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<document id="92CEF81D027C9A60391995A40603024B" ID-DOI="10.5070/P9371050859" ID-ISSN="0031-0298" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1726019095505" checkinUser="felipe" docAuthor="Marsh, Adam D. &amp; Parker, William G." docDate="2022" docId="038C8F41F820096E62B1D5F1FAF58F37" docLanguage="en" docName="PaleoBios.37.50859.pdf" docOrigin="PaleoBios 37" docStyle="DocumentStyle:2D14AFDE223D5FD9EB1703D6A63DEF6C.1:PaleoBios.2018-2021.journal_article" docStyleId="2D14AFDE223D5FD9EB1703D6A63DEF6C" docStyleName="PaleoBios.2018-2021.journal_article" docStyleVersion="1" docTitle="Archosauria COPE, 1869 SENSU NESBITT, 2011 AVEMETATARSALIA BENTON 1999" docType="treatment" docVersion="1" lastPageNumber="9" masterDocId="FFB5F739F82A09646206D72CFFC88C26" masterDocTitle="New dinosauromorph specimens from Petrified Forest National Park and a global biostratigraphic review of Triassic dinosauromorph body fossils" masterLastPageNumber="56" masterPageNumber="1" pageNumber="9" updateTime="1726078387551" updateUser="felipe" zenodo-license-document="CC-BY-NC-SA-4.0">
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<mods:title id="AB8FC64A44756366C90AE1C8638A97F1">New dinosauromorph specimens from Petrified Forest National Park and a global biostratigraphic review of Triassic dinosauromorph body fossils</mods:title>
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<mods:namePart id="3CBD357B108781580518620B0844FA88">Marsh, Adam D.</mods:namePart>
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<mods:namePart id="827A592C4CD4ADE9D4E8670D2A3F6D76">Parker, William G.</mods:namePart>
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<mods:title id="B28D723B170B5CE7C8267BF012BBE219">PaleoBios</mods:title>
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<mods:date id="9F40C66B0A4B051176F62480822F3C94">2022</mods:date>
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<mods:number id="F08B8D23C83D10EA354795C8E1D7C7B4">37</mods:number>
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<mods:identifier id="136F621DBC8E3BCF0C3486EE45BC72AC" type="ISSN">0031-0298</mods:identifier>
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<treatment id="038C8F41F820096E62B1D5F1FAF58F37" LSID="urn:lsid:plazi:treatment:038C8F41F820096E62B1D5F1FAF58F37" httpUri="http://treatment.plazi.org/id/038C8F41F820096E62B1D5F1FAF58F37" lastPageNumber="9" pageId="10" pageNumber="9">
<subSubSection id="C33F6DDCF820096E62B1D5F1FCD18F73" pageId="10" pageNumber="9" type="nomenclature">
<paragraph id="8B9A3E57F820096E62B1D5F1FCC38F33" blockId="10.[119,793,733,853]" pageId="10" pageNumber="9">
<heading id="D0D2893BF820096E62B1D5F1FCC38F33" allCaps="true" centered="true" fontSize="10" level="2" pageId="10" pageNumber="9" reason="2">
<taxonomicName id="4C2545D4F820096E62B1D5F1FCC08F33" authority="COPE, 1869 SENSU NESBITT, 2011 AVEMETATARSALIA BENTON, 1999 SENSU NESBITT ET AL." authorityName="COPE, 1869 SENSU NESBITT, 2011 AVEMETATARSALIA BENTON" authorityYear="1999" class="Archosauria" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="9" phylum="Chordata" rank="class">
ARCHOSAURIA COPE, 1869
<emphasis id="B951E245F820096E63E8D5F1FDF88ED2" box="[494,560,733,756]" italics="true" pageId="10" pageNumber="9">SENSU</emphasis>
NESBITT, 2011 AVEMETATARSALIA BENTON, 1999
<emphasis id="B951E245F820096E6010D5D1FD908F32" box="[534,600,765,788]" italics="true" pageId="10" pageNumber="9">SENSU</emphasis>
NESBITT ET AL.
