373 lines
55 KiB
XML
373 lines
55 KiB
XML
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<mods:title id="A45740688557B47B00CCFF47AD091312">Pareisactus evrostos, a new basal iguanodontian (Dinosauria: Ornithopoda) from the Upper Cretaceous of southwestern Europe</mods:title>
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<mods:namePart id="9CE8C52A2DEDAE8FEB1EE69739B7404B">Párraga, Javier</mods:namePart>
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<mods:namePart id="5FEEBDE7C951E67C9C3C17B194D2BDB6">Prieto-Márquez, Albert</mods:namePart>
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<mods:title id="1D4161AA291198FF6539560777537762">Zootaxa</mods:title>
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<mods:date id="08431A548DAABC9AF6D8C3A8C5624732">2019</mods:date>
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<treatment id="03C758683815FFEFFF23FDE59290AAD0" ID-DOI="http://doi.org/10.5281/zenodo.5936048" ID-GBIF-Taxon="154533654" ID-Zenodo-Dep="5936048" LSID="urn:lsid:plazi:treatment:03C758683815FFEFFF23FDE59290AAD0" httpUri="http://treatment.plazi.org/id/03C758683815FFEFFF23FDE59290AAD0" lastPageId="4" lastPageNumber="251" pageId="2" pageNumber="249">
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<subSubSection id="C374BAF53815FFE9FF23FDE592B4AFFE" box="[151,497,631,658]" pageId="2" pageNumber="249" type="nomenclature">
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<paragraph id="8BD1E97E3815FFE9FF23FDE592B4AFFE" blockId="2.[151,497,631,691]" box="[151,497,631,658]" pageId="2" pageNumber="249">
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<heading id="D0995E123815FFE9FF23FDE592B4AFFE" bold="true" box="[151,497,631,658]" fontSize="11" level="1" pageId="2" pageNumber="249" reason="1">
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<emphasis id="B91A356C3815FFE9FF23FDE592B4AFFE" bold="true" box="[151,497,631,658]" pageId="2" pageNumber="249">
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<taxonomicName id="4C6E92FD3815FFE9FF23FDE592C9AFFD" authority="Párraga & Prieto-Márquez, 2019" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[151,396,631,657]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="249" phylum="Chordata" rank="species" species="evrostos" status="sp. nov.">
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<emphasis id="B91A356C3815FFE9FF23FDE592C9AFFD" bold="true" box="[151,396,631,657]" italics="true" pageId="2" pageNumber="249">Pareisactus evrostos</emphasis>
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</taxonomicName>
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<taxonomicNameLabel id="A22988173815FFE9FE27FDEA92B4AFFE" box="[403,497,632,658]" pageId="2" pageNumber="249" rank="species">sp. nov.</taxonomicNameLabel>
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</emphasis>
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="C374BAF53815FFE9FF23FD08927FAFDF" box="[151,314,666,691]" pageId="2" pageNumber="249" type="description">
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<paragraph id="8BD1E97E3815FFE9FF23FD08927FAFDF" blockId="2.[151,497,631,691]" box="[151,314,666,691]" pageId="2" pageNumber="249">
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(
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<figureCitation id="1355F5FB3815FFE9FF2BFD0893A9AFDF" box="[159,236,666,691]" captionStart="FIGURE 2" captionStartId="3.[151,250,1868,1890]" captionTargetBox="[177,1410,254,1733]" captionTargetId="figure@3.[151,1436,223,1773]" captionTargetPageId="3" captionText="FIGURE 2. Holotype specimen of Pareisactus evrostos, a nearly complete left scapula (MCD 5371) in (A) dorsal, (B) lateral, (C) medial, and (D) ventral views." httpUri="https://zenodo.org/record/2624341/files/figure.png" pageId="2" pageNumber="249">Figs. 2</figureCitation>
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and
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<figureCitation id="1355F5FB3815FFE9FE97FD099274AFDF" box="[291,305,667,691]" captionStart="FIGURE 3" captionStartId="5.[151,250,1703,1725]" captionTargetBox="[151,1436,195,1651]" captionTargetId="figure@5.[151,1436,195,1651]" captionTargetPageId="5" captionText="FIGURE 3. Strict consensus tree derived from the parsimony analysis showing the position of Pareisactus evrostos among iguanodontian ornithopods. Depicted specimens in left lateral view are: Bolong yixianensis, YHZ-001 (line drawing based on Wu 2010, plate III); Camptosaurus dispar, USNM 5955; Corythosaurus casuarius, ROM 845; Cumnoria prestwichii, OXFUM J.3303; Eolambia caroljonesa, CEUM 52097 (reversed, line drawing based on McDonald et al. 2012, fig. 27C); Gilmoreosaurus mongoliensis, AMNH FARB 30725; Haya griva, IGM 100-2015; Iguanodon bernissartensis, IRSNB 1534; Jinzhousaurus yangi, IVPP V12691 (line drawing based on Wang et al. 2011, fig. 4); Lophorhothon atopus, AUMP 2295; Orodromeus makelai, MOR 294 (reversed); Oryctodromeus cubicularis, MOR 1636; Ouranosaurus nigeriensis, cast of MNHN GDF 300; Pareisactus evrostos, MCD 5371; Rhabdodon priscus, MDE-C3 1013; Tenontosaurus tilletti, MOR 2571; Uteodon aphanoecetes, CM 11337; Zalmoxes robustus, line drawing from Weishampel et al. [2003, fig. 19C]); Z. shqiperorum, UBB NVZ1-24 (reversed, line drawing based on Godefroit et al. 2009, fig. 15A)." httpUri="https://zenodo.org/record/2624343/files/figure.png" pageId="2" pageNumber="249">3</figureCitation>
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)
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</paragraph>
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</subSubSection>
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<subSubSection id="C374BAF53815FFE9FF23FD7091B0AF97" box="[151,757,738,763]" pageId="2" pageNumber="249" type="materials_examined">
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<paragraph id="8BD1E97E3815FFE9FF23FD7091B0AF97" blockId="2.[151,1436,738,1447]" box="[151,757,738,763]" pageId="2" pageNumber="249">
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<materialsCitation id="3B06E3233815FFE9FF23FD7091B0AF97" ID-GBIF-Occurrence="2234220157" box="[151,757,738,763]" collectionCode="MCD" pageId="2" pageNumber="249" specimenCode="MCD 5371" specimenCount="1" typeStatus="holotype">
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<emphasis id="B91A356C3815FFE9FF23FD709241AF97" bold="true" box="[151,260,738,763]" pageId="2" pageNumber="249">
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<typeStatus id="54D557DC3815FFE9FF23FD709241AF97" box="[151,260,738,763]" pageId="2" pageNumber="249" type="holotype">Holotype</typeStatus>
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</emphasis>
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:
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<specimenCode id="DBC841053815FFE9FEA7FD7192D1AF97" box="[275,404,738,763]" collectionCode="MCD" country="0" httpUri="http://grbio.org/cool/ar0g-m2yd" name="Muzeul Civilizatiei Dacice si Romane Deva" pageId="2" pageNumber="249">MCD 5371</specimenCode>
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, a nearly complete left scapula.
