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<document id="34A4EA86E7DA5D7DDAAB6665AC8A93F6" ID-DOI="10.1206/0003-0082(2003)395&lt;0001:ANSOPG&gt;2.0.CO;2" ID-ISSN="0003-0082" ID-Zenodo-Dep="5373383" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630273424663" checkinUser="carolina" docAuthor="HILL, ROBERT V., WITMER, LAWRENCE M. &amp; NORELL, MARK A." docDate="2003" docId="03C587BFE91FFFF8FF2B03C7FDDB5D17" docLanguage="en" docName="N3395.pdf" docOrigin="American Museum Novitates 3395" docSource="http://www.bioone.org/doi/abs/10.1206/0003-0082%282003%29395%3C0001%3AANSOPG%3E2.0.CO%3B2" docStyle="DocumentStyle:70DDF1BDF6846024993F05262BAE3001.3:AmMusNovit.2000-2010.journal_article.type1" docStyleId="70DDF1BDF6846024993F05262BAE3001" docStyleName="AmMusNovit.2000-2010.journal_article.type1" docStyleVersion="3" docTitle="Syrmosaurus viminicaudus Maleev 1952" docType="treatment" docVersion="3" lastPageNumber="4" masterDocId="FFFCFFC7E91CFFFCFFA1061DFF9D586A" masterDocTitle="A New Specimen of Pinacosaurus grangeri (Dinosauria: Ornithischia) from the Late Cretaceous of Mongolia: Ontogeny and Phylogeny of Ankylosaurs" masterLastPageNumber="30" masterPageNumber="1" pageNumber="4" updateTime="1699208312011" updateUser="plazi" zenodo-license-document="CC-BY-4.0">
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<mods:title id="948E424E8B21B069949C4A1B71640E74">A New Specimen of Pinacosaurus grangeri (Dinosauria: Ornithischia) from the Late Cretaceous of Mongolia: Ontogeny and Phylogeny of Ankylosaurs</mods:title>
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<mods:namePart id="4050D7CBEEA6C09F5E6FE0594E5245E3">HILL, ROBERT V.</mods:namePart>
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<mods:namePart id="508D241E3C0129B504F52537C5D65116">WITMER, LAWRENCE M.</mods:namePart>
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<mods:namePart id="8EB1A501F087DE9FBBAA40AEDD4E1058">NORELL, MARK A.</mods:namePart>
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<mods:title id="81798211D7FBD6C2B71249F8EC039F5D">American Museum Novitates</mods:title>
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<mods:date id="683087EC36D608BEF5621FA63BB0B36A">2003</mods:date>
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<mods:number id="A9EC1F3FD31A92BFFF584F2400552432">2003-02-19</mods:number>
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<mods:detail id="CE7D8784CCFDE5E7685C96C525BD9DBB" type="volume">
<mods:number id="A3DB5407E2FB76294AAA2AACED55CFBC">3395</mods:number>
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<mods:url id="05A0A2AB816ABD5143253C7E9763826B">http://www.bioone.org/doi/abs/10.1206/0003-0082%282003%29395%3C0001%3AANSOPG%3E2.0.CO%3B2</mods:url>
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<mods:identifier id="8E9320B9199B8734FD3CDB2538E7B6F9" type="DOI">10.1206/0003-0082(2003)395&lt;0001:ANSOPG&gt;2.0.CO;2</mods:identifier>
<mods:identifier id="07A5B7DFED0864EAD29AACD151E241C6" type="ISSN">0003-0082</mods:identifier>