</taxonomicName>
,
</heading>
</paragraph>
<paragraph id="8B9A3E57F820096E63ACD431FCD18F73" blockId="10.[119,793,733,853]" pageId="10" pageNumber="9">
2017 ORNITHODIRA GAUTHIER, 1986
<emphasis id="B951E245F820096E63E2D411FDEE8F72" box="[484,550,829,852]" italics="true" pageId="10" pageNumber="9">SENSU</emphasis>
NESBITT ET AL., 2017
</paragraph>
</subSubSection>
<subSubSection id="C33F6DDCF820096E6390D44BFE318F59" box="[406,505,871,895]" pageId="10" pageNumber="9" type="description">
<paragraph id="8B9A3E57F820096E6390D44BFE318F59" blockId="10.[406,505,871,895]" box="[406,505,871,895]" pageId="10" pageNumber="9">
<figureCitation id="131E22D2F820096E6390D44BFE318F59" box="[406,505,871,895]" captionStart="Figure 4" captionStartId="11.[112,183,1791,1813]" captionTargetBox="[769,1416,205,1767]" captionTargetId="figure-230@11.[808,1105,863,1764]" captionTargetPageId="11" captionText="Figure 4. A–E. Left ornithodiran metatarsal I, PEFO 34741 in lateral (A), dorsal (B), medial (C), proximal (D), and distal (E) view. F–J. Right ornithodiran metatarsal I, PEFO 44217 in lateral (F), dorsal (G), medial (H), proximal (I), and distal (J) view. K–O. Right dinosauromorph tibia, PEFO 44476 in medial (K), medial (L), proximal (M), posterior (N), and distal (O) view.P. Right dinosauriform scapula, PEFO 26678 in lateral view.Q. Right dinosauriform scapulocoracoid, PEFO 46207 in lateral view. Abbreviations: cc, cnemial crest; cf, coracoid foramen; co, coracoid; cs, cross-section; de, distal end; gl, glenoid; lc, lateral condyle; lp, lip; mc, medial condyle; pe, proximal end; r, ridge; sc, scapula. Scale bars=1 cm, arrows point in anterior direction (note that PEFO 44476 is broken in two pieces within the block, that those are perpendicular to one another in the block, and the written arrow on the matrix in L is a preparation mark and does not indicate direction)." pageId="10" pageNumber="9">FIG. 4A–J</figureCitation>
</paragraph>
</subSubSection>
<subSubSection id="C33F6DDCF820096E6296D4BFFE4B883E" pageId="10" pageNumber="9" type="materials_examined">
<paragraph id="8B9A3E57F820096E6296D4BFFE4B883E" blockId="10.[112,801,915,1950]" pageId="10" pageNumber="9">
<emphasis id="B951E245F820096E6296D4BFFDAB8F8B" bold="true" box="[144,611,915,942]" pageId="10" pageNumber="9">Referred specimens and localities</emphasis>
—
<collectionCode id="ED34A692F820096E6079D4B8FD0C8F88" box="[639,708,916,942]" country="0" httpUri="http://grbio.org/cool/7x98-r3em" name="Petrified Forest" pageId="10" pageNumber="9">PEFO</collectionCode>
36741 (
<figureCitation id="131E22D2F820096E627FD49BFF3B8FF7" box="[121,243,951,977]" captionStart="Figure 4" captionStartId="11.[112,183,1791,1813]" captionTargetBox="[769,1416,205,1767]" captionTargetId="figure-230@11.[808,1105,863,1764]" captionTargetPageId="11" captionText="Figure 4. A–E. Left ornithodiran metatarsal I, PEFO 34741 in lateral (A), dorsal (B), medial (C), proximal (D), and distal (E) view. F–J. Right ornithodiran metatarsal I, PEFO 44217 in lateral (F), dorsal (G), medial (H), proximal (I), and distal (J) view. K–O. Right dinosauromorph tibia, PEFO 44476 in medial (K), medial (L), proximal (M), posterior (N), and distal (O) view.P. Right dinosauriform scapula, PEFO 26678 in lateral view.Q. Right dinosauriform scapulocoracoid, PEFO 46207 in lateral view. Abbreviations: cc, cnemial crest; cf, coracoid foramen; co, coracoid; cs, cross-section; de, distal end; gl, glenoid; lc, lateral condyle; lp, lip; mc, medial condyle; pe, proximal end; r, ridge; sc, scapula. Scale bars=1 cm, arrows point in anterior direction (note that PEFO 44476 is broken in two pieces within the block, that those are perpendicular to one another in the block, and the written arrow on the matrix in L is a preparation mark and does not indicate direction)." pageId="10" pageNumber="9">Fig. 4A–E</figureCitation>
), left metatarsal I,
<collectionCode id="ED34A692F820096E63D5D49BFDCD8FF7" box="[467,517,951,977]" pageId="10" pageNumber="9">PFV</collectionCode>
215: Zuni Well Mound,
<collectionCode id="ED34A692F820096E6276D4F6FF638FD2" box="[112,171,986,1012]" pageId="10" pageNumber="9">PFM</collectionCode>
;