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</materialsCitation>
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</paragraph>
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</subSubSection>
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<subSubSection id="C374BAF53815FFE9FF73FC94908CAE2F" pageId="2" pageNumber="249" type="etymology">
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<paragraph id="8BD1E97E3815FFE9FF73FC94908CAE2F" blockId="2.[151,1436,738,1447]" pageId="2" pageNumber="249">
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<emphasis id="B91A356C3815FFE9FF73FC94920DAE73" bold="true" box="[199,328,774,799]" pageId="2" pageNumber="249">Etymology</emphasis>
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:
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<emphasis id="B91A356C3815FFE9FEECFC94928DAE73" box="[344,456,774,799]" italics="true" pageId="2" pageNumber="249">Eύρωστoς</emphasis>
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(
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<taxonomicName id="4C6E92FD3815FFE9FE6EFC959173AE72" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[474,566,775,798]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="249" phylum="Chordata" rank="species" species="evrostos">
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<emphasis id="B91A356C3815FFE9FE6EFC959173AE72" box="[474,566,775,798]" italics="true" pageId="2" pageNumber="249">evrostos</emphasis>
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</taxonomicName>
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) means ‘robust’ in Greek, in reference to the great mediolateral thickness of the proximal extent of the scapula and massive build of the deltoid ridge.
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</paragraph>
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</subSubSection>
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<subSubSection id="C374BAF53815FFE9FF73FCDD900CAEC3" pageId="2" pageNumber="249" type="materials_examined">
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<paragraph id="8BD1E97E3815FFE9FF73FCDD96C2AE0B" blockId="2.[151,1436,738,1447]" box="[199,1415,846,871]" pageId="2" pageNumber="249">
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<emphasis id="B91A356C3815FFE9FF73FCDD9213AE0B" box="[199,342,846,871]" italics="true" pageId="2" pageNumber="249">Type locality</emphasis>
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: Basturs Poble, near the village of the same name, Lleida Province, northeastern
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<collectingCountry id="F379A9EE3815FFE9FB52FCDC9663AE0B" box="[1254,1318,846,871]" name="Spain" pageId="2" pageNumber="249">Spain</collectingCountry>
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(
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<figureCitation id="1355F5FB3815FFE9FA81FCDC9633AE0B" box="[1333,1398,846,871]" captionStart="FIGURE 1" captionStartId="1.[151,250,1392,1414]" captionTargetBox="[151,1436,666,1371]" captionTargetId="figure@1.[151,1436,660,1371]" captionTargetPageId="1" captionText="FIGURE 1. Geographic location and stratigraphic position of the Basturs Poble bonebed, the type locality of Pareisactus evrostos." httpUri="https://zenodo.org/record/2624339/files/figure.png" pageId="2" pageNumber="249">Fig. 1</figureCitation>
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).
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</paragraph>
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<paragraph id="8BD1E97E3815FFE9FF73FCE6900CAEC3" blockId="2.[151,1436,738,1447]" pageId="2" pageNumber="249">
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<emphasis id="B91A356C3815FFE9FF73FCE69219AEE7" box="[199,348,882,907]" italics="true" pageId="2" pageNumber="249">Type horizon</emphasis>
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: Upper lower Maastrichtian strata of the Conques Formation, Tremp Group, eastern Tremp Syncline, northeastern
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<collectingCountry id="F379A9EE3815FFE9FE2CFC04929DAEC3" box="[408,472,918,943]" name="Spain" pageId="2" pageNumber="249">Spain</collectingCountry>
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(
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<bibRefCitation id="EFFF948F3815FFE9FE53FC0491A5AEC3" author="Fondevilla, V. & Dinares-Turell, J. & Oms, O." box="[487,736,918,943]" pageId="2" pageNumber="249" pagination="55 - 68" refId="ref5538" refString="Fondevilla, V., Dinares-Turell, J. & Oms, O. (2016) The chronostratigraphic framework of the South-Pyrenean Maastrichtian succession reappraised: implications for basin development and end-Cretaceous dinosaur faunal turnover. Sediment Geology, 337, 55 - 68. https: // doi. org / 10.1016 / j. sedgeo. 2016.03.006" type="journal article" year="2016">
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Fondevilla
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<emphasis id="B91A356C3815FFE9FDD3FC0591DDAEC3" box="[615,664,918,943]" italics="true" pageId="2" pageNumber="249">et al</emphasis>
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. 2016
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</bibRefCitation>
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) (
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<figureCitation id="1355F5FB3815FFE9FD43FC04907DAEC3" box="[759,824,918,943]" captionStart="FIGURE 1" captionStartId="1.[151,250,1392,1414]" captionTargetBox="[151,1436,666,1371]" captionTargetId="figure@1.[151,1436,660,1371]" captionTargetPageId="1" captionText="FIGURE 1. Geographic location and stratigraphic position of the Basturs Poble bonebed, the type locality of Pareisactus evrostos." httpUri="https://zenodo.org/record/2624339/files/figure.png" pageId="2" pageNumber="249">Fig. 1</figureCitation>
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).
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</paragraph>
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</subSubSection>
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<subSubSection id="C374BAF53815FFE9FF73FC289670A9C7" pageId="2" pageNumber="249" type="diagnosis">
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<paragraph id="8BD1E97E3815FFE9FF73FC289670A9C7" blockId="2.[151,1436,738,1447]" pageId="2" pageNumber="249">
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<emphasis id="B91A356C3815FFE9FF73FC28927FAEBF" bold="true" box="[199,314,954,979]" pageId="2" pageNumber="249">Diagnosis</emphasis>
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. Basal iguanodontian ornithopod possessing the following autapomorphies: scapula with dorsoventral extent of proximal constriction (scapular ‘neck’) as deep as it is thick; lateral surface of proximal scapula under acromion process forming deep and thick roof over extremely deep deltoid fossa; long glenoid apex accounting for half of maximum depth of proximal region of scapula.
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<taxonomicName id="4C6E92FD3815FFE9FBB9FBB597B9A952" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[1037,1276,1063,1086]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="249" phylum="Chordata" rank="species" species="evrostos">
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<emphasis id="B91A356C3815FFE9FBB9FBB597B9A952" box="[1037,1276,1063,1086]" italics="true" pageId="2" pageNumber="249">Pareisactus evrostos</emphasis>
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</taxonomicName>
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differs from dryomorphan ornithopods in combining realtively long craniodorsally projected acromion process, craniocaudally short scapular ‘neck’, and glenoid apex that is strongly offset caudally from level of craniodorsal tip of the acromion process. It differs from other pre-dryomorphan ornithopods in the autapomorphies listed above.