<mods:identifier id="B4B9DED2A105944A9E2324A605C945C8" type="Zenodo-Dep">5373383</mods:identifier>
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<treatment id="03C587BFE91FFFF8FF2B03C7FDDB5D17" LSID="urn:lsid:plazi:treatment:03C587BFE91FFFF8FF2B03C7FDDB5D17" httpUri="http://treatment.plazi.org/id/03C587BFE91FFFF8FF2B03C7FDDB5D17" lastPageId="4" lastPageNumber="4" pageId="3" pageNumber="4">
<subSubSection id="C3766522E91FFFFFFF2B03C7FDE65D9B" box="[138,635,1498,1521]" pageId="3" pageNumber="4" type="nomenclature">
<paragraph id="8BD336A9E91FFFFFFF2B03C7FDE65D9B" blockId="3.[138,635,1498,1521]" box="[138,635,1498,1521]" pageId="3" pageNumber="4">
<heading id="D09B81C5E91FFFFFFF2B03C7FDE65D9B" box="[138,635,1498,1521]" centered="true" fontSize="36" level="2" pageId="3" pageNumber="4" reason="8">
<taxonomicName id="4C6C4D2AE91FFFFFFF2B03C7FDEE5D9B" authority="Maleev, 1952" authorityName="Maleev" authorityYear="1952" box="[138,627,1498,1521]" class="Reptilia" family="Ankylosauridae" genus="Syrmosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="viminicaudus">
<emphasis id="B918EABBE91FFFFFFF2B03C7FE5A5D9B" box="[138,455,1498,1521]" italics="true" pageId="3" pageNumber="4">Syrmosaurus viminicaudus</emphasis>
<bibRefCitation id="EFFD4B58E91FFFFFFE7103C7FDEE5D9B" author="Maleev, E. A." box="[464,627,1498,1521]" pageId="3" pageNumber="4" pagination="131 - 134" refId="ref12520" refString="Maleev, E. A. 1952. A new family of armored dinosaurs from the Upper Cretaceous of Mongolia. Doklady Akademii Nauk SSSR 87: 131 - 134. [in Russian]" type="journal article" year="1952">Maleev, 1952</bibRefCitation>
</taxonomicName>
)
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C3766522E91FFFFFFF39001BFC5F5927" pageId="3" pageNumber="4" type="materials_examined">
<paragraph id="8BD336A9E91FFFFFFF39001BFDBF5E32" blockId="3.[123,651,1542,1741]" pageId="3" pageNumber="4">
<materialsCitation id="3B043CF4E91FFFFFFF39001BFDBF5E32" collectionCode="AMNH" pageId="3" pageNumber="4" specimenCode="AMNH 6523" specimenCount="1" typeStatus="holotype">
<typeStatus id="54D7880BE91FFFFFFF39001BFE935E77" box="[152,270,1542,1565]" pageId="3" pageNumber="4" type="holotype">HOLOTYPE</typeStatus>
:
<specimenCode id="DBCA9ED2E91FFFFFFE86001BFE535E77" box="[295,462,1542,1565]" collectionCode="AMNH" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34925" lsid="urn:lsid:biocol.org:col:34925" name="American Museum of Natural History" pageId="3" pageNumber="4">AMNH 6523</specimenCode>
, dorsoventrally crushed skull and mandible with several associated osteoderms, atlas and axis.
</materialsCitation>
</paragraph>
<paragraph id="8BD336A9E91FFFFFFF390043FE925EDA" blockId="3.[123,651,1542,1741]" pageId="3" pageNumber="4">
TYPE LOCALITY: Djadokhta Formation, Shabarakh Usu,
<collectingCountry id="F37B7639E91FFFFFFE9E0061FE2D5EF9" box="[319,432,1660,1683]" name="Mongolia" pageId="3" pageNumber="4">Mongolia</collectingCountry>
(e.g., ‘‘The Flaming Cliffs’’).
</paragraph>
<paragraph id="8BD336A9E91FFFFFFF3900ABFD165EA7" blockId="3.[123,651,1542,1741]" box="[152,651,1718,1741]" pageId="3" pageNumber="4">
REFERRED SPECIMENS: Uncataloged
<collectionCode id="ED7DAE6CE91FFFFFFDED00ABFD165EA7" box="[588,651,1718,1741]" country="China" httpUri="http://grbio.org/cool/wd8c-kdma" name="Institute of Vertebrate Paleontology and Paleoanthropology" pageId="3" pageNumber="4">IVPP</collectionCode>
</paragraph>
<paragraph id="8BD336A9E91FFFFFFD1A06DCFC5F5927" blockId="3.[699,1228,193,1741]" pageId="3" pageNumber="4">
specimens;
<collectionCode id="ED7DAE6CE91FFFFFFCE606DCFC1D58B2" box="[839,896,193,216]" country="Mongolia" httpUri="http://grbio.org/cool/0h5a-qjh4" name="Geological Institute, Mongolian Academy of Sciences" pageId="3" pageNumber="4">IGM</collectionCode>
100/1014;
<collectionCode id="ED7DAE6CE91FFFFFFBAA06DCFBA758B2" box="[1035,1082,193,216]" country="Russia" httpUri="http://grbio.org/cool/kjjc-ziy5" name="Paleontological Institute, Russian Academy of Sciences" pageId="3" pageNumber="4">PIN</collectionCode>
614, uncataloged
<collectionCode id="ED7DAE6CE91FFFFFFCBC06C3FCD1589F" box="[797,844,222,245]" country="Russia" httpUri="http://grbio.org/cool/kjjc-ziy5" name="Paleontological Institute, Russian Academy of Sciences" pageId="3" pageNumber="4">PIN</collectionCode>
specimens;
<collectionCode id="ED7DAE6CE91FFFFFFC5606C3FBA3589F" box="[1015,1086,222,245]" country="Poland" httpUri="http://grbio.org/cool/yd4q-n37a" name="Zoological Institute of Paleobiology, Polish Academy of Sciences" pageId="3" pageNumber="4">ZPAL</collectionCode>
MgD­II/1,
<collectionCode id="ED7DAE6CE91FFFFFFD1A06E1FC9F5979" box="[699,770,252,275]" country="Poland" httpUri="http://grbio.org/cool/yd4q-n37a" name="Zoological Institute of Paleobiology, Polish Academy of Sciences" pageId="3" pageNumber="4">ZPAL</collectionCode>
MgD­II/2,
<collectionCode id="ED7DAE6CE91FFFFFFC3E06E1FC7B5979" box="[927,998,252,275]" country="Poland" httpUri="http://grbio.org/cool/yd4q-n37a" name="Zoological Institute of Paleobiology, Polish Academy of Sciences" pageId="3" pageNumber="4">ZPAL</collectionCode>
MgD­II/7,
<collectionCode id="ED7DAE6CE91FFFFFFB2506E1FB565979" box="[1156,1227,252,275]" country="Poland" httpUri="http://grbio.org/cool/yd4q-n37a" name="Zoological Institute of Paleobiology, Polish Academy of Sciences" pageId="3" pageNumber="4">ZPAL</collectionCode>
MgD­II/9,
<collectionCode id="ED7DAE6CE91FFFFFFC9A0704FC1F595A" box="[827,898,281,304]" country="Poland" httpUri="http://grbio.org/cool/yd4q-n37a" name="Zoological Institute of Paleobiology, Polish Academy of Sciences" pageId="3" pageNumber="4">ZPAL</collectionCode>
MgD­II/27,
<collectionCode id="ED7DAE6CE91FFFFFFBB80704FBFD595A" box="[1049,1120,281,304]" country="Poland" httpUri="http://grbio.org/cool/yd4q-n37a" name="Zoological Institute of Paleobiology, Polish Academy of Sciences" pageId="3" pageNumber="4">ZPAL</collectionCode>
MgD­II/ 31,
<collectionCode id="ED7DAE6CE91FFFFFFD48072BFCAD5927" box="[745,816,310,333]" country="Poland" httpUri="http://grbio.org/cool/yd4q-n37a" name="Zoological Institute of Paleobiology, Polish Academy of Sciences" pageId="3" pageNumber="4">ZPAL</collectionCode>
MgD­II/32.
</paragraph>
</subSubSection>
<subSubSection id="C3766522E91FFFFFFD790749FBD759A9" pageId="3" pageNumber="4" type="diagnosis">
<paragraph id="8BD336A9E91FFFFFFD790749FBD759A9" blockId="3.[699,1228,193,1741]" pageId="3" pageNumber="4">
DIAGNOSIS: As for genus;
<taxonomicName id="4C6C4D2AE91FFFFFFB8A0749FB565901" authorityName="Gilmore" authorityYear="1933" box="[1067,1227,340,363]" class="Reptilia" family="Ankylosauridae" genus="Pinacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B918EABBE91FFFFFFB8A0749FB565901" box="[1067,1227,340,363]" italics="true" pageId="3" pageNumber="4">Pinacosaurus</emphasis>
</taxonomicName>
with squamosal dermal ossifications present as weakly developed pyramids, not long spines as in
<taxonomicName id="4C6C4D2AE91FFFFFFCF107B1FBDE59A9" box="[848,1091,428,451]" class="Reptilia" family="Ankylosauridae" genus="Pinacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="mephistocephalus">
<emphasis id="B918EABBE91FFFFFFCF107B1FBDE59A9" box="[848,1091,428,451]" italics="true" pageId="3" pageNumber="4">P. mephistocephalus</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="C3766522E91FFFF8FD7907D4FDDB5D17" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4" type="discussion">
<paragraph id="8BD336A9E91FFFFFFD7907D4FD6B5D7F" blockId="3.[699,1228,193,1741]" pageId="3" pageNumber="4">
DISCUSSION: Abundant material collected by the Sino­Swedish, Polish­Mongolian, Sino­Canadian, and Russian­Mongolian expeditions has been referred to
<taxonomicName id="4C6C4D2AE91FFFFFFB82043CFB375A52" authorityName="Gilmore" authorityYear="1933" box="[1059,1194,545,568]" class="Reptilia" family="Ankylosauridae" genus="Pinacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="grangeri">
<emphasis id="B918EABBE91FFFFFFB82043CFB375A52" box="[1059,1194,545,568]" italics="true" pageId="3" pageNumber="4">P. grangeri</emphasis>
</taxonomicName>
or
<taxonomicName id="4C6C4D2AE91FFFFFFD1A0422FC1E5A3F" box="[699,899,574,598]" class="Reptilia" family="Ankylosauridae" genus="Pinacosaurus" kingdom="Animalia" order="Ornithischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B918EABBE91FFFFFFD1A0422FCC65A3C" box="[699,859,575,598]" italics="true" pageId="3" pageNumber="4">Pinacosaurus</emphasis>
sp.
</taxonomicName>
Most of the cranial material collected by these teams has represented juvenile or subadult individuals. All specimens of
<taxonomicName id="4C6C4D2AE91FFFFFFD7D048AFCE15AC4" authorityName="Gilmore" authorityYear="1933" box="[732,892,663,686]" class="Reptilia" family="Ankylosauridae" genus="Pinacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B918EABBE91FFFFFFD7D048AFCE15AC4" box="[732,892,663,686]" italics="true" pageId="3" pageNumber="4">Pinacosaurus</emphasis>
</taxonomicName>
feature the distinctive set of characters found in the
<typeStatus id="54D7880BE91FFFFFFC7304A9FBA65AA1" box="[978,1083,692,715]" pageId="3" pageNumber="4" type="holotype">holotype</typeStatus>
, including a highly specialized narial region with at least three pairs of openings, prominent hornlike protuberances above the orbits, and lack of secondary dermal ossifications on the external rostroventral portion of the premaxillary beak. The
<typeStatus id="54D7880BE91FFFFFFC9A0579FC3C5B11" box="[827,929,868,891]" pageId="3" pageNumber="4" type="holotype">holotype</typeStatus>
skull, however, also differs considerably from the numerous subadult crania that have been referred to the genus. The skull is longer than wide, with a narrow premaxillary beak and palate.
<bibRefCitation id="EFFD4B58E91FFFFFFB3305C4FCFD5C67" author="Maryanska, T." pageId="3" pageNumber="4" pagination="85 - 151" refId="ref12581" refString="Maryanska, T. 1977. Ankylosauridae (Dinosauria) from Mongolia. Palaeontologia Polonica 37: 85 - 151." type="journal article" year="1977">Maryańska (1977)</bibRefCitation>
has attributed this condition to taphonomic processes, but the relatively mild distortion of the pterygoids and premaxillae suggests that the postdepositional reworking of this specimen was limited to dorsoventral crushing. Nevertheless, skulls subsequently referred to
<taxonomicName id="4C6C4D2AE91FFFFFFCE802BAFC485CD4" authorityName="Gilmore" authorityYear="1933" box="[841,981,1191,1214]" class="Reptilia" family="Ankylosauridae" genus="Pinacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="grangeri">
<emphasis id="B918EABBE91FFFFFFCE802BAFC485CD4" box="[841,981,1191,1214]" italics="true" pageId="3" pageNumber="4">P. grangeri</emphasis>
</taxonomicName>
consistently exhibit the typical ankylosaurid condition of a skull and premaxillary palate that is wider than long.
</paragraph>
<paragraph id="8BD336A9E91FFFF8FD790301FDDB5D17" blockId="3.[699,1228,193,1741]" lastBlockId="4.[124,653,1236,1405]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
A narrow skull can be found in more basal thyreophorans such as
<taxonomicName id="4C6C4D2AE91FFFFFFC410324FB595D3A" authorityName="Haubold" authorityYear="1990" box="[992,1220,1337,1360]" class="Reptilia" genus="Emausaurus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="ernsti">
<emphasis id="B918EABBE91FFFFFFC410324FB595D3A" box="[992,1220,1337,1360]" italics="true" pageId="3" pageNumber="4">Emausaurus ernsti</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6C4D2AE91FFFFFFD1A034AFC6F5D04" authorityName="Owen" authorityYear="1861" box="[699,1010,1367,1390]" class="Reptilia" family="Scelidosauridae" genus="Scelidosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="harrisonii">
<emphasis id="B918EABBE91FFFFFFD1A034AFC6F5D04" box="[699,1010,1367,1390]" italics="true" pageId="3" pageNumber="4">Scelidosaurus harrisonii</emphasis>
</taxonomicName>
, and stegosaurs. The primitive ankylosaurids
<taxonomicName id="4C6C4D2AE91FFFFFFB900369FCF15DC2" authorityName="Tumanova" authorityYear="1993" class="Reptilia" family="Ankylosauridae" genus="Tsagantegia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="longicranialis">
<emphasis id="B918EABBE91FFFFFFB900369FCF15DC2" italics="true" pageId="3" pageNumber="4">Tsagantegia longicranialis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6C4D2AE91FFFFFFC10038CFB565DC2" authorityName="Tumanova" authorityYear="1983" box="[945,1227,1425,1448]" class="Reptilia" family="Ankylosauridae" genus="Shamosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="scutatus">
<emphasis id="B918EABBE91FFFFFFC10038CFB565DC2" box="[945,1227,1425,1448]" italics="true" pageId="3" pageNumber="4">Shamosaurus scutatus</emphasis>
</taxonomicName>
also exhibit this condition. The presence of a narrow skull in adult
<taxonomicName id="4C6C4D2AE91FFFFFFC6C03D1FBF05D89" authorityName="Gilmore" authorityYear="1933" box="[973,1133,1484,1507]" class="Reptilia" family="Ankylosauridae" genus="Pinacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
<emphasis id="B918EABBE91FFFFFFC6C03D1FBF05D89" box="[973,1133,1484,1507]" italics="true" pageId="3" pageNumber="4">Pinacosaurus</emphasis>
</taxonomicName>
may be just one indicator of a phylogenetic affinity with these more primitive genera that has previously been obscured, as many of the characters considered diagnostic for the genus (and indeed, for suprageneric ankylosaurian taxa) have been scored from juvenile skulls that do not exhibit all features of the adult. In particular, the exceptionally com­ plete skull of
<taxonomicName id="4C6C4D2AE918FFF8FE8202C9FE365C81" authorityName="Gilmore" authorityYear="1933" box="[291,427,1236,1259]" class="Reptilia" family="Ankylosauridae" genus="Pinacosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Ornithischia" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="grangeri">
<emphasis id="B918EABBE918FFF8FE8202C9FE365C81" box="[291,427,1236,1259]" italics="true" pageId="4" pageNumber="5">P. grangeri</emphasis>
</taxonomicName>
described by
<bibRefCitation id="EFFD4B58E918FFF8FDF502C9FE865D62" author="Maryanska, T." pageId="4" pageNumber="5" pagination="45 - 53" refId="ref12555" refString="Maryanska, T. 1971. New data on the skull of Pinacosaurus grangeri (Ankylosauria). Palaeontologia Polonica 25: 45 - 53." type="journal article" year="1971">Maryańska (1971</bibRefCitation>
; 1977), ZPAL MgD­II/1, is frequently cited as displaying typical ankylosaurid morphology, despite the fact that it represents a juvenile individual (e.g.,
<bibRefCitation id="EFFD4B58E918FFF8FD940354FF275D17" author="Sereno, P. C." pageId="4" pageNumber="5" pagination="234 - 256" refId="ref13065" refString="Sereno, P. C. 1986. Phylogeny of the bird-hipped dinosaurs (Order Ornithischia). National Geographic Society Research 2: 234 - 256." type="journal article" year="1986">Sereno, 1986</bibRefCitation>
;
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).
</paragraph>
</subSubSection>
</treatment>
</document>
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