<collectionCode id="ED34A692F820096E62B2D4F6FF3F8FD2" box="[180,247,986,1012]" country="0" httpUri="http://grbio.org/cool/7x98-r3em" name="Petrified Forest" pageId="10" pageNumber="9">PEFO</collectionCode>
44217 (
<figureCitation id="131E22D2F820096E635FD4F6FE0D8FD2" box="[345,453,986,1012]" captionStart="Figure 4" captionStartId="11.[112,183,1791,1813]" captionTargetBox="[769,1416,205,1767]" captionTargetId="figure-230@11.[808,1105,863,1764]" captionTargetPageId="11" captionText="Figure 4. A–E. Left ornithodiran metatarsal I, PEFO 34741 in lateral (A), dorsal (B), medial (C), proximal (D), and distal (E) view. F–J. Right ornithodiran metatarsal I, PEFO 44217 in lateral (F), dorsal (G), medial (H), proximal (I), and distal (J) view. K–O. Right dinosauromorph tibia, PEFO 44476 in medial (K), medial (L), proximal (M), posterior (N), and distal (O) view.P. Right dinosauriform scapula, PEFO 26678 in lateral view.Q. Right dinosauriform scapulocoracoid, PEFO 46207 in lateral view. Abbreviations: cc, cnemial crest; cf, coracoid foramen; co, coracoid; cs, cross-section; de, distal end; gl, glenoid; lc, lateral condyle; lp, lip; mc, medial condyle; pe, proximal end; r, ridge; sc, scapula. Scale bars=1 cm, arrows point in anterior direction (note that PEFO 44476 is broken in two pieces within the block, that those are perpendicular to one another in the block, and the written arrow on the matrix in L is a preparation mark and does not indicate direction)." pageId="10" pageNumber="9">Fig. 4F–J</figureCitation>
), right metatarsal I,
<collectionCode id="ED34A692F820096E60B4D4F6FD2C8FD2" box="[690,740,986,1012]" pageId="10" pageNumber="9">PFV</collectionCode>
217: Dinosaur Wash,
<collectionCode id="ED34A692F820096E633DD4D2FE48883E" box="[315,384,1022,1048]" country="Germany" httpUri="http://grbio.org/cool/e0n2-eyvk" name="Buergermeister Mueller, Museum" pageId="10" pageNumber="9">BMM</collectionCode>
.
</paragraph>
</subSubSection>
<subSubSection id="C33F6DDCF820096E6296D333FAF58F37" pageId="10" pageNumber="9" type="description">
<paragraph id="8B9A3E57F820096E6296D333FAF58F37" blockId="10.[112,801,915,1950]" lastBlockId="10.[832,1520,195,785]" pageId="10" pageNumber="9">
<emphasis id="B951E245F820096E6296D333FD54881F" bold="true" box="[144,668,1055,1082]" pageId="10" pageNumber="9">Description and rationale for assignment</emphasis>
—Metatarsals I of PEFO 36741 and PEFO 44217 are proportionally long and slender metapodials and, as observed by the crushing of PEFO 34741, hollow. The proximal end of PEFO 34371 is twisted with respect to the distal end. In proximal view, the medial surface is nearly straight where it would articulate with metatarsal II, and the lateral edge is rounded, making a subtriangular proximal outline (
<figureCitation id="131E22D2F820096E60F4D23EFF678969" captionStart="Figure 4" captionStartId="11.[112,183,1791,1813]" captionTargetBox="[769,1416,205,1767]" captionTargetId="figure-230@11.[808,1105,863,1764]" captionTargetPageId="11" captionText="Figure 4. A–E. Left ornithodiran metatarsal I, PEFO 34741 in lateral (A), dorsal (B), medial (C), proximal (D), and distal (E) view. F–J. Right ornithodiran metatarsal I, PEFO 44217 in lateral (F), dorsal (G), medial (H), proximal (I), and distal (J) view. K–O. Right dinosauromorph tibia, PEFO 44476 in medial (K), medial (L), proximal (M), posterior (N), and distal (O) view.P. Right dinosauriform scapula, PEFO 26678 in lateral view.Q. Right dinosauriform scapulocoracoid, PEFO 46207 in lateral view. Abbreviations: cc, cnemial crest; cf, coracoid foramen; co, coracoid; cs, cross-section; de, distal end; gl, glenoid; lc, lateral condyle; lp, lip; mc, medial condyle; pe, proximal end; r, ridge; sc, scapula. Scale bars=1 cm, arrows point in anterior direction (note that PEFO 44476 is broken in two pieces within the block, that those are perpendicular to one another in the block, and the written arrow on the matrix in L is a preparation mark and does not indicate direction)." pageId="10" pageNumber="9">Fig. 4D, I</figureCitation>
). The proximal outline is mediolaterally narrow, which indicates that the metatarsus was compact, not splayed (Ezcurra 2016:565-1). A compact metatarsus is present in ornithodirans and some early crocodylomorphs (Nesbitt 2011, Ezcurra 2016). The proximal surface of PEFO 36741 and PEFO 44217 is flat, would articulate with the distal tarsals, and would reach the proximal extent of metatarsal II, unlike in neotheropods where metatarsal I is more distally placed along the back of the metatarsus (Rauhut 2003, Nesbitt 2011:385-1). The distal end is asymmetrical, such that the medial condyle is more prominent than the lateral (
<figureCitation id="131E22D2F820096E60A6D19EFCDB8AEB" box="[672,787,1714,1741]" captionStart="Figure 4" captionStartId="11.[112,183,1791,1813]" captionTargetBox="[769,1416,205,1767]" captionTargetId="figure-230@11.[808,1105,863,1764]" captionTargetPageId="11" captionText="Figure 4. A–E. Left ornithodiran metatarsal I, PEFO 34741 in lateral (A), dorsal (B), medial (C), proximal (D), and distal (E) view. F–J. Right ornithodiran metatarsal I, PEFO 44217 in lateral (F), dorsal (G), medial (H), proximal (I), and distal (J) view. K–O. Right dinosauromorph tibia, PEFO 44476 in medial (K), medial (L), proximal (M), posterior (N), and distal (O) view.P. Right dinosauriform scapula, PEFO 26678 in lateral view.Q. Right dinosauriform scapulocoracoid, PEFO 46207 in lateral view. Abbreviations: cc, cnemial crest; cf, coracoid foramen; co, coracoid; cs, cross-section; de, distal end; gl, glenoid; lc, lateral condyle; lp, lip; mc, medial condyle; pe, proximal end; r, ridge; sc, scapula. Scale bars=1 cm, arrows point in anterior direction (note that PEFO 44476 is broken in two pieces within the block, that those are perpendicular to one another in the block, and the written arrow on the matrix in L is a preparation mark and does not indicate direction)." pageId="10" pageNumber="9">Fig. 4E, J</figureCitation>
). Both distal condyles have a collateral ligament fossa. In dorsal view, the medial condyle projects distally further than the lateral. The distal condyles of metatarsal I in the crocodylomorphs
<taxonomicName id="4C2545D4F820096E6388D010FF0F8B5C" authority="Brown (1933)" authorityName="Brown" authorityYear="1933" class="Reptilia" family="Protosuchidae" genus="Protosuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Crocodylia" pageId="10" pageNumber="9" phylum="Chordata" rank="species" species="richardsoni">
<emphasis id="B951E245F820096E6388D010FD088B7E" box="[398,704,1852,1880]" italics="true" pageId="10" pageNumber="9">Protosuchus richardsoni</emphasis>
Brown (1933)
</taxonomicName>
(AMNH FR 3024), Colbert and Mook 1951: fig. 20b) and
<taxonomicName id="4C2545D4F820096E62EAD0ADFD748BBA" authority="Crush (1984)" authorityName="Crush" authorityYear="1984" box="[236,700,1921,1950]" class="Reptilia" family="Saltoposuchidae" genus="Terrestrisuchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Prolacertiformes" pageId="10" pageNumber="9" phylum="Chordata" rank="species" species="gracilis">
<emphasis id="B951E245F820096E62EAD0ADFDC38BB8" box="[236,523,1921,1950]" italics="true" pageId="10" pageNumber="9">Terrestrisuchus gracilis</emphasis>
Crush (1984)
</taxonomicName>
are not asymmetrical (Crush 1984: fig. 10e). Asymmetrical distal condyles of metatarsal I are found in
<taxonomicName id="4C2545D4F820096E6703D7C9FB4E8D05" authority="Crompton and Charig" authorityName="Crompton and Charig" class="Reptilia" family="Heterodontosauridae" genus="Heterodontosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="10" pageNumber="9" phylum="Chordata" rank="species" species="tucki">
<emphasis id="B951E245F820096E6703D7C9FCB48D02" italics="true" pageId="10" pageNumber="9">Heterodontosaurus tucki</emphasis>
Crompton and Charig
</taxonomicName>
(1962, SAM-PK-K1332) (Santa Lucas 1980: fig. 20b),
<taxonomicName id="4C2545D4F820096E6670D607FA388D61" authority="Sereno et al." authorityName="Sereno" box="[1142,1520,299,327]" class="Reptilia" genus="Eoraptor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="10" pageNumber="9" phylum="Chordata" rank="species" species="lunensis">
<emphasis id="B951E245F820096E6670D607FA868D61" box="[1142,1358,299,327]" italics="true" pageId="10" pageNumber="9">Eoraptor lunensis</emphasis>
Sereno et al.
</taxonomicName>
(1993, PVSJ 512) (Sereno et al. 2012: fig. 91b),
<taxonomicName id="4C2545D4F820096E6776D67CFBA38DAB" authority="Langer et al." authorityName="Langer" class="Reptilia" genus="Saturnalia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="10" pageNumber="9" phylum="Chordata" rank="species" species="tupiniquim">
<emphasis id="B951E245F820096E6776D67CFC0F8DA8" italics="true" pageId="10" pageNumber="9">Saturnalia tupiniquim</emphasis>
Langer et al.
</taxonomicName>
(1999, MCP PV 3844) (Langer 2003: fig. 7e),
<taxonomicName id="4C2545D4F820096E660FD6B9FA278D96" authority="Reig" authorityName="Reig" authorityYear="1963" box="[1033,1519,405,435]" class="Reptilia" family="Herrerasauridae" genus="Herrerasaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="10" pageNumber="9" phylum="Chordata" rank="species" species="ischigualastensis">
<emphasis id="B951E245F820096E660FD6B9FA638D95" box="[1033,1451,405,435]" italics="true" pageId="10" pageNumber="9">Herrerasaurus ischigualastensis</emphasis>
Reig
</taxonomicName>
(1963, PVSJ 373) (Novas 1993: fig. 9j) and
<taxonomicName id="4C2545D4F820096E6762D696FC4A8DD1" authorityName="Cope" authorityYear="1887" class="Reptilia" family="Coelophysidae" genus="Coelophysis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Saurischia" pageId="10" pageNumber="9" phylum="Chordata" rank="species" species="bauri">
<emphasis id="B951E245F820096E6762D696FC4A8DD1" italics="true" pageId="10" pageNumber="9">Coelophysis bauri</emphasis>
</taxonomicName>
(MNA V3320, Nesbitt 2011: fig. 48f), but not in the non-dinosaur dinosauriform
<taxonomicName id="4C2545D4F820096E66AED6D2FCB38E1B" authority="Nesbitt et al." authorityName="Nesbitt" family="Silesauridae" genus="Asilisaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="10" pageNumber="9" phylum="Chordata" rank="species" species="kongwe">
<emphasis id="B951E245F820096E66AED6D2FA598E3A" box="[1192,1425,510,540]" italics="true" pageId="10" pageNumber="9">Asilisaurus kongwe</emphasis>
Nesbitt et al.
</taxonomicName>
(2010, NMT RB159) (Nesbitt et al. 2019: fig. 53e). The PEFO specimens are identical to metatarsal I of GR 1033 (Nesbitt et al. 2009a: fig. 2j), a partial skeleton of
<emphasis id="B951E245F820096E6146D5A7FC4A8E81" box="[832,898,651,679]" italics="true" pageId="10" pageNumber="9">Tawa</emphasis>
<taxonomicName id="4C2545D4F820096E6180D5A7FB1A8E81" authority="Nesbitt et al. (2009 a)" authorityName="Nesbitt" authorityYear="2009" box="[902,1234,651,679]" class="Reptilia" family="Stagonolepididae" genus="Typothorax" higherTaxonomySource="GBIF" kingdom="Animalia" order="Aetosauria" pageId="10" pageNumber="9" phylum="Chordata" rank="species" species="hallae">
<emphasis id="B951E245F820096E6180D5A7FC188E81" box="[902,976,651,679]" italics="true" pageId="10" pageNumber="9">hallae</emphasis>
Nesbitt et al. (2009a)
</taxonomicName>
. We assign PEFO 36741 and PEFO 44217 to non-neotheropod Ornithodira owing to the presence of a compact metatarsal I that is not distally positioned down the metatarsus.
</paragraph>
</subSubSection>
</treatment>
</document>
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