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</paragraph>
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</subSubSection>
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<subSubSection id="C374BAF53815FFEFFF73FB249290AAD0" lastPageId="4" lastPageNumber="251" pageId="2" pageNumber="249" type="description">
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||
<paragraph id="8BD1E97E3815FFE9FF73FB2492BCA8CB" blockId="2.[151,1436,738,1447]" pageId="2" pageNumber="249">
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<emphasis id="B91A356C3815FFE9FF73FB249214A9A3" bold="true" box="[199,337,1206,1231]" pageId="2" pageNumber="249">Description</emphasis>
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. MCD 5371 is a nearly complete left scapula, missing part of the coracoid facet, the cranial margin of the glenoid, and a substantial part of the caudodorsal extent and caudoventral margin of the distal region of the scapular blade (
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<figureCitation id="1355F5FB3815FFE9FE3AFB6C9291A87B" box="[398,468,1278,1303]" captionStart="FIGURE 2" captionStartId="3.[151,250,1868,1890]" captionTargetBox="[177,1410,254,1733]" captionTargetId="figure@3.[151,1436,223,1773]" captionTargetPageId="3" captionText="FIGURE 2. Holotype specimen of Pareisactus evrostos, a nearly complete left scapula (MCD 5371) in (A) dorsal, (B) lateral, (C) medial, and (D) ventral views." httpUri="https://zenodo.org/record/2624341/files/figure.png" pageId="2" pageNumber="249">Fig. 2</figureCitation>
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; Table 1). At about two-fifths of the scapular length, there is a fracture that crosses the bone dorsoventrally and that has exaggerated the longitudinal medial curvature of the scapula. The lateral surface of the scapula is strongly convex dorsoventrally, particularly along its thickest proximal extent (
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<figureCitation id="1355F5FB3815FFE9FB76FAD4965FA833" box="[1218,1306,1350,1375]" captionStart="FIGURE 2" captionStartId="3.[151,250,1868,1890]" captionTargetBox="[177,1410,254,1733]" captionTargetId="figure@3.[151,1436,223,1773]" captionTargetPageId="3" captionText="FIGURE 2. Holotype specimen of Pareisactus evrostos, a nearly complete left scapula (MCD 5371) in (A) dorsal, (B) lateral, (C) medial, and (D) ventral views." httpUri="https://zenodo.org/record/2624341/files/figure.png" pageId="2" pageNumber="249">Fig. 2B</figureCitation>
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), becoming flatter distally along the scapular blade. In contrast, the medial surface of the scapula is flat throughout the entire length of the element (
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<figureCitation id="1355F5FB3815FFE9FE27FA1C92A9A8CB" box="[403,492,1422,1447]" captionStart="FIGURE 2" captionStartId="3.[151,250,1868,1890]" captionTargetBox="[177,1410,254,1733]" captionTargetId="figure@3.[151,1436,223,1773]" captionTargetPageId="3" captionText="FIGURE 2. Holotype specimen of Pareisactus evrostos, a nearly complete left scapula (MCD 5371) in (A) dorsal, (B) lateral, (C) medial, and (D) ventral views." httpUri="https://zenodo.org/record/2624341/files/figure.png" pageId="2" pageNumber="249">Fig. 2C</figureCitation>
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).
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</paragraph>
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<paragraph id="8BD1E97E3815FFE9FF2BF996971EABB2" pageId="2" pageNumber="249">
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<table id="F96E1BDE38150014FF2BF9969789ABB2" box="[159,1228,1540,1758]" gridcols="2" gridrows="5" pageId="2" pageNumber="249">
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<tr id="355EEB3C38150014FF2BF9969789AB76" box="[159,1228,1540,1562]" gridrow="0" pageId="2" pageNumber="249">
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<th id="768F824038150014FF2BF996976EAB76" box="[159,1067,1540,1562]" gridcol="0" gridrow="0" pageId="2" pageNumber="249">Scapular dimension</th>
|
||
<th id="768F824038150014FBF6F9969789AB76" box="[1090,1228,1540,1562]" gridcol="1" gridrow="0" pageId="2" pageNumber="249">Measurement</th>
|
||
</tr>
|
||
<tr id="355EEB3C38150014FF2BF9BF9789AB0E" box="[159,1228,1581,1634]" gridrow="1" pageId="2" pageNumber="249">
|
||
<td id="768F824038150014FF2BF9BF976EAB0E" box="[159,1067,1581,1634]" gridcol="0" gridrow="1" pageId="2" pageNumber="249">Length from the craniodorsal margin of the proximal end to the preserved distal-most end of the scapular blade</td>
|
||
<td id="768F824038150014FBF6F9BF9789AB0E" box="[1090,1228,1581,1634]" gridcol="1" gridrow="1" pageId="2" pageNumber="249">307</td>
|
||
</tr>
|
||
<tr id="355EEB3C38150014FF2BF9E79789ABE7" box="[159,1228,1653,1675]" gridrow="2" pageId="2" pageNumber="249">
|
||
<td id="768F824038150014FF2BF9E7976EABE7" box="[159,1067,1653,1675]" gridcol="0" gridrow="2" pageId="2" pageNumber="249">Distance from the dorsal tip of the acromion process to the apex of the glenoid</td>
|
||
<td id="768F824038150014FBF6F9E79789ABE7" box="[1090,1228,1653,1675]" gridcol="1" gridrow="2" pageId="2" pageNumber="249">128</td>
|
||
</tr>
|
||
<tr id="355EEB3C38150014FF2BF90C9789ABD8" box="[159,1228,1694,1716]" gridrow="3" pageId="2" pageNumber="249">
|
||
<td id="768F824038150014FF2BF90C976EABD8" box="[159,1067,1694,1716]" gridcol="0" gridrow="3" pageId="2" pageNumber="249">Minimum width of the scapular ‘neck’</td>
|
||
<td id="768F824038150014FBF6F90C9789ABD8" box="[1090,1228,1694,1716]" gridcol="1" gridrow="3" pageId="2" pageNumber="249">40</td>
|
||
</tr>
|
||
<tr id="355EEB3C38150014FF2BF95A9789ABB2" box="[159,1228,1736,1758]" gridrow="4" pageId="2" pageNumber="249">
|
||
<td id="768F824038150014FF2BF95A976EABB2" box="[159,1067,1736,1758]" gridcol="0" gridrow="4" pageId="2" pageNumber="249">Maximum width of the distal scapular blade (as preserved)</td>
|
||
<td id="768F824038150014FBF6F95A9789ABB2" box="[1090,1228,1736,1758]" gridcol="1" gridrow="4" pageId="2" pageNumber="249">98</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<caption id="DF11B9F63814FFE8FF23F8DE92B0AAED" httpUri="https://zenodo.org/record/2624341/files/figure.png" pageId="3" pageNumber="250" startId="3.[151,250,1868,1890]" targetBox="[177,1410,254,1733]" targetPageId="3">
|
||
<paragraph id="8BD1E97E3814FFE8FF23F8DE92B0AAED" blockId="3.[151,1436,1868,1921]" pageId="3" pageNumber="250">
|
||
<emphasis id="B91A356C3814FFE8FF23F8DE9248AA0E" bold="true" box="[151,269,1868,1890]" pageId="3" pageNumber="250">FIGURE 2</emphasis>
|
||
. Holotype specimen of
|
||
<taxonomicName id="4C6E92FD3814FFE8FDB4F8DC9194AA0F" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[512,721,1870,1891]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="250" phylum="Chordata" rank="species" species="evrostos">
|
||
<emphasis id="B91A356C3814FFE8FDB4F8DC9194AA0F" box="[512,721,1870,1891]" italics="true" pageId="3" pageNumber="250">Pareisactus evrostos</emphasis>
|
||
</taxonomicName>
|
||
, a nearly complete left scapula (MCD 5371) in (
|
||
<emphasis id="B91A356C3814FFE8FB76F8DE9791AA0E" bold="true" box="[1218,1236,1868,1890]" pageId="3" pageNumber="250">A</emphasis>
|
||
) dorsal, (
|
||
<emphasis id="B91A356C3814FFE8FA82F8DE9602AA0E" bold="true" box="[1334,1351,1868,1890]" pageId="3" pageNumber="250">B</emphasis>
|
||
) lateral, (
|
||
<emphasis id="B91A356C3814FFE8FF2BF8F993F4AAED" bold="true" box="[159,177,1899,1921]" pageId="3" pageNumber="250">C</emphasis>
|
||
) medial, and (
|
||
<emphasis id="B91A356C3814FFE8FEF1F8F99212AAED" bold="true" box="[325,343,1899,1921]" pageId="3" pageNumber="250">D</emphasis>
|
||
) ventral views.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BD1E97E3813FFEFFF73FF05905EAF9C" blockId="4.[151,1437,151,1980]" pageId="4" pageNumber="251">
|
||
The dorsal region of the proximal region of the scapula of
|
||
<taxonomicName id="4C6E92FD3813FFEFFC07FF0B97E7ADDC" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[947,1186,153,176]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="species" species="evrostos">
|
||
<emphasis id="B91A356C3813FFEFFC07FF0B97E7ADDC" box="[947,1186,153,176]" italics="true" pageId="4" pageNumber="251">Pareisactus evrostos</emphasis>
|
||
</taxonomicName>
|
||
is greatly expanded mediolaterally, so that the proximal constriction (‘scapular neck’) is as dorsoventrally deep as it is mediolaterally thick (
|
||
<figureCitation id="1355F5FB3813FFEFFF6AFF4D923AAD94" box="[222,383,223,248]" captionStart="FIGURE 2" captionStartId="3.[151,250,1868,1890]" captionTargetBox="[177,1410,254,1733]" captionTargetId="figure@3.[151,1436,223,1773]" captionTargetPageId="3" captionText="FIGURE 2. Holotype specimen of Pareisactus evrostos, a nearly complete left scapula (MCD 5371) in (A) dorsal, (B) lateral, (C) medial, and (D) ventral views." httpUri="https://zenodo.org/record/2624341/files/figure.png" pageId="4" pageNumber="251">Fig. 2A and B</figureCitation>
|
||
). The deltoid fossa is extremely deep and its lateral surface projects ventrally to form the apex of the glenoid (
|
||
<figureCitation id="1355F5FB3813FFEFFEFCFE9692E7AC70" box="[328,418,260,285]" captionStart="FIGURE 2" captionStartId="3.[151,250,1868,1890]" captionTargetBox="[177,1410,254,1733]" captionTargetId="figure@3.[151,1436,223,1773]" captionTargetPageId="3" captionText="FIGURE 2. Holotype specimen of Pareisactus evrostos, a nearly complete left scapula (MCD 5371) in (A) dorsal, (B) lateral, (C) medial, and (D) ventral views." httpUri="https://zenodo.org/record/2624341/files/figure.png" pageId="4" pageNumber="251">Fig. 2B</figureCitation>
|
||
). This lateral surface under the deltoid ridge leading into the glenoid apex is remarkably long, accounting for half of the maximum depth of the proximal region of the scapula including the acromion process (
|
||
<figureCitation id="1355F5FB3813FFEFFF4AFEDE921CAC08" box="[254,345,332,357]" captionStart="FIGURE 2" captionStartId="3.[151,250,1868,1890]" captionTargetBox="[177,1410,254,1733]" captionTargetId="figure@3.[151,1436,223,1773]" captionTargetPageId="3" captionText="FIGURE 2. Holotype specimen of Pareisactus evrostos, a nearly complete left scapula (MCD 5371) in (A) dorsal, (B) lateral, (C) medial, and (D) ventral views." httpUri="https://zenodo.org/record/2624341/files/figure.png" pageId="4" pageNumber="251">Fig. 2B</figureCitation>
|
||
). The deltoid fossa is roofed by a prominent deltoid ridge. This ridge extends caudoventrally becoming continuous with the ventral margin of the scapular blade (
|
||
<figureCitation id="1355F5FB3813FFEFFC20FEFD90A9ACE4" box="[916,1004,367,392]" captionStart="FIGURE 2" captionStartId="3.[151,250,1868,1890]" captionTargetBox="[177,1410,254,1733]" captionTargetId="figure@3.[151,1436,223,1773]" captionTargetPageId="3" captionText="FIGURE 2. Holotype specimen of Pareisactus evrostos, a nearly complete left scapula (MCD 5371) in (A) dorsal, (B) lateral, (C) medial, and (D) ventral views." httpUri="https://zenodo.org/record/2624341/files/figure.png" pageId="4" pageNumber="251">Fig. 2B</figureCitation>
|
||
). At the cranial end of the scapula, the dorsal margin curves dorsally and laterally to form a mediolaterally broad acromion process (
|
||
<figureCitation id="1355F5FB3813FFEFFB06FE069610ACC0" box="[1202,1365,404,429]" captionStart="FIGURE 2" captionStartId="3.[151,250,1868,1890]" captionTargetBox="[177,1410,254,1733]" captionTargetId="figure@3.[151,1436,223,1773]" captionTargetPageId="3" captionText="FIGURE 2. Holotype specimen of Pareisactus evrostos, a nearly complete left scapula (MCD 5371) in (A) dorsal, (B) lateral, (C) medial, and (D) ventral views." httpUri="https://zenodo.org/record/2624341/files/figure.png" pageId="4" pageNumber="251">Fig. 2A and B</figureCitation>
|
||
). This process projects craniodorsally, as well as craniolaterally, and its lateral profile appears nearly parallel to the eroded cranial margin of the glenoid (
|
||
<figureCitation id="1355F5FB3813FFEFFE5DFE4E9105AC98" box="[489,576,476,501]" captionStart="FIGURE 2" captionStartId="3.[151,250,1868,1890]" captionTargetBox="[177,1410,254,1733]" captionTargetId="figure@3.[151,1436,223,1773]" captionTargetPageId="3" captionText="FIGURE 2. Holotype specimen of Pareisactus evrostos, a nearly complete left scapula (MCD 5371) in (A) dorsal, (B) lateral, (C) medial, and (D) ventral views." httpUri="https://zenodo.org/record/2624341/files/figure.png" pageId="4" pageNumber="251">Fig. 2B</figureCitation>
|
||
). The acromion process of
|
||
<taxonomicName id="4C6E92FD3813FFEFFCDBFE4F90A6AC98" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[879,995,477,500]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="species" species="evrostos">
|
||
<emphasis id="B91A356C3813FFEFFCDBFE4F90A6AC98" box="[879,995,477,500]" italics="true" pageId="4" pageNumber="251">P. evrostos</emphasis>
|
||
</taxonomicName>
|
||
extends above the dorsal margin of the scapula a shorter distance than in other ornithopods such as
|
||
<taxonomicName id="4C6E92FD3813FFEFFCF6FE6D97A7AF74" authority="(Scheetz 1999)" authorityName="Scheetz" authorityYear="1999" baseAuthorityName="Scheetz" baseAuthorityYear="1999" box="[834,1250,511,536]" class="Reptilia" family="Thescelosauridae" genus="Orodromeus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="species" species="makelai">
|
||
<emphasis id="B91A356C3813FFEFFCF6FE6D976BAF74" box="[834,1070,511,536]" italics="true" pageId="4" pageNumber="251">Orodromeus makelai</emphasis>
|
||
(
|
||
<bibRefCitation id="EFFF948F3813FFEFFB89FE6D979FAF74" author="Scheetz, R. D." box="[1085,1242,511,536]" pageId="4" pageNumber="251" refId="ref7246" refString="Scheetz, R. D. (1999) Osteology of Orodromeus makelai and the phylogeny of basal ornithopod dinosaurs. Montana State University, Bozeman, 186 pp." type="book" year="1999">Scheetz 1999</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="4C6E92FD3813FFEFFB5BFE6D9161AF50" authority="(Varricchio et al. 2007)" authorityName="Varricchio" authorityYear="2007" baseAuthorityName="Varricchio" baseAuthorityYear="2007" class="Reptilia" family="Thescelosauridae" genus="Oryctodromeus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="species" species="cubicularis">
|
||
<emphasis id="B91A356C3813FFEFFB5BFE6D9250AF51" italics="true" pageId="4" pageNumber="251">Oryctodromeus cubicularis</emphasis>
|
||
(
|
||
<bibRefCitation id="EFFF948F3813FFEFFE97FDB69159AF50" author="Varricchio, D. J. & Martin, A. J. & Katsura, Y." box="[291,540,548,573]" pageId="4" pageNumber="251" pagination="1361 - 1368" refId="ref7496" refString="Varricchio, D. J., Martin, A. J. & Katsura, Y. (2007) First trace and body fossil evidence of a burrowing, denning dinosaur. Proceedings of the Royal Society B, 274, 1361 - 1368. https: // doi. org / 10.1098 / rspb. 2006.0443" type="journal article" year="2007">
|
||
Varricchio
|
||
<emphasis id="B91A356C3813FFEFFE15FDB7929FAF51" box="[417,474,548,573]" italics="true" pageId="4" pageNumber="251">et al.</emphasis>
|
||
2007
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
,
|
||
<emphasis id="B91A356C3813FFEFFD84FDB791EAAF50" box="[560,687,549,572]" italics="true" pageId="4" pageNumber="251">Haya griva</emphasis>
|
||
(
|
||
<bibRefCitation id="EFFF948F3813FFEFFD0AFDB690FAAF50" author="Makovicky, P. J. & Kilbourne, B. M. & Sadleir, R. W. & Norell, M. A." box="[702,959,548,573]" pageId="4" pageNumber="251" pagination="626 - 640" refId="ref6165" refString="Makovicky, P. J., Kilbourne, B. M., Sadleir, R. W. & Norell, M. A. (2011) A new basal ornithopod (Dinosauria, Ornithischia) from the Late Cretaceous of Mongolia. Journal of Fertebrate Paleontology, 31, 626 - 640. https: // doi. org / 10.1080 / 02724634.2011.557114" type="journal article" year="2011">
|
||
Makovicky
|
||
<emphasis id="B91A356C3813FFEFFCF3FDB79032AF51" box="[839,887,548,573]" italics="true" pageId="4" pageNumber="251">et al</emphasis>
|
||
. 2011
|
||
</bibRefCitation>
|
||
) or
|
||
<emphasis id="B91A356C3813FFEFFC58FDB7979AAF51" box="[1004,1247,548,573]" italics="true" pageId="4" pageNumber="251">Camptosaurus dispar</emphasis>
|
||
(
|
||
<bibRefCitation id="EFFF948F3813FFEFFB59FDB696D5AF50" author="Gilmore, C. W." box="[1261,1424,548,573]" pageId="4" pageNumber="251" pagination="197 - 332" refId="ref5847" refString="Gilmore, C. W. (1909) Osteology of the Jurassic reptile Camptosaurus, with a revision of the species of the genus, and descriptions of two new species. Proceedings of the United States National Museum, 36, 197 - 332. https: // doi. org / 10.5479 / si. 00963801.36 - 1666.197" type="journal article" year="1909">Gilmore 1909</bibRefCitation>
|
||
), but a longer distance than in Hadrosauriformes (
|
||
<bibRefCitation id="EFFF948F3813FFEFFD45FDDA9724AF0C" author="Brett-Surman, M. K. & Wagner, J. R." box="[753,1121,584,608]" pageId="4" pageNumber="251" pagination="135 - 169" refId="ref4318" refString="Brett-Surman, M. K. & Wagner, J. R. (2007) Discussion of character analysis of the appendicular anatomy in Campanian and Maastrichtian North American hadrosaurids-variation and ontogeny. In: Carpenter, K. (Ed.), Horns and Beaks. Ceratopsian and Ornithopod Dinosaurs. Indiana University Press, Bloomington, pp. 135 - 169." type="book chapter" year="2007">Brett-Surman & Wagner 2007</bibRefCitation>
|
||
) and at least some basal styracosternans like
|
||
<taxonomicName id="4C6E92FD3813FFEFFE3CFDFE91B4AFE8" authority="(Wu 2010)" authorityName="Wu" authorityYear="2010" baseAuthorityName="Wu" baseAuthorityYear="2010" box="[392,753,620,645]" class="Reptilia" family="Iguanodontidae" genus="Bolong" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="species" species="yixianensis">
|
||
<emphasis id="B91A356C3813FFEFFE3CFDFE9126AFE8" box="[392,611,620,645]" italics="true" pageId="4" pageNumber="251">Bolong yixianensis</emphasis>
|
||
(
|
||
<bibRefCitation id="EFFF948F3813FFEFFDCCFDFE91ACAFE8" author="Wu, W. - H." box="[632,745,620,644]" pageId="4" pageNumber="251" refId="ref7841" refString="Wu, W. - H. (2010) Bolong yixianensis Wu et al. 2010 from the Yixian Formation in western Liaoning, China and the phylogeny of basal Iguanodontia. Yilin University, Chanchung, 154 pp." type="book" year="2010">Wu 2010</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="1355F5FB3813FFEFFCB2FDFE9015AFE8" box="[774,848,620,645]" captionStart="FIGURE 3" captionStartId="5.[151,250,1703,1725]" captionTargetBox="[151,1436,195,1651]" captionTargetId="figure@5.[151,1436,195,1651]" captionTargetPageId="5" captionText="FIGURE 3. Strict consensus tree derived from the parsimony analysis showing the position of Pareisactus evrostos among iguanodontian ornithopods. Depicted specimens in left lateral view are: Bolong yixianensis, YHZ-001 (line drawing based on Wu 2010, plate III); Camptosaurus dispar, USNM 5955; Corythosaurus casuarius, ROM 845; Cumnoria prestwichii, OXFUM J.3303; Eolambia caroljonesa, CEUM 52097 (reversed, line drawing based on McDonald et al. 2012, fig. 27C); Gilmoreosaurus mongoliensis, AMNH FARB 30725; Haya griva, IGM 100-2015; Iguanodon bernissartensis, IRSNB 1534; Jinzhousaurus yangi, IVPP V12691 (line drawing based on Wang et al. 2011, fig. 4); Lophorhothon atopus, AUMP 2295; Orodromeus makelai, MOR 294 (reversed); Oryctodromeus cubicularis, MOR 1636; Ouranosaurus nigeriensis, cast of MNHN GDF 300; Pareisactus evrostos, MCD 5371; Rhabdodon priscus, MDE-C3 1013; Tenontosaurus tilletti, MOR 2571; Uteodon aphanoecetes, CM 11337; Zalmoxes robustus, line drawing from Weishampel et al. [2003, fig. 19C]); Z. shqiperorum, UBB NVZ1-24 (reversed, line drawing based on Godefroit et al. 2009, fig. 15A)." httpUri="https://zenodo.org/record/2624343/files/figure.png" pageId="4" pageNumber="251">Fig. 3</figureCitation>
|
||
). As in non-hadrosauriform iguanodontians (e.g.,
|
||
<bibRefCitation id="EFFF948F3813FFEFFF23FD1D9214AFC4" author="McDonald, A. T." box="[151,337,655,680]" pageId="4" pageNumber="251" pagination="52 - 68" refId="ref6484" refString="McDonald, A. T. (2011) The taxonomy of species assigned to Camptosaurus (Dinosauria: Ornithopoda). Zootaxa, 2783, 52 - 68." type="journal article" year="2011">
|
||
<collectingCountry id="F379A9EE3813FFEFFF23FD1D924AAFC4" box="[151,271,655,680]" name="Heard Island and McDonald Islands" pageId="4" pageNumber="251">McDonald</collectingCountry>
|
||
2011
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFFF948F3813FFEFFEEBFD0292BAAFC4" author="Norman, D. B." box="[351,511,656,680]" pageId="4" pageNumber="251" pagination="165 - 194" refId="ref6637" refString="Norman, D. B. (2011) On the osteology of the lower Wealden (Valanginian) ornithopod Barilium dawsoni (Iguanodontia: Styracosterna). Palaeontology, 86, 165 - 194." type="journal article" year="2011">Norman 2011</bibRefCitation>
|
||
), and particularly more distant outgroups such as rhabdodontids (e.g.,
|
||
<bibRefCitation id="EFFF948F3813FFEFFA90FD1D9394AFA0" author="Chanthasit, P." pageId="4" pageNumber="251" refId="ref4765" refString="Chanthasit, P. (2010) The ornithopod dinosaur Rhabdodon from the Late Cretaceous of France: anatomy, systematics and paleobiology. Universite Claude Bernard, Lyon, 196 pp." type="book" year="2010">Chanthasit 2010</bibRefCitation>
|
||
) or jeholosaurids (e.g.,
|
||
<bibRefCitation id="EFFF948F3813FFEFFE5FFD2691FAAFA0" author="Butler, R. J. & Jin, L. & Chen, J. & Godefroit, P." box="[491,703,692,717]" pageId="4" pageNumber="251" pagination="667 - 683" refId="ref4491" refString="Butler, R. J., Jin, L., Chen, J. & Godefroit, P. (2011) Phylogenetic position of the small ornithischian dinosaur Changchunsaurus parvus from the Quantou Formation (Cretaceous: Aptian-Cenomanian) of Jilin Province, north-eastern China. Palaeontology, 54, 667 - 683. https: // doi. org / 10.1111 / j. 1475 - 4983.2011.01046. x" type="journal article" year="2011">
|
||
Butler
|
||
<emphasis id="B91A356C3813FFEFFD88FD279137AFA1" box="[572,626,692,717]" italics="true" pageId="4" pageNumber="251">et al</emphasis>
|
||
. 2011
|
||
</bibRefCitation>
|
||
), the apex of the acromion process is strongly offset cranially relative to the position of the apex of the glenoid (
|
||
<figureCitation id="1355F5FB3813FFEFFD73FD459049AF9C" box="[711,780,727,752]" captionStart="FIGURE 3" captionStartId="5.[151,250,1703,1725]" captionTargetBox="[151,1436,195,1651]" captionTargetId="figure@5.[151,1436,195,1651]" captionTargetPageId="5" captionText="FIGURE 3. Strict consensus tree derived from the parsimony analysis showing the position of Pareisactus evrostos among iguanodontian ornithopods. Depicted specimens in left lateral view are: Bolong yixianensis, YHZ-001 (line drawing based on Wu 2010, plate III); Camptosaurus dispar, USNM 5955; Corythosaurus casuarius, ROM 845; Cumnoria prestwichii, OXFUM J.3303; Eolambia caroljonesa, CEUM 52097 (reversed, line drawing based on McDonald et al. 2012, fig. 27C); Gilmoreosaurus mongoliensis, AMNH FARB 30725; Haya griva, IGM 100-2015; Iguanodon bernissartensis, IRSNB 1534; Jinzhousaurus yangi, IVPP V12691 (line drawing based on Wang et al. 2011, fig. 4); Lophorhothon atopus, AUMP 2295; Orodromeus makelai, MOR 294 (reversed); Oryctodromeus cubicularis, MOR 1636; Ouranosaurus nigeriensis, cast of MNHN GDF 300; Pareisactus evrostos, MCD 5371; Rhabdodon priscus, MDE-C3 1013; Tenontosaurus tilletti, MOR 2571; Uteodon aphanoecetes, CM 11337; Zalmoxes robustus, line drawing from Weishampel et al. [2003, fig. 19C]); Z. shqiperorum, UBB NVZ1-24 (reversed, line drawing based on Godefroit et al. 2009, fig. 15A)." httpUri="https://zenodo.org/record/2624343/files/figure.png" pageId="4" pageNumber="251">Fig. 3</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BD1E97E3813FFEFFF73FD6E9070A958" blockId="4.[151,1437,151,1980]" pageId="4" pageNumber="251">
|
||
Caudally, the paddle-shaped scapular blade gradually expands dorsoventrally, more ventrally than dorsally, while progressively thinning mediolaterally to become a delicate lamina at its distal end. As preserved, the maximum distal expansion of the scapular blade is 90 percent of the maximum depth of the proximal end (from the apex of the acromion process to the apex of the glenoid) and two and half times deeper than the proximal constriction. The total length of the scapula (from the apex of the acromion process to the distal end of the scapular blade) is only 2.4 times the maximum depth of the proximal end. Such relatively short scapulae, with length/ proximal depth ratios less than three, are typically found in styracosternan-outgroup ornithopods such as
|
||
<taxonomicName id="4C6E92FD3813FFEFFF23FC65917AA97C" authority="(Scheetz 1999)" authorityName="Scheetz" authorityYear="1999" baseAuthorityName="Scheetz" baseAuthorityYear="1999" box="[151,575,1015,1040]" class="Reptilia" family="Thescelosauridae" genus="Orodromeus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="species" species="makelai">
|
||
<emphasis id="B91A356C3813FFEFFF23FC6592C0A97C" box="[151,389,1015,1040]" italics="true" pageId="4" pageNumber="251">Orodromeus makelai</emphasis>
|
||
(
|
||
<bibRefCitation id="EFFF948F3813FFEFFE2CFC659172A97C" author="Scheetz, R. D." box="[408,567,1015,1040]" pageId="4" pageNumber="251" refId="ref7246" refString="Scheetz, R. D. (1999) Osteology of Orodromeus makelai and the phylogeny of basal ornithopod dinosaurs. Montana State University, Bozeman, 186 pp." type="book" year="1999">Scheetz 1999</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
, jeholosaurids (
|
||
<bibRefCitation id="EFFF948F3813FFEFFD42FC65908CA97C" author="Butler, R. J. & Jin, L. & Chen, J. & Godefroit, P." box="[758,969,1015,1040]" pageId="4" pageNumber="251" pagination="667 - 683" refId="ref4491" refString="Butler, R. J., Jin, L., Chen, J. & Godefroit, P. (2011) Phylogenetic position of the small ornithischian dinosaur Changchunsaurus parvus from the Quantou Formation (Cretaceous: Aptian-Cenomanian) of Jilin Province, north-eastern China. Palaeontology, 54, 667 - 683. https: // doi. org / 10.1111 / j. 1475 - 4983.2011.01046. x" type="journal article" year="2011">
|
||
Butler
|
||
<emphasis id="B91A356C3813FFEFFCFDFC6B90C0A97C" box="[841,901,1015,1040]" italics="true" pageId="4" pageNumber="251">et al.</emphasis>
|
||
2011
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFFF948F3813FFEFFC6DFC6597A0A97C" author="Makovicky, P. J. & Kilbourne, B. M. & Sadleir, R. W. & Norell, M. A." box="[985,1253,1015,1040]" pageId="4" pageNumber="251" pagination="626 - 640" refId="ref6165" refString="Makovicky, P. J., Kilbourne, B. M., Sadleir, R. W. & Norell, M. A. (2011) A new basal ornithopod (Dinosauria, Ornithischia) from the Late Cretaceous of Mongolia. Journal of Fertebrate Paleontology, 31, 626 - 640. https: // doi. org / 10.1080 / 02724634.2011.557114" type="journal article" year="2011">
|
||
Makovicky
|
||
<emphasis id="B91A356C3813FFEFFBD7FC6B97DAA97C" box="[1123,1183,1015,1040]" italics="true" pageId="4" pageNumber="251">et al.</emphasis>
|
||
2011
|
||
</bibRefCitation>
|
||
), rhabdodontids (
|
||
<bibRefCitation id="EFFF948F3813FFEFFF2AFB8E921CA958" author="Chanthasit, P." box="[158,345,1052,1077]" pageId="4" pageNumber="251" refId="ref4765" refString="Chanthasit, P. (2010) The ornithopod dinosaur Rhabdodon from the Late Cretaceous of France: anatomy, systematics and paleobiology. Universite Claude Bernard, Lyon, 196 pp." type="book" year="2010">Chanthasit 2010</bibRefCitation>
|
||
) or
|
||
<emphasis id="B91A356C3813FFEFFE31FB8F913DA959" box="[389,632,1052,1077]" italics="true" pageId="4" pageNumber="251">Camptosaurus dispar</emphasis>
|
||
(
|
||
<bibRefCitation id="EFFF948F3813FFEFFD32FB8E906DA958" author="Gilmore, C. W." box="[646,808,1052,1077]" pageId="4" pageNumber="251" pagination="197 - 332" refId="ref5847" refString="Gilmore, C. W. (1909) Osteology of the Jurassic reptile Camptosaurus, with a revision of the species of the genus, and descriptions of two new species. Proceedings of the United States National Museum, 36, 197 - 332. https: // doi. org / 10.5479 / si. 00963801.36 - 1666.197" type="journal article" year="1909">Gilmore 1909</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BD1E97E3813FFEFFF73FBD1978FAB38" blockId="4.[151,1437,151,1980]" pageId="4" pageNumber="251">
|
||
<emphasis id="B91A356C3813FFEFFF73FBD191C7A937" bold="true" box="[199,642,1091,1116]" pageId="4" pageNumber="251">
|
||
Relationships of
|
||
<emphasis id="B91A356C3813FFEFFE25FBD691C7A937" bold="true" box="[401,642,1092,1116]" italics="true" pageId="4" pageNumber="251">
|
||
<taxonomicName id="4C6E92FD3813FFEFFE25FBD6913AA937" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[401,639,1092,1116]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="species" species="evrostos">Pareisactus evrostos</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
</emphasis>
|
||
The phylogenetic position of
|
||
<taxonomicName id="4C6E92FD3813FFEFFC6EFBD79787A930" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[986,1218,1093,1116]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="species" species="evrostos">
|
||
<emphasis id="B91A356C3813FFEFFC6EFBD79787A930" box="[986,1218,1093,1116]" italics="true" pageId="4" pageNumber="251">Pareisactus evrostos</emphasis>
|
||
</taxonomicName>
|
||
was inferred using Parsimony. The taxonomic sample included 52 ornithopod species and
|
||
<emphasis id="B91A356C3813FFEFFC64FBF5964EA9EC" box="[976,1291,1127,1152]" italics="true" pageId="4" pageNumber="251">Lesothosaurus diagnosticus</emphasis>
|
||
as outgroup taxon. We included the unnamed lambeosaurine (
|
||
<bibRefCitation id="EFFF948F3813FFEFFD7AFB19908AA9C8" author="Fondevilla, V. & Dalla Vecchia, F. M. & Gaete, R. & Galobart, A. & Moncunill-Sole, B. & Kohler, M." box="[718,975,1163,1188]" pageId="4" pageNumber="251" pagination="0206287" refId="ref5598" refString="Fondevilla, V., Dalla Vecchia, F. M., Gaete, R., Galobart, A., Moncunill-Sole, B. & Kohler, M. (2018) Ontogeny and taxonomy of the hadrosaur (Dinosauria, Ornithopoda) remains from Basturs Poble bonebed (late early Maastrichtian, Tremp Syncline, Spain). PLoS ONE, 12, e 0206287. https: // doi. org / 10.1371 / journal. pone. 0206287" type="journal article" year="2018">
|
||
Fondevilla
|
||
<emphasis id="B91A356C3813FFEFFCE5FB1F90C9A9C8" box="[849,908,1163,1188]" italics="true" pageId="4" pageNumber="251">et al.</emphasis>
|
||
2018
|
||
</bibRefCitation>
|
||
) from the Basturs Poble bonebed to the analysis. The subadult elements of the Basturs Poble lambeosaurine were scored only for those characters the states of which do not vary during ontogeny, according to Prieto-Márquez (2011) and Prieto-Márquez & Guenther (2018). The data set (see Supporting Material) consisted of the 134 morphological characters of
|
||
<bibRefCitation id="EFFF948F3813FFEFFAB6FB6593A1A858" author="Madzia, D. & Boyd, C. A. & Mazuch, M." pageId="4" pageNumber="251" pagination="967 - 979" refId="ref6113" refString="Madzia, D., Boyd, C. A. & Mazuch, M. (2018) A basal ornithopod dinosaur from the Cenomanian of the Czech Republic. Journal of Systematic Palaeontology, 16, 967 - 979. https: // doi. org / 10.1080 / 14772019.2017.1371258" type="journal article" year="2018">
|
||
Madzia
|
||
<emphasis id="B91A356C3813FFEFFAD4FB6B96D9A87C" box="[1376,1436,1271,1296]" italics="true" pageId="4" pageNumber="251">et al.</emphasis>
|
||
(2018)
|
||
</bibRefCitation>
|
||
, to which six more characters were added (totaling 94 cranial and 46 postcranial discrete morphological characters). Multistate characters containing states that are not mutually exclusive, following a natural morphocline, were ordered. This criterion allows for ‘crediting’ shared intermediate states. The search for the optimal tree(s) was conducted in TNT version 1.1 (
|
||
<bibRefCitation id="EFFF948F3813FFEFFD55FA159094A8CC" author="Goloboff, P. A. & Farris, J. S. & Nixon, K. C." box="[737,977,1415,1440]" pageId="4" pageNumber="251" pagination="774 - 786" refId="ref5974" refString="Goloboff, P. A., Farris, J. S. & Nixon, K. C. (2008) TNT, a free program for phylogenetic analysis. Cladistics, 24, 774 - 786. https: // doi. org / 10.1111 / j. 1096 - 0031.2008.00217. x" type="journal article" year="2008">
|
||
Goloboff
|
||
<emphasis id="B91A356C3813FFEFFCE6FA1B90C9A8CC" box="[850,908,1415,1440]" italics="true" pageId="4" pageNumber="251">et al.</emphasis>
|
||
2008
|
||
</bibRefCitation>
|
||
). A heuristic search of 10,000 replicates using random additional sequences was performed, followed by branch swapping by tree-bisection-reconnection holding ten trees per replicate. Bremer support (
|
||
<bibRefCitation id="EFFF948F3813FFEFFD53FA4290C9A884" author="Bremer, K." box="[743,908,1488,1512]" pageId="4" pageNumber="251" pagination="795 - 803" refId="ref4272" refString="Bremer, K. (1988) The limits of amino acid sequence data in angiosperm phylogenetic reconstruction. Evolution, 42, 795 - 803. https: // doi. org / 10.1111 / j. 1558 - 5646.1988. tb 02497. x" type="journal article" year="1988">Bremer, 1988</bibRefCitation>
|
||
) was assessed by computing decay indices (
|
||
<bibRefCitation id="EFFF948F3813FFEFFF2AFA6192E5AB60" author="Donoghue, M. J. & Olmstead, R. G. & Smith, J. F. & Palmer, J. D." box="[158,416,1523,1548]" pageId="4" pageNumber="251" pagination="672 - 685" refId="ref5164" refString="Donoghue, M. J., Olmstead, R. G., Smith, J. F. & Palmer, J. D. (1992) Phylogenetic relationships of Dipsacales based on rbcL sequences. Annals of the Missouri Botanical Garden, 79, 672 - 685. https: // doi. org / 10.2307 / 2399772" type="journal article" year="1992">
|
||
Donoghue
|
||
<emphasis id="B91A356C3813FFEFFE94FA679219AB60" box="[288,348,1523,1548]" italics="true" pageId="4" pageNumber="251">et al.</emphasis>
|
||
1992
|
||
</bibRefCitation>
|
||
) using TNT. Bootstrap proportions (
|
||
<bibRefCitation id="EFFF948F3813FFEFFCF8FA619755AB60" author="Felsenstein, J." box="[844,1040,1523,1548]" pageId="4" pageNumber="251" pagination="783 - 791" refId="ref5383" refString="Felsenstein, J. (1985) Confidence limits on phylogenies: an approach using the bootstrap. Evolution, 39, 783 - 791. https: // doi. org / 10.1111 / j. 1558 - 5646.1985. tb 00420. x" type="journal article" year="1985">Felsenstein 1985</bibRefCitation>
|
||
) were also calculated using TNT, setting the analysis for 5,000 replicates using heuristic searches, in which each search was conducted using random additional sequences with branch-swapping by subtree pruning and regrafting and 25 replicates.
|
||
</paragraph>
|
||
<paragraph id="8BD1E97E3813FFEFFF73F9CD9290AAD0" blockId="4.[151,1437,151,1980]" pageId="4" pageNumber="251">
|
||
The phylogenetic analysis resulted in 12 most parsimonious trees of 504 steps each (C.I. = 0.45, R.I. = 0.80); the best score was found in 904 of the 10,000 replicates.
|
||
<taxonomicName id="4C6E92FD3813FFEFFCDFF917971CABF0" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[875,1113,1669,1692]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="species" species="evrostos">
|
||
<emphasis id="B91A356C3813FFEFFCDFF917971CABF0" box="[875,1113,1669,1692]" italics="true" pageId="4" pageNumber="251">Pareisactus evrostos</emphasis>
|
||
</taxonomicName>
|
||
was recovered as a basal iguanodontian, nested within
|
||
<taxonomicName id="4C6E92FD3813FFEFFE5CF93591DAABAC" box="[488,671,1703,1728]" class="Reptilia" family="Rhabdodontidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="family">Rhabdodontidae</taxonomicName>
|
||
as the sister taxon to
|
||
<emphasis id="B91A356C3813FFEFFC29F935973DABAC" box="[925,1144,1703,1728]" italics="true" pageId="4" pageNumber="251">Rhabdodon priscus</emphasis>
|
||
(
|
||
<figureCitation id="1355F5FB3813FFEFFB3EF9359795ABAC" box="[1162,1232,1703,1728]" captionStart="FIGURE 3" captionStartId="5.[151,250,1703,1725]" captionTargetBox="[151,1436,195,1651]" captionTargetId="figure@5.[151,1436,195,1651]" captionTargetPageId="5" captionText="FIGURE 3. Strict consensus tree derived from the parsimony analysis showing the position of Pareisactus evrostos among iguanodontian ornithopods. Depicted specimens in left lateral view are: Bolong yixianensis, YHZ-001 (line drawing based on Wu 2010, plate III); Camptosaurus dispar, USNM 5955; Corythosaurus casuarius, ROM 845; Cumnoria prestwichii, OXFUM J.3303; Eolambia caroljonesa, CEUM 52097 (reversed, line drawing based on McDonald et al. 2012, fig. 27C); Gilmoreosaurus mongoliensis, AMNH FARB 30725; Haya griva, IGM 100-2015; Iguanodon bernissartensis, IRSNB 1534; Jinzhousaurus yangi, IVPP V12691 (line drawing based on Wang et al. 2011, fig. 4); Lophorhothon atopus, AUMP 2295; Orodromeus makelai, MOR 294 (reversed); Oryctodromeus cubicularis, MOR 1636; Ouranosaurus nigeriensis, cast of MNHN GDF 300; Pareisactus evrostos, MCD 5371; Rhabdodon priscus, MDE-C3 1013; Tenontosaurus tilletti, MOR 2571; Uteodon aphanoecetes, CM 11337; Zalmoxes robustus, line drawing from Weishampel et al. [2003, fig. 19C]); Z. shqiperorum, UBB NVZ1-24 (reversed, line drawing based on Godefroit et al. 2009, fig. 15A)." httpUri="https://zenodo.org/record/2624343/files/figure.png" pageId="4" pageNumber="251">Fig. 3</figureCitation>
|
||
).
|
||
<taxonomicName id="4C6E92FD3813FFEFFB53F93B962FABAC" authority="Párraga & Prieto-Márquez, 2019" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[1255,1386,1705,1728]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="249" phylum="Chordata" rank="genus" status="gen. nov.">
|
||
<emphasis id="B91A356C3813FFEFFB53F93B962FABAC" box="[1255,1386,1705,1728]" italics="true" pageId="4" pageNumber="251">Pareisactus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis id="B91A356C3813FFEFFF23F959925DAB88" box="[151,280,1739,1764]" italics="true" pageId="4" pageNumber="251">Rhabdodon</emphasis>
|
||
are unambiguously united by the presence of a well-demarcated deltoid ridge. However, this sister relationship is weakly supported, as indicated by the low decay indices and Boostrap proportions below 50%. We tested how many more steps would less parsimonious trees have when forcing
|
||
<taxonomicName id="4C6E92FD3813FFEFFB97F88797E3AA40" authority="Párraga & Prieto-Márquez, 2019" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[1059,1190,1813,1836]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="249" phylum="Chordata" rank="genus" status="gen. nov.">
|
||
<emphasis id="B91A356C3813FFEFFB97F88797E3AA40" box="[1059,1190,1813,1836]" italics="true" pageId="4" pageNumber="251">Pareisactus</emphasis>
|
||
</taxonomicName>
|
||
to be placed in other areas of the iguanodontian tree. In particular, moving
|
||
<taxonomicName id="4C6E92FD3813FFEFFCB5F8AB90C1AA3C" authority="Párraga & Prieto-Márquez, 2019" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[769,900,1849,1872]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="249" phylum="Chordata" rank="genus" status="gen. nov.">
|
||
<emphasis id="B91A356C3813FFEFFCB5F8AB90C1AA3C" box="[769,900,1849,1872]" italics="true" pageId="4" pageNumber="251">Pareisactus</emphasis>
|
||
</taxonomicName>
|
||
outside Iguanodontia caused the tree length to increase in one more step, as it did when positioned as sister to
|
||
<taxonomicName id="4C6E92FD3813FFEFFCDAF8C99759AA18" authorityName="Gilmore" authorityYear="1913" box="[878,1052,1883,1908]" class="Reptilia" family="Thescelosauridae" genus="Thescelosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91A356C3813FFEFFCDAF8C99759AA18" box="[878,1052,1883,1908]" italics="true" pageId="4" pageNumber="251">Thescelosaurus</emphasis>
|
||
</taxonomicName>
|
||
. Notably, tree length increased in seven, six and again six steps when placing
|
||
<taxonomicName id="4C6E92FD3813FFEFFD69F8139024AAF4" authority="Párraga & Prieto-Márquez, 2019" authorityName="Párraga & Prieto-Márquez" authorityYear="2019" box="[733,865,1921,1944]" class="Reptilia" family="Rhabdodontidae" genus="Pareisactus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="249" phylum="Chordata" rank="genus" status="gen. nov.">
|
||
<emphasis id="B91A356C3813FFEFFD69F8139024AAF4" box="[733,865,1921,1944]" italics="true" pageId="4" pageNumber="251">Pareisactus</emphasis>
|
||
</taxonomicName>
|
||
within Hadrosauriformes, Hadrosauroidea and
|
||
<taxonomicName id="4C6E92FD3813FFEFFF23F8319278AAD0" box="[151,317,1955,1980]" class="Reptilia" family="Hadrosauridae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="251" phylum="Chordata" rank="family">Hadrosauridae</taxonomicName>
|
||
, respectively.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |