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<document id="CDBA670C97C6845496F1423F18047384" ID-DOI="10.1206/0003-0082(2005)481[0001:ZUNSAN]2.0.CO;2" ID-ISSN="0003-0082" ID-Zenodo-Dep="5383072" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" checkinTime="1630295965375" checkinUser="carolina" docAuthor="JOYCE, WALTER G. &amp; NORELL, MARK A." docDate="2005" docId="5F0C87EB1937FF8FD5DE2899FD7B9AA2" docLanguage="en" docName="N3481.pdf" docOrigin="American Museum Novitates 3481" docSource="http://www.bioone.org/doi/abs/10.1206/0003-0082%282005%29481%5B0001%3AZUNSAN%5D2.0.CO%3B2" docStyle="DocumentStyle:70DDF1BDF6846024993F05262BAE3001.3:AmMusNovit.2000-2010.journal_article.type1" docStyleId="70DDF1BDF6846024993F05262BAE3001" docStyleName="AmMusNovit.2000-2010.journal_article.type1" docStyleVersion="3" docTitle="Zangerlia ukhaachelys JOYCE &amp; NORELL 2005, new species" docType="treatment" docVersion="3" lastPageNumber="7" masterDocId="A335FF931934FF89D5772C49FFF79C6F" masterDocTitle="Zangerlia ukhaachelys, New Species, a Nanhsiungchelyid Turtle from the Late Cretaceous of Ukhaa Tolgod, Mongolia" masterLastPageNumber="20" masterPageNumber="1" pageNumber="4" updateTime="1699218809662" updateUser="plazi" zenodo-license-document="CC-BY-4.0">
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<mods:title id="26F1566B5E51B30A97EFE706799DD506">Zangerlia ukhaachelys, New Species, a Nanhsiungchelyid Turtle from the Late Cretaceous of Ukhaa Tolgod, Mongolia</mods:title>
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<mods:namePart id="E630EB14E04930C4041DD92013F43E4E">JOYCE, WALTER G.</mods:namePart>
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<mods:namePart id="BBE081B1D417CC9E599B0615AB6FA47A">NORELL, MARK A.</mods:namePart>
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<mods:title id="1D53862F074384F89437EE20933504D9">American Museum Novitates</mods:title>
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<mods:date id="38B86ECBA476FAFEE64F0B34BA2FCAAF">2005</mods:date>
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<mods:number id="074D1E0DD6EBB109EA958AF50D667E56">2005-07-25</mods:number>
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<mods:number id="DA00EFEA14E4A24DBFCC1478C7F39727">3481</mods:number>
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<mods:url id="90BB3C07CE8D80FBE2BBE91B3298C846">http://www.bioone.org/doi/abs/10.1206/0003-0082%282005%29481%5B0001%3AZUNSAN%5D2.0.CO%3B2</mods:url>
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<mods:identifier id="74B400D91A995162C5241D1B2EB98BE4" type="DOI">10.1206/0003-0082(2005)481[0001:ZUNSAN]2.0.CO;2</mods:identifier>
<mods:identifier id="D62A6AABAEBE509949B8A70CD62C530B" type="ISSN">0003-0082</mods:identifier>
<mods:identifier id="E996E45807E2ACF89F6A7B1C137491C8" type="Zenodo-Dep">5383072</mods:identifier>
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<treatment id="5F0C87EB1937FF8FD5DE2899FD7B9AA2" LSID="urn:lsid:plazi:treatment:5F0C87EB1937FF8FD5DE2899FD7B9AA2" httpUri="http://treatment.plazi.org/id/5F0C87EB1937FF8FD5DE2899FD7B9AA2" lastPageId="6" lastPageNumber="7" pageId="3" pageNumber="4">
<subSubSection id="9FBF65761937FF8AD5DE2899FDAA9888" box="[169,605,1232,1256]" pageId="3" pageNumber="4" type="nomenclature">
<paragraph id="D71A36FD1937FF8AD5DE2899FDAA9888" blockId="3.[169,605,1232,1256]" box="[169,605,1232,1256]" pageId="3" pageNumber="4">
<heading id="8C5281911937FF8AD5DE2899FDAA9888" box="[169,605,1232,1256]" centered="true" fontSize="36" level="2" pageId="3" pageNumber="4" reason="8">
<emphasis id="E5D1EAEF1937FF8AD5DE2899FE329887" bold="true" box="[169,453,1232,1256]" pageId="3" pageNumber="4">
<taxonomicName id="10A54D7E1937FF8AD5DE2899FE499887" authority="JOYCE &amp; NORELL, 2005" authorityName="JOYCE &amp; NORELL" authorityYear="2005" box="[169,446,1232,1256]" class="Reptilia" family="Nanhsiungchelyidae" genus="Zangerlia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="ukhaachelys" status="sp. nov.">
<emphasis id="E5D1EAEF1937FF8AD5DE2899FE499887" bold="true" box="[169,446,1232,1256]" italics="true" pageId="3" pageNumber="4">Zangerlia ukhaachelys</emphasis>
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,
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<taxonomicNameLabel id="FEE257941937FF8AD4B82899FDAA9888" box="[463,605,1232,1255]" pageId="3" pageNumber="4" rank="species">new species</taxonomicNameLabel>
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</paragraph>
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<subSubSection id="9FBF65761937FF8AD5EF28B7FD7C9A37" pageId="3" pageNumber="4" type="materials_examined">
<paragraph id="D71A36FD1937FF8AD5EF28B7FE3F9902" blockId="3.[123,652,1278,1742]" pageId="3" pageNumber="4">
<materialsCitation id="67CD3CA01937FF8AD5EF28B7FE3F9902" collectionCode="IGM" pageId="3" pageNumber="4" specimenCode="IGM 90" specimenCount="1" typeStatus="holotype">
<typeStatus id="081E885F1937FF8AD5EF28B7FEF9997A" box="[152,270,1278,1301]" pageId="3" pageNumber="4" type="holotype">HOLOTYPE</typeStatus>
:
<specimenCode id="87039E861937FF8AD45528B7FE70997A" box="[290,391,1278,1301]" collectionCode="IGM" country="Mongolia" httpUri="http://grbio.org/cool/0h5a-qjh4" name="Geological Institute, Mongolian Academy of Sciences" pageId="3" pageNumber="4">IGM 90</specimenCode>
/1 (
<figureCitation id="4F9E2A781937FF8AD4D828B7FDEA997A" box="[431,541,1278,1301]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="4.[150,188,1130,1151]" captionStartId-1="5.[149,187,1505,1526]" captionStartId-2="9.[149,187,1385,1406]" captionStartId-3="10.[150,188,1385,1406]" captionTargetBox-0="[125,1229,187,1109]" captionTargetBox-1="[123,1227,187,1485]" captionTargetBox-2="[123,1227,187,1365]" captionTargetBox-3="[129,1210,259,1361]" captionTargetId-0="figure-15@4.[125,1229,187,1109]" captionTargetId-1="figure-11@5.[123,1227,187,1485]" captionTargetId-2="figure-13@9.[123,1227,187,1365]" captionTargetId-3="figure-15@10.[125,1229,187,1365]" captionTargetPageId-0="4" captionTargetPageId-1="5" captionTargetPageId-2="9" captionTargetPageId-3="10" captionText-0="Fig. 1. Zangerlia ukhaachelys. IGM 90/1, holotype, Upper Cretaceous of Mongolia. Photographs were manipulated digitally to enhance sutures. A, Oblique dorsal view of mandible; B, dorsal view of mandible; C, dorsal view of cranium; D, ventral view of cranium. Abbreviations: desc proc, descending process of frontal; ex, exoccipital; fr, frontal; mx, maxilla; op, opisthotic; pa, parietal; pf, prefrontal; pm, premaxilla; po, postorbital; pro, prootic; qu, quadrate; rec lab op, recessus labyrinthicus opisthoticus; rec lab pro, recessus labyrinthicus prooticus; so, supraoccipital; vo, vomer;?, bone of uncertain homology." captionText-1="Fig. 2. Zangerlia ukhaachelys, cranium of IGM 90/1, holotype, Upper Cretaceous of Mongolia. Photographs were manipulated digitally to enhance sutures. A, Left lateral view; B, right lateral view; C, frontal view; D, caudal view. See figure 1 for abbreviations." captionText-2="Fig. 3. Zangerlia ukhaachelys, IGM 90/1, holotype, Upper Cretaceous of Mongolia. Ventral view of plastron." captionText-3="Fig. 4. Zangerlia ukhaachelys, IGM 90/1, holotype, Upper Cretaceous of Mongolia. Ventral view of plastron. Abbreviations: AB, abdominal scute; AN, anal scute; ent, entoplastron; epi, epiplastron; FE, femoral scute; HU, humeral scute; hyo, hyoplastron; hyp, hypoplastron; IM, inframarginal scute; PE, pectoral scute; per, peripheral; xi, xiphiplastron." figureDoi-0="http://doi.org/10.5281/zenodo.5383074" figureDoi-1="http://doi.org/10.5281/zenodo.5383076" figureDoi-2="http://doi.org/10.5281/zenodo.5383078" figureDoi-3="http://doi.org/10.5281/zenodo.5383080" httpUri-0="https://zenodo.org/record/5383074/files/figure.png" httpUri-1="https://zenodo.org/record/5383076/files/figure.png" httpUri-2="https://zenodo.org/record/5383078/files/figure.png" httpUri-3="https://zenodo.org/record/5383080/files/figure.png" pageId="3" pageNumber="4">figs. 1–4</figureCitation>
), incomplete skeleton consisting of partial cranium, peripherals, plastron, and fragmentary other remains of the postcranium.
</materialsCitation>
</paragraph>
<paragraph id="D71A36FD1937FF8AD5EF293DFD7C9A37" blockId="3.[123,652,1278,1742]" pageId="3" pageNumber="4">
TYPE LOCALITY: Ukhaa Tolgod, just south of Xanadu, Omongov Aimag,
<collectingCountry id="AFB2766D1937FF8AD4A929D8FDA499C7" box="[478,595,1425,1448]" name="Mongolia" pageId="3" pageNumber="4">Mongolia</collectingCountry>
. The Ukhaa Tolgod beds have been considered to be roughly equivalent to Djadoktha (
<bibRefCitation id="B3344B0C1937FF8AD7482985FEFB9A6F" author="Loope, D. B. &amp; L. Dingus &amp; C. C. Swisher III &amp; C. Minjin" pageId="3" pageNumber="4" pagination="27 - 30" refId="ref8429" refString="Loope, D. B., L. Dingus, C. C. Swisher III, and C. Minjin. 1998. Life and death in a Cretaceous dune field, Nemegt Basin, Mongolia. Geology 26: 27 - 30." type="journal article" year="1998">Loope et al., 1998</bibRefCitation>
;
<bibRefCitation id="B3344B0C1937FF8AD46929A0FDC99A6F" author="Dashzeveg, D. &amp; M. J. Novacek &amp; M. A. Norell &amp; J. M. Clark &amp; L. M. Chiappe &amp; A. Davidson &amp; L. Dingus &amp; C. Swisher &amp; P. Altangerel" box="[286,574,1513,1536]" pageId="3" pageNumber="4" pagination="446 - 449" refId="ref7866" refString="Dashzeveg, D., M. J. Novacek, M. A. Norell, J. M. Clark, L. M. Chiappe, A. Davidson, M. C. Mc- Kenna, L. Dingus, C. Swisher, and P. Altangerel. 1995. Extraordinary preservation in a new vertebrate assemblage from the Late Cretaceous of Mongolia. Nature 374: 446 - 449." type="journal article" year="1995">Dashzeveg et al., 1995</bibRefCitation>
). Djadoktha beds elsewhere in
<collectingCountry id="AFB2766D1937FF8AD4C72A4FFDD69A72" box="[432,545,1542,1565]" name="Mongolia" pageId="3" pageNumber="4">Mongolia</collectingCountry>
are considered to be Late Campanian (
<bibRefCitation id="B3344B0C1937FF8AD7732A6DFE999A37" author="Lillegraven, J. A. &amp; M. C. McKenna" pageId="3" pageNumber="4" pagination="1 - 68" refId="ref8349" refString="Lillegraven, J. A., and M. C. McKenna. 1986. Fossil mammals from the '' Mesaverde' ' Formation (Late Cretaceous, Judithian) of the Bighorn and Wind River Basins, Wyoming, with definitions of Late Cretaceous North American Land Mammal '' Ages' '. American Museum Novitates 2840: 1 - 68." type="journal article" year="1986">Lillegraven and McKenna, 1986</bibRefCitation>
;
<bibRefCitation id="B3344B0C1937FF8AD40C2A08FD889A37" author="Gao, K. - Q. &amp; M. A. Norell" box="[379,639,1601,1624]" pageId="3" pageNumber="4" pagination="1 - 118" refId="ref8049" refString="Gao, K. - Q., and M. A. Norell. 2000. Taxonomic composition and systematics of Late Cretaceous lizard assemblages from Ukhaa Tolgod and adjacent localities, Mongolian Gobi Desert. Bulletin of the American Museum of Natural History 249: 1 - 118." type="journal article" year="2000">Gao and Norell, 2000</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="9FBF65761937FF8AD5EF2A17FE4D9ADF" pageId="3" pageNumber="4" type="etymology">
<paragraph id="D71A36FD1937FF8AD5EF2A17FE4D9ADF" blockId="3.[123,652,1278,1742]" pageId="3" pageNumber="4">
ETYMOLOGY: ‘‘Ukhaa’’, in reference to the fossil locality Ukhaa Tolgod,
<collectingCountry id="AFB2766D1937FF8AD4A82A35FDA49AFC" box="[479,595,1660,1683]" name="Mongolia" pageId="3" pageNumber="4">Mongolia</collectingCountry>
, and ‘‘chelys’’, Greek for turtle.
</paragraph>
</subSubSection>
<subSubSection id="9FBF65761937FF8AD5EF2AFFFC7C9F14" pageId="3" pageNumber="4" type="diagnosis">
<paragraph id="D71A36FD1937FF8AD5EF2AFFFC7C9F14" blockId="3.[123,652,1278,1742]" lastBlockId="3.[699,1228,193,891]" pageId="3" pageNumber="4">
DIAGNOSIS:
<taxonomicName id="10A54D7E1937FF8AD45F2AFEFDCD9AA1" authorityName="JOYCE &amp; NORELL" authorityYear="2005" box="[296,570,1719,1742]" class="Reptilia" family="Nanhsiungchelyidae" genus="Zangerlia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="ukhaachelys">
<emphasis id="E5D1EAEF1937FF8AD45F2AFEFDCD9AA1" box="[296,570,1719,1742]" italics="true" pageId="3" pageNumber="4">Zangerlia ukhaachelys</emphasis>
</taxonomicName>
is diagnosed by a single autapomorphy, the presence of an anteromedial process of the hyoplastron that limits contact between the entoplastron and epiplastron. In addition,
<taxonomicName id="10A54D7E1937FF8AD1E32D50FC469D21" authorityName="JOYCE &amp; NORELL" authorityYear="2005" class="Reptilia" family="Nanhsiungchelyidae" genus="Zangerlia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="3" pageNumber="4" phylum="Chordata" rank="species" species="ukhaachelys">
<emphasis id="E5D1EAEF1937FF8AD1E32D50FC469D21" italics="true" pageId="3" pageNumber="4">Zangerlia ukhaachelys</emphasis>
</taxonomicName>
is diagnosed by the following list of synapomorphies and symplesiomorphies: cranium with enlarged nasal cavity; wide fissura ethmoidalis; well­developed upper temporal emargination; frontals do not contribute to orbit and produce descending processes that almost surround sulcus olfactorius ventrally; lingual ridges absent; antorbital groove present along anteroventral rim of orbit; shell covered with deep pock­marks, deep nuchal notch formed by small and trapezoid nuchal and first peripheral; vertebral and pleural scutes do not overlap onto peripherals; entoplastron dissected by humero­pectoral sulcus; four pairs of inframarginals present which fully separate marginals from plastral scutes; marginal VI not expanded ventromedially; pectoral does not contribute to axillary notch; midline plastral sulci straight.
</paragraph>
</subSubSection>
<subSubSection id="9FBF65761937FF8FD7C82FD6FD7B9AA2" lastPageId="6" lastPageNumber="7" pageId="3" pageNumber="4" type="description">
<paragraph id="D71A36FD1937FF8AD7C82FD6FB309FD9" blockId="3.[703,1223,927,950]" box="[703,1223,927,950]" pageId="3" pageNumber="4">DESCRIPTION AND COMPARISON OF IGM 90/1</paragraph>
<paragraph id="D71A36FD1937FF8AD7CC2F82FCA89F8D" blockId="3.[699,863,971,994]" box="[699,863,971,994]" pageId="3" pageNumber="4">PRESERVATION</paragraph>
<paragraph id="D71A36FD1937FF8AD7AF2FBFFB5F9A54" blockId="3.[699,1228,1014,1741]" pageId="3" pageNumber="4">
Unlike many other fossils from the locality of Ukhaa Tolgod, IGM 90/1 is only moderately well preserved and shows signs of pre­ or postdepositional decay due to scavenging and/or postburial insect activity. Most limb elements are absent, and only fragmentary remains of the girdles were found within the shell. No traces of vertebral column elements, including the neural elements of the carapace, are present. Despite the lack of these elements, parts of the skull remain intact and display details in morphology never seen before in a nanhsiungchelyid turtle. The skull was found within the carapace just above the entoplastron. This position, however, is not considered positive evidence for full neck retraction in
<taxonomicName id="10A54D7E1937FF8AD6B12985FB33998D" authorityName="YEH" authorityYear="1966" box="[966,1220,1484,1507]" class="Reptilia" family="Nanhsiungchelyidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="3" pageNumber="4" phylum="Chordata" rank="family">NANHSIUNGCHELYIDAE</taxonomicName>
, but may be coincidental, because the cranium is not in articulation with the neck and may have been moved before final deposition.
</paragraph>
<paragraph id="D71A36FD1937FF8DD7AF2A08FE919A7A" blockId="3.[699,1228,1014,1741]" lastBlockId="4.[124,653,1358,1557]" lastPageId="4" lastPageNumber="5" pageId="3" pageNumber="4">
The shell is incompletely preserved and many areas that are useful in diagnosing NANHSIUNGCHELYID species are missing. Only the anterior and lateral peripherals remain of the carapace, thus obscuring the morphology of most carapacial scutes and the nuchal and pygal region. The plastron is more complete, but significant parts of the anterior lobe and the bridges are missing or they are heavily weathered. The larvae of large scavenging beetles likely produced the holes seen in the plastron (
<figureCitation id="4F9E2A781930FF8DD59B29B7FEAE9A7A" box="[236,345,1534,1557]" captionStart-0="Fig" captionStart-1="Fig" captionStart-2="Fig" captionStart-3="Fig" captionStartId-0="4.[150,188,1130,1151]" captionStartId-1="5.[149,187,1505,1526]" captionStartId-2="9.[149,187,1385,1406]" captionStartId-3="10.[150,188,1385,1406]" captionTargetBox-0="[125,1229,187,1109]" captionTargetBox-1="[123,1227,187,1485]" captionTargetBox-2="[123,1227,187,1365]" captionTargetBox-3="[129,1210,259,1361]" captionTargetId-0="figure-15@4.[125,1229,187,1109]" captionTargetId-1="figure-11@5.[123,1227,187,1485]" captionTargetId-2="figure-13@9.[123,1227,187,1365]" captionTargetId-3="figure-15@10.[125,1229,187,1365]" captionTargetPageId-0="4" captionTargetPageId-1="5" captionTargetPageId-2="9" captionTargetPageId-3="10" captionText-0="Fig. 1. Zangerlia ukhaachelys. IGM 90/1, holotype, Upper Cretaceous of Mongolia. Photographs were manipulated digitally to enhance sutures. A, Oblique dorsal view of mandible; B, dorsal view of mandible; C, dorsal view of cranium; D, ventral view of cranium. Abbreviations: desc proc, descending process of frontal; ex, exoccipital; fr, frontal; mx, maxilla; op, opisthotic; pa, parietal; pf, prefrontal; pm, premaxilla; po, postorbital; pro, prootic; qu, quadrate; rec lab op, recessus labyrinthicus opisthoticus; rec lab pro, recessus labyrinthicus prooticus; so, supraoccipital; vo, vomer;?, bone of uncertain homology." captionText-1="Fig. 2. Zangerlia ukhaachelys, cranium of IGM 90/1, holotype, Upper Cretaceous of Mongolia. Photographs were manipulated digitally to enhance sutures. A, Left lateral view; B, right lateral view; C, frontal view; D, caudal view. See figure 1 for abbreviations." captionText-2="Fig. 3. Zangerlia ukhaachelys, IGM 90/1, holotype, Upper Cretaceous of Mongolia. Ventral view of plastron." captionText-3="Fig. 4. Zangerlia ukhaachelys, IGM 90/1, holotype, Upper Cretaceous of Mongolia. Ventral view of plastron. Abbreviations: AB, abdominal scute; AN, anal scute; ent, entoplastron; epi, epiplastron; FE, femoral scute; HU, humeral scute; hyo, hyoplastron; hyp, hypoplastron; IM, inframarginal scute; PE, pectoral scute; per, peripheral; xi, xiphiplastron." figureDoi-0="http://doi.org/10.5281/zenodo.5383074" figureDoi-1="http://doi.org/10.5281/zenodo.5383076" figureDoi-2="http://doi.org/10.5281/zenodo.5383078" figureDoi-3="http://doi.org/10.5281/zenodo.5383080" httpUri-0="https://zenodo.org/record/5383074/files/figure.png" httpUri-1="https://zenodo.org/record/5383076/files/figure.png" httpUri-2="https://zenodo.org/record/5383078/files/figure.png" httpUri-3="https://zenodo.org/record/5383080/files/figure.png" pageId="4" pageNumber="5">figs. 1–4</figureCitation>
).
</paragraph>
<caption id="83DA66751930FF8DD5E12823FF1A9970" ID-DOI="http://doi.org/10.5281/zenodo.5383074" ID-Zenodo-Dep="5383074" httpUri="https://zenodo.org/record/5383074/files/figure.png" pageId="4" pageNumber="5" startId="4.[150,188,1130,1151]" targetBox="[125,1229,187,1109]" targetPageId="4">
<paragraph id="D71A36FD1930FF8DD5E12823FF1A9970" blockId="4.[124,1229,1130,1311]" pageId="4" pageNumber="5">
Fig. 1.
<taxonomicName id="10A54D7E1930FF8DD58F2823FE1C9810" authorityName="JOYCE &amp; NORELL" authorityYear="2005" box="[248,491,1130,1151]" class="Reptilia" family="Nanhsiungchelyidae" genus="Zangerlia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="4" pageNumber="5" phylum="Chordata" rank="species" species="ukhaachelys">
<emphasis id="E5D1EAEF1930FF8DD58F2823FE1C9810" box="[248,491,1130,1151]" italics="true" pageId="4" pageNumber="5">Zangerlia ukhaachelys</emphasis>
</taxonomicName>
. IGM 90/1, holotype, Upper Cretaceous of Mongolia. Photographs were manipulated digitally to enhance sutures.
<emphasis id="E5D1EAEF1930FF8DD70D28CDFD6498F6" bold="true" box="[634,659,1156,1177]" pageId="4" pageNumber="5">A,</emphasis>
Oblique dorsal view of mandible;
<emphasis id="E5D1EAEF1930FF8DD16728CDFBDF98F6" bold="true" box="[1040,1064,1156,1177]" pageId="4" pageNumber="5">B,</emphasis>
dorsal view of mandible;
<emphasis id="E5D1EAEF1930FF8DD59B28D6FEF198DB" bold="true" box="[236,262,1183,1204]" pageId="4" pageNumber="5">C,</emphasis>
dorsal view of cranium;
<emphasis id="E5D1EAEF1930FF8DD76628D6FDDD98DB" bold="true" box="[529,554,1183,1204]" pageId="4" pageNumber="5">D,</emphasis>
ventral view of cranium. Abbreviations:
<emphasis id="E5D1EAEF1930FF8DD6A828D6FBA798DB" bold="true" box="[991,1104,1183,1204]" pageId="4" pageNumber="5">desc proc,</emphasis>
descending process of frontal;
<emphasis id="E5D1EAEF1930FF8DD43C28F3FE9D98A0" bold="true" box="[331,362,1210,1231]" pageId="4" pageNumber="5">ex,</emphasis>
exoccipital;
<emphasis id="E5D1EAEF1930FF8DD48F28F3FDE498A0" bold="true" box="[504,531,1210,1231]" pageId="4" pageNumber="5">fr,</emphasis>
frontal;
<emphasis id="E5D1EAEF1930FF8DD70328F3FD6A98A0" bold="true" box="[628,669,1210,1231]" pageId="4" pageNumber="5">mx,</emphasis>
maxilla;
<emphasis id="E5D1EAEF1930FF8DD67028F3FCDE98A0" bold="true" box="[775,809,1210,1231]" pageId="4" pageNumber="5">op,</emphasis>
opisthotic;
<emphasis id="E5D1EAEF1930FF8DD6DC28F3FC3A98A0" bold="true" box="[939,973,1210,1231]" pageId="4" pageNumber="5">pa,</emphasis>
parietal;
<emphasis id="E5D1EAEF1930FF8DD14028F3FBA298A0" bold="true" box="[1079,1109,1210,1231]" pageId="4" pageNumber="5">pf,</emphasis>
prefrontal;
<emphasis id="E5D1EAEF1930FF8DD50B289DFF509886" bold="true" box="[124,167,1236,1257]" pageId="4" pageNumber="5">pm,</emphasis>
premaxilla;
<emphasis id="E5D1EAEF1930FF8DD447289DFEA59886" bold="true" box="[304,338,1236,1257]" pageId="4" pageNumber="5">po,</emphasis>
postorbital;
<emphasis id="E5D1EAEF1930FF8DD4AC289DFDFE9886" bold="true" box="[475,521,1236,1257]" pageId="4" pageNumber="5">pro,</emphasis>
prootic;
<emphasis id="E5D1EAEF1930FF8DD71A289DFD679886" bold="true" box="[621,656,1236,1257]" pageId="4" pageNumber="5">qu,</emphasis>
quadrate;
<emphasis id="E5D1EAEF1930FF8DD674289DFC8A9886" bold="true" box="[771,893,1236,1257]" pageId="4" pageNumber="5">rec lab op,</emphasis>
recessus labyrinthicus opisthoticus;
<emphasis id="E5D1EAEF1930FF8DD5C928A6FEB4996B" bold="true" box="[190,323,1263,1284]" pageId="4" pageNumber="5">rec lab pro,</emphasis>
recessus labyrinthicus prooticus;
<emphasis id="E5D1EAEF1930FF8DD7C728A6FD3A996B" bold="true" box="[688,717,1263,1284]" pageId="4" pageNumber="5">so,</emphasis>
supraoccipital;
<emphasis id="E5D1EAEF1930FF8DD60E28A6FC6D996B" bold="true" box="[889,922,1263,1284]" pageId="4" pageNumber="5">vo,</emphasis>
vomer;?, bone of uncertain homology.
</paragraph>
</caption>
<paragraph id="D71A36FD1930FF8DD50B2A7FFE579A23" blockId="4.[124,416,1590,1613]" box="[124,416,1590,1613]" pageId="4" pageNumber="5">CRANIUM AND MANDIBLE</paragraph>
<paragraph id="D71A36FD1930FF8DD5EE2A17FD7B9AA2" blockId="4.[124,653,1630,1741]" pageId="4" pageNumber="5">
The most interesting aspect of IGM 90/1 is its uncrushed, partial cranium, which is the best preserved of any nanhsiungchelyid and clearly exhibits all cranial sutures (
<figureCitation id="4F9E2A781930FF8DD7612AFFFDA89AA2" box="[534,607,1718,1741]" captionStart="Fig" captionStartId="4.[150,188,1130,1151]" captionTargetBox="[125,1229,187,1109]" captionTargetId="figure-15@4.[125,1229,187,1109]" captionTargetPageId="4" captionText="Fig. 1. Zangerlia ukhaachelys. IGM 90/1, holotype, Upper Cretaceous of Mongolia. Photographs were manipulated digitally to enhance sutures. A, Oblique dorsal view of mandible; B, dorsal view of mandible; C, dorsal view of cranium; D, ventral view of cranium. Abbreviations: desc proc, descending process of frontal; ex, exoccipital; fr, frontal; mx, maxilla; op, opisthotic; pa, parietal; pf, prefrontal; pm, premaxilla; po, postorbital; pro, prootic; qu, quadrate; rec lab op, recessus labyrinthicus opisthoticus; rec lab pro, recessus labyrinthicus prooticus; so, supraoccipital; vo, vomer;?, bone of uncertain homology." figureDoi="http://doi.org/10.5281/zenodo.5383074" httpUri="https://zenodo.org/record/5383074/files/figure.png" pageId="4" pageNumber="5">figs. 1</figureCitation>
,
<figureCitation id="4F9E2A781930FF8DD71A2AFFFD899AA2" box="[621,638,1718,1741]" captionStart="Fig" captionStartId="5.[149,187,1505,1526]" captionTargetBox="[123,1227,187,1485]" captionTargetId="figure-11@5.[123,1227,187,1485]" captionTargetPageId="5" captionText="Fig. 2. Zangerlia ukhaachelys, cranium of IGM 90/1, holotype, Upper Cretaceous of Mongolia. Photographs were manipulated digitally to enhance sutures. A, Left lateral view; B, right lateral view; C, frontal view; D, caudal view. See figure 1 for abbreviations." figureDoi="http://doi.org/10.5281/zenodo.5383076" httpUri="https://zenodo.org/record/5383076/files/figure.png" pageId="4" pageNumber="5">2</figureCitation>
).
</paragraph>
<paragraph id="D71A36FD1930FF8FD7AE2907FE269E4E" blockId="4.[700,1229,1358,1741]" lastBlockId="6.[124,653,193,1741]" lastPageId="6" lastPageNumber="7" pageId="4" pageNumber="5">PREFRONTAL: The prefrontal of IGM 90/1 consists of a dorsal plate, which forms the roof of a broad fossa nasalis, and a vertical plate, which forms significant portions of the anterior orbit wall. In dorsal view, the prefrontal has a posteromedial contact with the frontal, a medial contact with its counterpart, and a small posterolateral contact with the postorbital, thus excluding the frontal from contributing to the orbital rim. The anterior rims of both prefrontals are weathered away, making it unclear if nasals were present anteriorly. In lateral view, the vertical plate of the prefrontal meets the ascending process of the maxilla along a short interdigitated contact to form the lateral wall of the fossa nasalis. An additional, elongate contact between these bones exists along the anteroventral rim of the orbit. Although the distal ends of the ascending process of the prefrontals are weathered, the left one is better preserved and shows the presence of a moderately sized foramen orbito­nasale. The fissura ethmoidalis and the sulcus olfactorius are apparent and are notably wide.</paragraph>
<caption id="83DA66751931FF8CD5E229A8FCD19A43" ID-DOI="http://doi.org/10.5281/zenodo.5383076" ID-Zenodo-Dep="5383076" httpUri="https://zenodo.org/record/5383076/files/figure.png" pageId="5" pageNumber="6" startId="5.[149,187,1505,1526]" targetBox="[123,1227,187,1485]" targetPageId="5">
<paragraph id="D71A36FD1931FF8CD5E229A8FCD19A43" blockId="5.[123,1228,1505,1580]" pageId="5" pageNumber="6">
Fig. 2.
<taxonomicName id="10A54D7E1931FF8CD58F29ABFE1B9998" authorityName="JOYCE &amp; NORELL" authorityYear="2005" box="[248,492,1506,1527]" class="Reptilia" family="Nanhsiungchelyidae" genus="Zangerlia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="5" pageNumber="6" phylum="Chordata" rank="species" species="ukhaachelys">
<emphasis id="E5D1EAEF1931FF8CD58F29ABFE1B9998" box="[248,492,1506,1527]" italics="true" pageId="5" pageNumber="6">Zangerlia ukhaachelys</emphasis>
</taxonomicName>
, cranium of IGM 90/1, holotype, Upper Cretaceous of Mongolia. Photographs were manipulated digitally to enhance sutures.
<emphasis id="E5D1EAEF1931FF8CD67229B5FCE99A7E" bold="true" box="[773,798,1532,1553]" pageId="5" pageNumber="6">A,</emphasis>
Left lateral view;
<emphasis id="E5D1EAEF1931FF8CD69D29B5FBF49A7E" bold="true" box="[1002,1027,1532,1553]" pageId="5" pageNumber="6">B,</emphasis>
right lateral view;
<emphasis id="E5D1EAEF1931FF8CD50C2A5FFF639A44" bold="true" box="[123,148,1558,1579]" pageId="5" pageNumber="6">C,</emphasis>
frontal view;
<emphasis id="E5D1EAEF1931FF8CD4582A5FFEB09A44" bold="true" box="[303,327,1558,1579]" pageId="5" pageNumber="6">D,</emphasis>
caudal view. See figure 1 for abbreviations.
</paragraph>
</caption>
<paragraph id="D71A36FD1932FF8FD5EE2E6EFDDC9F42" blockId="6.[124,653,193,1741]" pageId="6" pageNumber="7">
The prefrontal of IGM 90/1 closely resembles that of
<taxonomicName id="10A54D7E1932FF8FD4702E0FFDC19E32" authorityName="Brinkman and Peng" authorityYear="1996" box="[263,566,582,605]" class="Reptilia" family="Nanhsiungchelyidae" genus="Zangerlia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="neimongolensis">
<emphasis id="E5D1EAEF1932FF8FD4702E0FFDC19E32" box="[263,566,582,605]" italics="true" pageId="6" pageNumber="7">Zangerlia neimongolensis</emphasis>
</taxonomicName>
. It also resembles the prefrontals of
<taxonomicName id="10A54D7E1932FF8FD4A32E2AFDA09E15" box="[468,599,611,634]" class="Reptilia" family="Adocidae" genus="Adocus" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="E5D1EAEF1932FF8FD4A32E2AFDDD9E15" box="[468,554,611,634]" italics="true" pageId="6" pageNumber="7">Adocus</emphasis>
sp.
</taxonomicName>
, but differs by having a contact with the postorbital and by covering a much larger proportion of the dorsal roofing of the skull between the orbits. The fissura ethmoidalis greatly resembles that of the
<taxonomicName id="10A54D7E1932FF8FD4B82EB0FD989F60" box="[463,623,761,784]" class="Reptilia" family="Testudinidae" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="7" phylum="Chordata" rank="family">TESTUDINIDAE</taxonomicName>
in being wide and not keyhole­shaped.
</paragraph>
<paragraph id="D71A36FD1932FF8FD5EE2F7DFEBC99CF" blockId="6.[124,653,193,1741]" pageId="6" pageNumber="7">FRONTAL: In dorsal view, the frontal of IGM 90/1 is flat and subtriangular. It meets the prefrontal anterolaterally, the postorbital laterally, the other frontal medially, and the parietal along an interdigitated suture posteriorly. It is a large element forming much of the skull roof, but it does not contribute to the orbital rim. The frontal generally has a smooth surface, but a faint sulcus crosses it in an arch. The morphology of the frontal is more complex in ventral view. Between the parietal and the prefrontal, it forms a significant portion of the sulcus olfactorius. From the rim of the sulcus, tapered processes ascend from both frontals that almost meet medially, practically rendering the sulcus olfactorius a canal. Transverse to the sulcus olfactorius, the prefrontal additionally forms a small ridge that runs parallel to its border with the parietal and forms the posterodorsal limit of the orbit.</paragraph>
<paragraph id="D71A36FD1932FF8FD5EE29EEFD7B9AA2" blockId="6.[124,653,193,1741]" pageId="6" pageNumber="7">
The skull materials of
<taxonomicName id="10A54D7E1932FF8FD4C329E1FD7A99D0" authorityName="Brinkman and Peng" authorityYear="1996" box="[436,653,1448,1471]" class="Reptilia" family="Nanhsiungchelyidae" genus="Zangerlia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="neimongolensis">
<emphasis id="E5D1EAEF1932FF8FD4C329E1FD7A99D0" box="[436,653,1448,1471]" italics="true" pageId="6" pageNumber="7">Z. neimongolensis</emphasis>
</taxonomicName>
and
<taxonomicName id="10A54D7E1932FF8FD5D9298CFE9A99B3" authorityName="Yeh" authorityYear="1966" box="[174,365,1477,1500]" class="Reptilia" family="Nanhsiungchelyidae" genus="Nanhsiungchelys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="wuchingensis">
<emphasis id="E5D1EAEF1932FF8FD5D9298CFE9A99B3" box="[174,365,1477,1500]" italics="true" pageId="6" pageNumber="7">N. wuchingensis</emphasis>
</taxonomicName>
are not prepared enough to assess the presence of a descending frontal process in these taxa. These processes are not known in
<taxonomicName id="10A54D7E1932FF8FD58C2A69FE8A9A58" box="[251,381,1568,1591]" class="Reptilia" family="Adocidae" genus="Adocus" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="E5D1EAEF1932FF8FD58C2A69FEA69A58" box="[251,337,1568,1591]" italics="true" pageId="6" pageNumber="7">Adocus</emphasis>
sp.
</taxonomicName>
Among living turtles, descending frontal processes are known from a number of TESTUDINOID turtles, especially terrestrial forms of the
<taxonomicName id="10A54D7E1932FF8FD4DE2A33FDBE9AFF" box="[425,585,1658,1681]" class="Reptilia" family="Testudinidae" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="7" phylum="Chordata" rank="family">TESTUDINIDAE</taxonomicName>
. The frontals of
<taxonomicName id="10A54D7E1932FF8FD5882AD1FE229AC0" authorityName="Brinkman and Peng" authorityYear="1996" box="[255,469,1688,1711]" class="Reptilia" family="Nanhsiungchelyidae" genus="Zangerlia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Testudines" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="neimongolensis">
<emphasis id="E5D1EAEF1932FF8FD5882AD1FE229AC0" box="[255,469,1688,1711]" italics="true" pageId="6" pageNumber="7">Z. neimongolensis</emphasis>
</taxonomicName>
do not contribute to the orbital rim, as seen in IGM 90/1.
</paragraph>
</subSubSection>
</treatment>
</document>
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