12581" type="journal article" year="2024">Ezcurra 2024</bibRefCitation>
|
||
), our following tree description and discussion is based on the strict consensus tree of the three MPTs resulting from this analysis. At least in respect to metriacanthosaurids, this tree was also the most consistent with the stratigraphic occurrences of the taxa.
|
||
</paragraph>
|
||
<paragraph id="8BE41551F15FE027FF3682E05FB417A7" blockId="41.[111,764,144,1954]" pageId="41" pageNumber="42">
|
||
The consensus tree from the implied weighing analysis with k = 12 was very well resolved, with just two polytomies in carnosaurs, one between
|
||
<taxonomicName id="4C5B6ED2F15FE027FE0882A15C6114AA" box="[435,515,1492,1515]" class="Reptilia" family="Spinosauridae" genus="Irritator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FE0882A15C6114AA" box="[435,515,1492,1515]" italics="true" pageId="41" pageNumber="42">Irritator</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FDA382A65CA714AA" box="[536,709,1491,1515]" family="Spinosauridae" genus="Vallibonavenatrix" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FDA382A65CA714AA" box="[536,709,1491,1515]" italics="true" pageId="41" pageNumber="42">Vallibonavenatrix</emphasis>
|
||
</taxonomicName>
|
||
, the
|
||
<taxonomicName id="4C5B6ED2F15FE027FFCA82865F69174A" baseAuthorityName="Charig & Milner" baseAuthorityYear="1986" box="[113,267,1523,1547]" class="Reptilia" family="Spinosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="subFamily" subFamily="Baryonychinae">Baryonychinae</taxonomicName>
|
||
, and a subclade containing
|
||
<taxonomicName id="4C5B6ED2F15FE027FDF482865CA7174B" box="[591,709,1523,1546]" class="Reptilia" family="Spinosauridae" genus="Spinosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FDF482865CA7174B" box="[591,709,1523,1546]" italics="true" pageId="41" pageNumber="42">Spinosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C5B6ED2F15FE027FFCA81675F63176B" authorityName="Allain, Xaisanavong, Richir & Khentavong" authorityYear="2012" box="[113,257,1554,1578]" family="Spinosauridae" genus="Ichthyovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FFCA81675F63176B" box="[113,257,1554,1578]" italics="true" pageId="41" pageNumber="42">Ichthyovenator</emphasis>
|
||
</taxonomicName>
|
||
, and one in metriacanthosaurids between
|
||
<taxonomicName id="4C5B6ED2F15FE027FFCA81475EAF1708" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[113,205,1586,1609]" class="Reptilia" family="Sinraptoridae" genus="Sinraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FFCA81475EAF1708" box="[113,205,1586,1609]" italics="true" pageId="41" pageNumber="42">Sinraptor</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FF6281475F0E1708" box="[217,364,1586,1609]" family="Metriacanthosauridae" genus="Siamotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FF6281475F0E1708" box="[217,364,1586,1609]" italics="true" pageId="41" pageNumber="42">Siamotyrannus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C5B6ED2F15FE027FE1881445C071708" box="[419,613,1585,1609]" class="Reptilia" family="Megalosauridae" genus="Metriacanthosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FE1881445C071708" box="[419,613,1585,1609]" italics="true" pageId="41" pageNumber="42">Metriacanthosaurus</emphasis>
|
||
</taxonomicName>
|
||
. Furthermore, therewasanadditionalthree-taxapolytomyoutsideCarnosauria, within megaraptorans. Apart from the better resolution, the results were almost identical to those found in the unweighted analysis, so this tree is used to discuss character distribution and possible evolutionary implications.
|
||
</paragraph>
|
||
<paragraph id="8BE41551F15FE027FF3681985B41139A" blockId="41.[111,764,144,1954]" lastBlockId="41.[808,1461,144,1829]" pageId="41" pageNumber="42">
|
||
Within Carnosauria, the implied weighting analysis found a basal dichotomy into Megalosauroidea and Allosauroidea. Within the former clade, a monophyletic
|
||
<taxonomicName id="4C5B6ED2F15FE027FFCA803E5F4F1622" box="[113,301,1867,1891]" family="Piatnitzkysauridae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Piatnitzkysauridae</taxonomicName>
|
||
formed the earliest branching subclade; this clade included the genera
|
||
<emphasis id="B92FC943F15FE027FE08801E5C3E16C3" box="[435,604,1899,1922]" italics="true" pageId="41" pageNumber="42">Yungyangosaurus</emphasis>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FDD0801E5C9716C2" authorityName="Madsen" authorityYear="1976" box="[619,757,1899,1923]" class="Reptilia" family="Piatnitzkysauridae" genus="Marshosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FDD0801E5C9716C2" box="[619,757,1899,1923]" italics="true" pageId="41" pageNumber="42">Marshosaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FFCA80FF5F7716E3" authorityName="Bonaparte" authorityYear="1979" box="[113,277,1930,1954]" class="Reptilia" family="Allosauridae" genus="Piatnitzkysaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FFCA80FF5F7716E3" box="[113,277,1930,1954]" italics="true" pageId="41" pageNumber="42">Piatnitzkysaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FE8A80FF5FB416E3" box="[305,470,1930,1954]" class="Reptilia" family="Piatnitzkysauridae" genus="Xuanhanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FE8A80FF5FB416E3" box="[305,470,1930,1954]" italics="true" pageId="41" pageNumber="42">Xuanhanosaurus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C5B6ED2F15FE027FD8B80FF5CD516E3" baseAuthorityName="Rauhut" baseAuthorityYear="2005" box="[560,695,1930,1954]" class="Reptilia" family="Piatnitzkysauridae" genus="Condorraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FD8B80FF5CD516E3" box="[560,695,1930,1954]" italics="true" pageId="41" pageNumber="42">Condorraptor</emphasis>
|
||
</taxonomicName>
|
||
. The other megalosauroids were placed in either the
|
||
<taxonomicName id="4C5B6ED2F15FE027FA9987E55BD111E9" authorityName="Stromer" authorityYear="1915" box="[1314,1459,144,168]" class="Reptilia" family="Spinosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Spinosauridae</taxonomicName>
|
||
or the
|
||
<taxonomicName id="4C5B6ED2F15FE027FCC087DA5A7D1186" box="[891,1055,175,199]" class="Reptilia" family="Megalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Megalosauridae</taxonomicName>
|
||
.
|
||
<taxonomicName id="4C5B6ED2F15FE027FB9587DA5AB21186" box="[1070,1232,175,199]" class="Reptilia" family="Megalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Megalosauridae</taxonomicName>
|
||
included the genera
|
||
<taxonomicName id="4C5B6ED2F15FE027FC9187BA5DBE11A6" authorityName="Mateus" authorityYear="1998" box="[810,988,207,231]" class="Reptilia" genus="Lourinhanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC9187BA5DBE11A6" box="[810,988,207,231]" italics="true" pageId="41" pageNumber="42">Lourinhanosaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FC4287BA5A1D11A6" baseAuthorityName="Benson" baseAuthorityYear="2010" box="[1017,1151,207,231]" class="Reptilia" family="Megalosauridae" genus="Megalosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC4287BA5A1D11A6" box="[1017,1151,207,231]" italics="true" pageId="41" pageNumber="42">Megalosaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FB2087BA5B6D11A6" box="[1179,1295,207,231]" class="Reptilia" family="Megalosauridae" genus="Afrovenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FB2087BA5B6D11A6" box="[1179,1295,207,231]" italics="true" pageId="41" pageNumber="42">Afrovenator</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FA9087BA5BCC11A7" authorityName="Benson" authorityYear="2008" box="[1323,1454,207,230]" class="Reptilia" family="Megalosauridae" genus="Duriavenator" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FA9087BA5BCC11A7" box="[1323,1454,207,230]" italics="true" pageId="41" pageNumber="42">Duriavenator</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FC91879B5DB31047" authorityName="Allain" authorityYear="2005" box="[810,977,238,262]" class="Reptilia" family="Megalosauridae" genus="Dubreuillosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC91879B5DB31047" box="[810,977,238,262]" italics="true" pageId="41" pageNumber="42">Dubreuillosaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FC5A879B5A101047" box="[993,1138,238,262]" family="Megalosaurinidae" genus="Wiehenvenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC5A879B5A101047" box="[993,1138,238,262]" italics="true" pageId="41" pageNumber="42">Wiehenvenator</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FB39879A5A981047" box="[1154,1274,239,262]" class="Reptilia" family="Megalosauridae" genus="Torvosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FB39879A5A981047" box="[1154,1274,239,262]" italics="true" pageId="41" pageNumber="42">Torvosaurus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FAB2879B5BCF1047" authorityName="Meyer" authorityYear="1830" box="[1289,1453,238,262]" class="Reptilia" genus="Streptospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FAB2879B5BCF1047" box="[1289,1453,238,262]" italics="true" pageId="41" pageNumber="42">Streptospondylus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C5B6ED2F15FE027FCE186785A761064" authorityName="Walker" authorityYear="1964" box="[858,1044,269,293]" class="Reptilia" family="Megalosauridae" genus="Eustreptospondylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FCE186785A761064" box="[858,1044,269,293]" italics="true" pageId="41" pageNumber="42">Eustreptospondylus</emphasis>
|
||
</taxonomicName>
|
||
, whereas
|
||
<taxonomicName id="4C5B6ED2F15FE027FB3A86785B4D1064" baseAuthorityName="Zhao and Currie" baseAuthorityYear="1993" box="[1153,1327,269,293]" class="Reptilia" genus="Monolophosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FB3A86785B4D1064" box="[1153,1327,269,293]" italics="true" pageId="41" pageNumber="42">Monolophosaurus</emphasis>
|
||
</taxonomicName>
|
||
and a clade including the poorly known
|
||
<taxonomicName id="4C5B6ED2F15FE027FBD186585A8C1005" authorityName="von Huene" authorityYear="1932" box="[1130,1262,301,324]" class="Reptilia" family="Megalosauridae" genus="Magnosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FBD186585A8C1005" box="[1130,1262,301,324]" italics="true" pageId="41" pageNumber="42">Magnosaurus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C5B6ED2F15FE027FA9586585BD11004" authorityName="Li, Peng, Ye, Jiang & Huang" authorityYear="2009" box="[1326,1459,301,325]" family="Sinraptoridae" genus="Leshansaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FA9586585BD11004" box="[1326,1459,301,325]" italics="true" pageId="41" pageNumber="42">Leshansaurus</emphasis>
|
||
</taxonomicName>
|
||
were found as an early branching
|
||
<taxonomicName id="4C5B6ED2F15FE027FB3386395B781025" authorityName="Stromer" authorityYear="1915" box="[1160,1306,332,356]" class="Reptilia" family="Spinosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Spinosauridae</taxonomicName>
|
||
, rather than in
|
||
<taxonomicName id="4C5B6ED2F15FE027FC97861E5DB210C2" box="[812,976,363,387]" class="Reptilia" family="Megalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Megalosauridae</taxonomicName>
|
||
. As in the unweighted analysis, Allosauroidea was subdivided into
|
||
<taxonomicName id="4C5B6ED2F15FE027FBA686FE5A9F10E2" authorityName="Paul" authorityYear="1988" box="[1053,1277,395,419]" class="Reptilia" family="Metriacanthosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Metriacanthosauridae</taxonomicName>
|
||
and Allosauria.
|
||
<taxonomicName id="4C5B6ED2F15FE027FC9186DF5DD51083" authorityName="Rauhut & Pol" authorityYear="2019" box="[810,951,426,450]" genus="Asfaltovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC9186DF5DD51083" box="[810,951,426,450]" italics="true" pageId="41" pageNumber="42">Asfaltovenator</emphasis>
|
||
</taxonomicName>
|
||
was here found as an allosaurid, sister taxon to
|
||
<taxonomicName id="4C5B6ED2F15FE027FC9186BC5DF310A0" baseAuthorityName="Madsen" baseAuthorityYear="1976" box="[810,913,457,481]" class="Reptilia" family="Allosauridae" genus="Allosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC9186BC5DF310A0" box="[810,913,457,481]" italics="true" pageId="41" pageNumber="42">Allosaurus</emphasis>
|
||
</taxonomicName>
|
||
, but otherwise the taxonomic composition and phylogenetic relationships within Allosauria correspond to those found in the unweighted analysis. Within metriacanthosaurids, the implied weighting analysis recovered the Chinese
|
||
<taxonomicName id="4C5B6ED2F15FE027FC9185325DCA131E" box="[810,936,583,607]" family="Metriacanthosauridae" genus="Shidaisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC9185325DCA131E" box="[810,936,583,607]" italics="true" pageId="41" pageNumber="42">Shidaisaurus</emphasis>
|
||
</taxonomicName>
|
||
as the earliest branching taxon, followed by the genus
|
||
<taxonomicName id="4C5B6ED2F15FE027FCD585135A40133F" box="[878,1058,614,638]" class="Reptilia" family="Ceratosauridae" genus="Yangchuanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FCD585135A40133F" box="[878,1058,614,638]" italics="true" pageId="41" pageNumber="42">Yangchuanosaurus</emphasis>
|
||
</taxonomicName>
|
||
with its two species.
|
||
<taxonomicName id="4C5B6ED2F15FE027FAB385135BE7133F" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[1288,1413,614,638]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FAB385135BE7133F" box="[1288,1413,614,638]" italics="true" pageId="41" pageNumber="42">Alpkarakush</emphasis>
|
||
</taxonomicName>
|
||
was confirmed as a metriacanthosaurid and represented the sister taxon to a clade uniting the remaining metriacanthosaurids
|
||
<taxonomicName id="4C5B6ED2F15FE027FC9185B15DE4139A" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[810,902,708,731]" class="Reptilia" family="Sinraptoridae" genus="Sinraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC9185B15DE4139A" box="[810,902,708,731]" italics="true" pageId="41" pageNumber="42">Sinraptor</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FC2A85B15A46139A" box="[913,1060,708,731]" family="Metriacanthosauridae" genus="Siamotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC2A85B15A46139A" box="[913,1060,708,731]" italics="true" pageId="41" pageNumber="42">Siamotyrannus</emphasis>
|
||
</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C5B6ED2F15FE027FBE185B15B7E139D" box="[1114,1308,708,732]" class="Reptilia" family="Megalosauridae" genus="Metriacanthosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FBE185B15B7E139D" box="[1114,1308,708,732]" italics="true" pageId="41" pageNumber="42">Metriacanthosaurus</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="8BE41551F15FE027FCFE85965BD615B0" blockId="41.[808,1461,144,1829]" pageId="41" pageNumber="42">
|
||
Characters that support the inclusion of
|
||
<taxonomicName id="4C5B6ED2F15FE027FB4A85965B0C13BA" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[1265,1390,739,763]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FB4A85965B0C13BA" box="[1265,1390,739,763]" italics="true" pageId="41" pageNumber="42">Alpkarakush</emphasis>
|
||
</taxonomicName>
|
||
in the
|
||
<taxonomicName id="4C5B6ED2F15FE027FC9184765A68125A" authorityName="Paul" authorityYear="1988" box="[810,1034,771,795]" class="Reptilia" family="Metriacanthosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Metriacanthosauridae</taxonomicName>
|
||
include the following: posterior margin of the infratemporal fenestra straight or only slightly convex (judged from the morphology of the quadratojugal and the facet for the contact with the squamosal on this bone; unknown in
|
||
<taxonomicName id="4C5B6ED2F15FE027FC9184F55DCA12D9" box="[810,936,896,920]" family="Metriacanthosauridae" genus="Shidaisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC9184F55DCA12D9" box="[810,936,896,920]" italics="true" pageId="41" pageNumber="42">Shidaisaurus</emphasis>
|
||
</taxonomicName>
|
||
); mesial carina of lateral teeth extended to base of crown (unknown in
|
||
<taxonomicName id="4C5B6ED2F15FE027FBB184EA5AEA12F6" box="[1034,1160,927,951]" family="Metriacanthosauridae" genus="Shidaisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FBB184EA5AEA12F6" box="[1034,1160,927,951]" italics="true" pageId="41" pageNumber="42">Shidaisaurus</emphasis>
|
||
</taxonomicName>
|
||
); flexor tubercle in manual unguals reduced and less than one-third of the height of the articular facet (apart from
|
||
<taxonomicName id="4C5B6ED2F15FE027FB9484AB5ACE12B7" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[1071,1196,990,1014]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FB9484AB5ACE12B7" box="[1071,1196,990,1014]" italics="true" pageId="41" pageNumber="42">Alpkarakush</emphasis>
|
||
</taxonomicName>
|
||
only known in
|
||
<taxonomicName id="4C5B6ED2F15FE027FAEC84AB5BD112B4" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[1367,1459,990,1013]" class="Reptilia" family="Sinraptoridae" genus="Sinraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FAEC84AB5BD112B4" box="[1367,1459,990,1013]" italics="true" pageId="41" pageNumber="42">Sinraptor</emphasis>
|
||
</taxonomicName>
|
||
within metriacanthosaurids; also present in
|
||
<taxonomicName id="4C5B6ED2F15FE027FAA884885BC21554" authorityName="Rauhut & Pol" authorityYear="2019" box="[1299,1440,1021,1045]" genus="Asfaltovenator" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FAA884885BC21554" box="[1299,1440,1021,1045]" italics="true" pageId="41" pageNumber="42">Asfaltovenator</emphasis>
|
||
</taxonomicName>
|
||
); medial brevis shelf poorly developed and not visible in lateral view; angle between proximal part of ischial shaft and pubic peduncle of ischium less than 110°; presence of a posterior midline crest on the articulated ischia; presence of a strongly expanded ischial boot; presence of an oval fossa on the base of the ascending process of the astragalus; pedal ungual II asymmetrical (shared with
|
||
<taxonomicName id="4C5B6ED2F15FE027FC1483AC5A6915B1" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[943,1035,1241,1264]" class="Reptilia" family="Sinraptoridae" genus="Sinraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC1483AC5A6915B1" box="[943,1035,1241,1264]" italics="true" pageId="41" pageNumber="42">Sinraptor</emphasis>
|
||
</taxonomicName>
|
||
, unknown in other metriacanthosaurids).
|
||
</paragraph>
|
||
<paragraph id="8BE41551F15FE027FCFE838D5B721664" blockId="41.[808,1461,144,1829]" pageId="41" pageNumber="42">
|
||
Within
|
||
<taxonomicName id="4C5B6ED2F15FE027FC1E838D5AE51451" authorityName="Paul" authorityYear="1988" box="[933,1159,1272,1296]" class="Reptilia" family="Metriacanthosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="family">Metriacanthosauridae</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15FE027FB1A838D5B7C1451" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[1185,1310,1272,1296]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FB1A838D5B7C1451" box="[1185,1310,1272,1296]" italics="true" pageId="41" pageNumber="42">Alpkarakush</emphasis>
|
||
</taxonomicName>
|
||
furthermore showsthefollowingsynapomorphieswithothertaxaorsubclades of this clade, which were absent in other members: ventral prong of the quadratojugal ends anterior to or level with the posterior rim of the infratemporal fenestra (shared with
|
||
<taxonomicName id="4C5B6ED2F15FE027FA9D82035BE014CC" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[1318,1410,1398,1421]" class="Reptilia" family="Sinraptoridae" genus="Sinraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FA9D82035BE014CC" box="[1318,1410,1398,1421]" italics="true" pageId="41" pageNumber="42">Sinraptor</emphasis>
|
||
</taxonomicName>
|
||
and unknown in
|
||
<taxonomicName id="4C5B6ED2F15FE027FC0282E05A2E14ED" box="[953,1100,1429,1452]" family="Metriacanthosauridae" genus="Siamotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC0282E05A2E14ED" box="[953,1100,1429,1452]" italics="true" pageId="41" pageNumber="42">Siamotyrannus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C5B6ED2F15FE027FB3082E05B2F14EC" box="[1163,1357,1429,1453]" class="Reptilia" family="Megalosauridae" genus="Metriacanthosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FB3082E05B2F14EC" box="[1163,1357,1429,1453]" italics="true" pageId="41" pageNumber="42">Metriacanthosaurus</emphasis>
|
||
</taxonomicName>
|
||
, whereas the prong extended further posteriorly in
|
||
<taxonomicName id="4C5B6ED2F15FE027FB4C82C15BC9148D" box="[1271,1451,1460,1484]" class="Reptilia" family="Ceratosauridae" genus="Yangchuanosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FB4C82C15BC9148D" box="[1271,1451,1460,1484]" italics="true" pageId="41" pageNumber="42">Yangchuanosaurus</emphasis>
|
||
</taxonomicName>
|
||
; although the jugal was not preserved, the length of its ventral posterior prong can be estimated from the articular facet on the quadratojugal); posterior process of the postorbital broader mediolaterally than high dorsoventrally (shared with
|
||
<taxonomicName id="4C5B6ED2F15FE027FAEC81475BD11708" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[1367,1459,1586,1609]" class="Reptilia" family="Sinraptoridae" genus="Sinraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FAEC81475BD11708" box="[1367,1459,1586,1609]" italics="true" pageId="41" pageNumber="42">Sinraptor</emphasis>
|
||
</taxonomicName>
|
||
and unknown in
|
||
<taxonomicName id="4C5B6ED2F15FE027FC5C81245A181729" box="[999,1146,1617,1640]" family="Metriacanthosauridae" genus="Siamotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC5C81245A181729" box="[999,1146,1617,1640]" italics="true" pageId="41" pageNumber="42">Siamotyrannus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C5B6ED2F15FE027FB0D81245B1A1728" box="[1206,1400,1617,1641]" class="Reptilia" family="Megalosauridae" genus="Metriacanthosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FB0D81245B1A1728" box="[1206,1400,1617,1641]" italics="true" pageId="41" pageNumber="42">Metriacanthosaurus</emphasis>
|
||
</taxonomicName>
|
||
; also present in derived carcharodontosaurids); transverse dimensions of mid-sacral vertebrae reduced in relation to first and last sacral (shared with all metriacanthosaurids more derived than
|
||
<taxonomicName id="4C5B6ED2F15FE027FC9181BB5DCA17A7" box="[810,936,1742,1766]" family="Metriacanthosauridae" genus="Shidaisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC9181BB5DCA17A7" box="[810,936,1742,1766]" italics="true" pageId="41" pageNumber="42">Shidaisaurus</emphasis>
|
||
</taxonomicName>
|
||
); lateral groove present on pubic boot (present in
|
||
<taxonomicName id="4C5B6ED2F15FE027FCFD819B5DC01644" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[838,930,1774,1797]" class="Reptilia" family="Sinraptoridae" genus="Sinraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FCFD819B5DC01644" box="[838,930,1774,1797]" italics="true" pageId="41" pageNumber="42">Sinraptor</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C5B6ED2F15FE027FC63819B5A091644" box="[984,1131,1774,1797]" family="Metriacanthosauridae" genus="Siamotyrannus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC63819B5A091644" box="[984,1131,1774,1797]" italics="true" pageId="41" pageNumber="42">Siamotyrannus</emphasis>
|
||
</taxonomicName>
|
||
); fibular crest on tibia bulbous (shared with
|
||
<taxonomicName id="4C5B6ED2F15FE027FC0B80785A6E1665" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[944,1036,1805,1828]" class="Reptilia" family="Sinraptoridae" genus="Sinraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FC0B80785A6E1665" box="[944,1036,1805,1828]" italics="true" pageId="41" pageNumber="42">Sinraptor</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C5B6ED2F15FE027FB8680785A9D1664" box="[1085,1279,1805,1829]" class="Reptilia" family="Megalosauridae" genus="Metriacanthosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="41" pageNumber="42" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15FE027FB8680785A9D1664" box="[1085,1279,1805,1829]" italics="true" pageId="41" pageNumber="42">Metriacanthosaurus</emphasis>
|
||
</taxonomicName>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BE41551F15FE027FC4E80315A8A161F" blockId="41.[1013,1256,1860,1886]" box="[1013,1256,1860,1886]" pageId="41" pageNumber="42">
|
||
<emphasis id="B92FC943F15FE027FC4E80315A8A161F" bold="true" box="[1013,1256,1860,1886]" pageId="41" pageNumber="42">Biogeographic analysis</emphasis>
|
||
</paragraph>
|
||
<paragraph id="8BE41551F15FE025FC91801E5E901005" blockId="41.[810,1459,1899,1954]" lastBlockId="43.[112,764,144,1045]" lastPageId="43" lastPageNumber="44" pageId="41" pageNumber="42">
|
||
As outlined in the Material and methods, the biogeographic analysis was carried out on the basis of the trees resulting from the phylogenetic analysis using implied weights with a k = 12. Since our focus lies on the phylogenetic position and biogeographic implications for
|
||
<taxonomicName id="4C5B6ED2F15DE025FEA187BA5FF411A6" baseAuthorityName="Currie and Zhao" baseAuthorityYear="1993" box="[282,406,207,231]" class="Reptilia" family="Metriacanthosauridae" genus="Alpkarakush" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="43" pageNumber="44" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15DE025FEA187BA5FF411A6" box="[282,406,207,231]" italics="true" pageId="43" pageNumber="44">Alpkarakush</emphasis>
|
||
</taxonomicName>
|
||
, we restrict the presentation of the biogeographic results to the Carnosauria. For a full documentation of the results of the S-DIVA analysis see the Supporting information.
|
||
</paragraph>
|
||
<caption id="DF2445D9F15CE024FF3A80595D4416D6" ID-DOI="http://doi.org/10.5281/zenodo.13774297" ID-Zenodo-Dep="13774297" httpUri="https://zenodo.org/record/13774297/files/figure.png" pageId="42" pageNumber="43" startId="42.[129,194,1836,1860]" targetBox="[174,1428,146,1806]" targetPageId="42" targetType="figure">
|
||
<paragraph id="8BE41551F15CE024FF3A80595D4416D6" blockId="42.[129,1470,1836,1944]" pageId="42" pageNumber="43">
|
||
<emphasis id="B92FC943F15CE024FF3A80595E871605" bold="true" box="[129,229,1836,1860]" pageId="42" pageNumber="43">Figure 27.</emphasis>
|
||
Biogeographic reconstructions for selected clades, resulting from the S-DIVA analysis. Legend: A, north-eastern Europe; B, southern South America; C, North America; D, northern Africa; E, north-eastern Asia; F, south-western Europe; G, Central Asia; H, Australia; I, Antarctica; J, South-East Asia; L, Central South America. For further details on these regions see the Material and methods section. For full results of the S-DIVA analysis see Fig. S7 in the Supporting information.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BE41551F15DE025FF3686395F7C1554" blockId="43.[112,764,144,1045]" pageId="43" pageNumber="44">
|
||
The S-DIVA analysis (
|
||
<figureCitation id="136009D4F15DE025FECD86395FDF1025" box="[374,445,332,356]" captionStart="Figure 27" captionStartId="42.[129,194,1836,1860]" captionTargetBox="[174,1428,146,1806]" captionTargetId="figure-7@42.[172,1430,144,1808]" captionTargetPageId="42" captionText="Figure 27. Biogeographic reconstructions for selected clades, resulting from the S-DIVA analysis. Legend:A, north-eastern Europe; B, southern South America; C, North America; D, northern Africa; E, north-eastern Asia; F, south-western Europe; G, Central Asia; H, Australia; I, Antarctica; J, South-East Asia; L, Central South America.For further details on these regions see the Material and methods section. For full results of the S-DIVA analysis see Fig. S7 in the Supporting information." figureDoi="http://doi.org/10.5281/zenodo.13774297" httpUri="https://zenodo.org/record/13774297/files/figure.png" pageId="43" pageNumber="44">Fig. 27</figureCitation>
|
||
) identified South-East Asia as the ancestral territory of Tetanurae, and eastern Asia in general (50% North-East Asia, 50% South-East Asia) as the original area for
|
||
<taxonomicName id="4C5B6ED2F15DE025FF6386DF5F0B1083" box="[216,361,426,450]" class="Reptilia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Avetheropoda" pageId="43" pageNumber="44" phylum="Chordata" rank="order">Avetheropoda</taxonomicName>
|
||
(the clade including Coelurosauria and Carnosauria). Likewise, a South-East Asian origin for Carnosauria was also indicated. Within Carnosauria, the area of origin of megalosauroids was problematic, as all three included clades (
|
||
<taxonomicName id="4C5B6ED2F15DE025FF0585525F1D137E" box="[190,383,551,575]" family="Piatnitzkysauridae" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="43" pageNumber="44" phylum="Chordata" rank="family">Piatnitzkysauridae</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C5B6ED2F15DE025FE3385525C78137E" authorityName="Stromer" authorityYear="1915" box="[392,538,551,575]" class="Reptilia" family="Spinosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="43" pageNumber="44" phylum="Chordata" rank="family">Spinosauridae</taxonomicName>
|
||
, and
|
||
<taxonomicName id="4C5B6ED2F15DE025FDEB85525C97137E" box="[592,757,551,575]" class="Reptilia" family="Megalosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="43" pageNumber="44" phylum="Chordata" rank="family">Megalosauridae</taxonomicName>
|
||
) have wide distributions, with megalosaurids being especially diverse in the Middle and Late Jurassic of Europe (see also Rauhut
|
||
<emphasis id="B92FC943F15DE025FF7185F35E9B13DC" box="[202,249,645,669]" italics="true" pageId="43" pageNumber="44">et al</emphasis>
|
||
. 2016, 2018, 2020), and consequently, Europe was identified as area of origin of this clade. However, given that the South-East Asian
|
||
<taxonomicName id="4C5B6ED2F15DE025FE2F85B05C53139D" box="[404,561,709,732]" genus="Yunyangosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="43" pageNumber="44" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15DE025FE2F85B05C53139D" box="[404,561,709,732]" italics="true" pageId="43" pageNumber="44">Yunyangosaurus</emphasis>
|
||
</taxonomicName>
|
||
was found as the earliest branching piatnitzkysaurid, and the central Asian
|
||
<taxonomicName id="4C5B6ED2F15DE025FFCA84765F7D125A" baseAuthorityName="Zhao and Currie" baseAuthorityYear="1993" box="[113,287,771,795]" class="Reptilia" genus="Monolophosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="43" pageNumber="44" phylum="Chordata" rank="genus">
|
||
<emphasis id="B92FC943F15DE025FFCA84765F7D125A" box="[113,287,771,795]" italics="true" pageId="43" pageNumber="44">Monolophosaurus</emphasis>
|
||
</taxonomicName>
|
||
as the earliest branching spinosaurid, an Asian origin of the Megalosauroidea in general, followed by a rapid dispersal over much of Pangaea, seems likely. The area of origin of Allosauroidea was identified as 50% South-East Asia and 50% south-west Europe; given that very little is known about Central Asian theropods, an origin somewhere between eastern Asia and Europe seems likely. For metriacanthosaurids, the area of origin was clearly identified as South-East Asia, followed by a dispersal into central Asia.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |