736 lines
152 KiB
XML
736 lines
152 KiB
XML
<document ID-CLB-Dataset="53678" ID-DOI="http://dx.doi.org/10.3897/fr.26.102128" ID-GBIF-Dataset="2e4c0683-f935-4898-ac8e-83bc8589c2d7" ID-Pensoft-Pub="2193-0074-1-117" ID-Pensoft-UUID="5075F649FC065E1F9F13996C8F1A5FB1" ID-ZooBank="141AAAE38F1D4117A64BD240B6BDF660" ModsDocID="2193-0074-26-1-117" checkinTime="1685804667582" checkinUser="pensoft" docAuthor="Joyce, Walter G., Evers, Serjoscha W., Ren, Sara, Rollot, Yann & Schwermann, Achim H." docDate="2023" docId="E50DF76F76415E50933786C8FC1B46DC" docLanguage="en" docName="FossilRecord 26(1): 117-133" docOrigin="Fossil Record 26 (1)" docPubDate="2023-06-02" docSource="http://dx.doi.org/10.3897/fr.26.102128" docTitle="Helochelydra nopcsai Lapparent de Broin & Murelaga 1999" docType="treatment" docVersion="6" id="5075F649FC065E1F9F13996C8F1A5FB1" lastPageNumber="117" masterDocId="5075F649FC065E1F9F13996C8F1A5FB1" masterDocTitle="The helochelydrid turtle Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) fissure fills of Balve, North Rhine-Westphalia, Germany, including a large sample of granicones" masterLastPageNumber="133" masterPageNumber="117" pageNumber="117" updateTime="1732867471117" updateUser="ExternalLinkService">
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<mods:titleInfo id="1ACCBD6567B6F311A2AD448E156504B7">
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<mods:title id="140A945A54AC9F8A116D64D6AC3405B0">The helochelydrid turtle Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) fissure fills of Balve, North Rhine-Westphalia, Germany, including a large sample of granicones</mods:title>
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<mods:namePart id="E5A2CA42651D9DB40AF5C71847B5BF0A">Joyce, Walter G.</mods:namePart>
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<mods:affiliation id="6045DF97A85E111879B54F4CE705AFCD">Department of Geosciences, University of Fribourg, 1700 Fribourg, Switzerland</mods:affiliation>
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<mods:nameIdentifier id="EF3466DA93B12798FA10D607612B2065" type="email">walter.joyce@unifr.ch</mods:nameIdentifier>
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<mods:roleTerm id="1C014C87125494452D38B760976719D5">Author</mods:roleTerm>
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<mods:namePart id="9D08840F4E2B795E8567312ECF38458F">Evers, Serjoscha W.</mods:namePart>
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<mods:nameIdentifier id="50F16537820172E85AD12DC69EEBCF76" type="ORCID">https://orcid.org/0000-0002-2393-5621</mods:nameIdentifier>
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<mods:affiliation id="6D9F9AE0A5A83510C6B1D9B20E50F63B">Department of Geosciences, University of Fribourg, 1700 Fribourg, Switzerland</mods:affiliation>
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<mods:name id="A5D99B7E609025E44B14B82D2D8791E6" type="personal">
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<mods:namePart id="ED7C0AECC015607A9B04C5F310E1C49E">Ren, Sara</mods:namePart>
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<mods:affiliation id="2004CF47C6CC7BEEE3210D198C4DAADB">Department of Geosciences, University of Fribourg, 1700 Fribourg, Switzerland</mods:affiliation>
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<mods:namePart id="16E8730C46BA50F8B0BFC93EC3898EA9">Rollot, Yann</mods:namePart>
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<mods:nameIdentifier id="8A18AEB32D00C3D960A0F1C224F07046" type="ORCID">https://orcid.org/0000-0002-2020-9456</mods:nameIdentifier>
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<mods:affiliation id="22979CD40BA42E5514418CF613069FD7">Department of Geosciences, University of Fribourg, 1700 Fribourg, Switzerland</mods:affiliation>
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<mods:namePart id="41939D65E15517F3D47B613D428A08B3">Schwermann, Achim H.</mods:namePart>
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<mods:nameIdentifier id="DF821DD09945C5E7056930CE5ED97B6D" type="ORCID">https://orcid.org/0000-0001-8928-8773</mods:nameIdentifier>
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<mods:affiliation id="3D878881BEE8A9BFD9D98E7EC7729D69">LWL-Museum of Natural History, Westphalian State Museum and Planetarium, 48161 Mu ̈ nster, Germany</mods:affiliation>
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<mods:typeOfResource id="310764470C37BCA86B637D889A36A4BF">text</mods:typeOfResource>
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<mods:titleInfo id="06BD2EE2EB333A7CE94D44C8BBA39A9D">
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<mods:title id="43C6552057CE6DBF4438AA4FF840A418">Fossil Record</mods:title>
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<mods:part id="83CD99418B2C18B13819BC318231DB8A">
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<mods:date id="0C2AB99F247035A45FC2519A34FCB393">2023</mods:date>
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<mods:number id="2FD9541A37E67B6AEB45238CC53B7583">2023-06-02</mods:number>
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<mods:number id="B721E08FDC6D04AEAB243CB8CE86B7BE">26</mods:number>
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</mods:detail>
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<mods:detail id="24B77DB818DCB20476FEB232048CA01A" type="issue">
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<mods:number id="801C7AC3C2C136DADE816C1E75119C9D">1</mods:number>
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<mods:extent id="8BF558D05B002180DCB76F8B9DE6AF13" unit="page">
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<mods:start id="33F25F663170644AE4512FF87B674DB8">117</mods:start>
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<mods:end id="19F70E1C1237BC4776CB79DED49A523B">133</mods:end>
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<mods:url id="D77750C4C40CCD32C6DA7ED1C4464E05">http://dx.doi.org/10.3897/fr.26.102128</mods:url>
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<mods:classification id="011970EC20710343550E45A8659462B7">journal article</mods:classification>
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<mods:identifier id="619A46889E4A13484EFED38BC4B30079" type="DOI">http://dx.doi.org/10.3897/fr.26.102128</mods:identifier>
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<mods:identifier id="AA339FADA37CC726273EA4CAB38C90CF" type="Pensoft-Pub">2193-0074-1-117</mods:identifier>
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</mods:mods>
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<treatment id="E50DF76F76415E50933786C8FC1B46DC" ID-GBIF-Taxon="209667779" LSID="urn:lsid:plazi:treatment:E50DF76F76415E50933786C8FC1B46DC" httpUri="http://treatment.plazi.org/id/E50DF76F76415E50933786C8FC1B46DC" lastPageNumber="117" pageId="0" pageNumber="117" scope_subClass="Testudinata">
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<subSubSection id="1DDBC38B6B682C2938A2DEBDAC971850" pageId="0" pageNumber="117" type="nomenclature">
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<paragraph id="0386DEFA555D31A066FA2A2886832AD5" pageId="0" pageNumber="117">
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<taxonomicName id="3B47D34CABDC25F096C2B5A31BDB9C30" LSID="E50DF76F-7641-5E50-9337-86C8FC1B46DC" authority="Lapparent de Broin & Murelaga, 1999" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">Helochelydra nopcsai Lapparent de Broin & Murelaga, 1999</taxonomicName>
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</paragraph>
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</subSubSection>
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<subSubSection id="86E6D3F6BB297A58926FA57F2B80A2A0" pageId="0" pageNumber="117" type="type material">
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<paragraph id="F49CEEF8F60815B4A0831531EAB90D96" pageId="0" pageNumber="117">Type material.</paragraph>
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<paragraph id="236344F17B8CFECE81837D649BF6E899" pageId="0" pageNumber="117">
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|
NHMUK R171 (holotype), a fragmentary shell associated with limb and girdle elements (
|
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<bibRefCitation id="B7981D070107F93E7F11FEEB13A31C3E" author="Nopcsa, F" journalOrPublisher="Geologica Hungarica, Palaeontology Series" pageId="0" pageNumber="117" pagination="1 - 84" refId="B27" refString="Nopcsa, F, 1928. Palaeontological notes on reptiles. Geologica Hungarica, Palaeontology Series 1: 1 - 84" title="Palaeontological notes on reptiles." volume="1" year="1928">Nopcsa 1928</bibRefCitation>
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).
|
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</paragraph>
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|
</subSubSection>
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<subSubSection id="6AA400BE5A4FF1AB17A5D31561387CB9" pageId="0" pageNumber="117" type="type locality">
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<paragraph id="57417AB136EA9312DC22CD7147E00B6C" pageId="0" pageNumber="117">Type locality.</paragraph>
|
|
<paragraph id="1C323E72A3C0A820DEBB64DFC23556D0" pageId="0" pageNumber="117">Isle of Wight, United Kingdom (Nopcsa, 1928).</paragraph>
|
|
</subSubSection>
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|
<subSubSection id="FC071E5783DF302B785DF2BD1367875C" pageId="0" pageNumber="117" type="type stratum">
|
|
<paragraph id="A7A84A3E2C36AFADE50A116635330C71" pageId="0" pageNumber="117">Type stratum.</paragraph>
|
|
<paragraph id="90C0F0732F60AC471ABE454E2772886A" pageId="0" pageNumber="117">
|
|
Wessex Formation, Wealden Group, Barremian, Early Cretaceous (
|
|
<bibRefCitation id="784580562FD274A149E06679C3328F4F" author="Joyce, WG" journalOrPublisher="Special Papers in Paleontology" pageId="0" pageNumber="117" pagination="75 - 97" refId="B15" refString="Joyce, WG, Chapman, SD, Moody, RTJ, Walker, CA, 2011. The skull of the solemydid turtle Helochelydra nopcsai from the Early Cretaceous (Barremian) of the Isle of Wight (UK) and a review of Solemydidae. Special Papers in Paleontology 86: 75 - 97" title="The skull of the solemydid turtle Helochelydra nopcsai from the Early Cretaceous (Barremian) of the Isle of Wight (UK) and a review of Solemydidae." volume="86" year="2011">Joyce et al. 2011</bibRefCitation>
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).
|
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</paragraph>
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|
</subSubSection>
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|
<subSubSection id="6CB37C6C400B4CB6A958920E389107D4" pageId="0" pageNumber="117" type="diagnosis">
|
|
<paragraph id="92B8FFEFF1A1D76A8BB6C6484BE4829E" pageId="0" pageNumber="117">Diagnosis.</paragraph>
|
|
<paragraph id="E264DA0CB5421E643E18AD13BBC8A7FD" pageId="0" pageNumber="117">
|
|
<taxonomicName id="48042098B3A330241C6582526FC7393A" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
|
|
<emphasis id="7A2067689705E99FDCBD83FBF739F081" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
|
|
</taxonomicName>
|
|
can most easily be diagnosed as a helochelydrid by the presence of a secondary pair of occipital tubercles that are formed by the pterygoids, a triangular fossa that is formed by the squamosal at the posterior margin of the skull, a shell covered by a surface texture consisting of distinct tubercles, expansion of the visceral cavity to the front and back of the peripheral series (also present in some compsemydids and pleurosternids), and presence of an entoplastral scute.
|
|
<taxonomicName id="BD0F011B6F3BC493E3992DD04901024F" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
|
|
<emphasis id="6CCE544680AAE5388879D1DD4F16CFA7" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
|
|
</taxonomicName>
|
|
can be distinguished from other helochelydrids by a shell surface texture consisting of evenly spaced tubercles that easily dislocate (also present in
|
|
<taxonomicName id="5C65A3213CC063D18C2523BB720456ED" authorityName="von Meyer" authorityYear="1854" class="Reptilia" genus="Helochelys" kingdom="Animalia" lsidName="Helochelys danubina" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="species" species="danubina">
|
|
<emphasis id="2D67372391BE84ED6DC8BD262149F689" italics="true" pageId="0" pageNumber="117">Helochelys danubina</emphasis>
|
|
</taxonomicName>
|
|
), cone-, hook-, and plate-shaped osteoderms (also present in
|
|
<taxonomicName id="B5D8C5C8BB520F2E8EC3315DF5E1DAFC" authorityName="Hay" authorityYear="1908" class="Reptilia" family="Pleurosternidae" genus="Naomichelys" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Naomichelys speciosa" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="species" species="speciosa">
|
|
<emphasis id="419D248B0F83BD349212EA521C45E225" italics="true" pageId="0" pageNumber="117">Naomichelys speciosa</emphasis>
|
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</taxonomicName>
|
|
), and pterygoids that fully cover the parabasisphenoid ventrally. No other helochelydrid is known to possess compound granicones.
|
|
<taxonomicName id="A34BB622F01DF84B3D1CB431C165E663" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
|
|
<emphasis id="FB447323D7172209719A2D2DDDA84266" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
|
|
</taxonomicName>
|
|
currently cannot be distinguished from
|
|
<taxonomicName id="C076CC324C34DAF1AC35A50AEC9D69FA" authorityName="von Meyer" authorityYear="1854" class="Reptilia" genus="Helochelys" kingdom="Animalia" lsidName="Helochelys danubina" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="species" species="danubina">
|
|
<emphasis id="28DEE26D2A10607FEEAF19BCAE763AD5" italics="true" pageId="0" pageNumber="117">Helochelys danubina</emphasis>
|
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</taxonomicName>
|
|
using morphological features, because the latter is poorly known and its holotype lost. Stratigraphic criteria (i.e., an age difference of more than 20 Ma) nevertheless suggest that they represent distinct taxa.
|
|
</paragraph>
|
|
</subSubSection>
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<subSubSection id="D3244A88D018E9561DD2AE1CE452E447" pageId="0" pageNumber="117" type="materials_examined">
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<paragraph id="8FB190F4A76BA7DD1BD6DC34494C357E" pageId="0" pageNumber="117">Referred material.</paragraph>
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<paragraph id="E58F2DA250D135BA56F243689192429C" pageId="0" pageNumber="117">
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<materialsCitation id="734507A20A58B379FD1E81554D7DC795" ID-GBIF-Occurrence="4126307301" collectingDate="2023-01-01" collectingDateMax="2023-12-31" collectingDateMin="2023-01-01" country="United Kingdom" location="Isle of Wight" specimenCount="1" stateProvince="Isle of Wight">
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Early Cretaceous (Barremian), Wessex Formation,
|
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<collectingRegion id="A8C618DC8F02472AF57E3ABE614F9033" country="United Kingdom" name="Isle of Wight">Isle of Wight</collectingRegion>
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,
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<collectingCountry id="FEEFDEA5D59BF6032BBDA90E2C8096C9" name="United Kingdom">United Kingdom</collectingCountry>
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(material referred by
|
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<bibRefCitation id="2E46A1D38E0E5F44ABF0CB4BD1342D3E" author="Lapparent de Broin, F de" journalOrPublisher="Estudios del Museo de Ciencias Naturales de Alava" pageId="0" pageNumber="117" pagination="135 - 211" refId="B20" refString="Lapparent de Broin, F de, Murelaga, X, 1999. Turtles from the Upper Cretaceous of Lano (Iberian peninsula). Estudios del Museo de Ciencias Naturales de Alava 14: 135 - 211" title="Turtles from the Upper Cretaceous of Lano (Iberian peninsula)." volume="14" year="1999">Lapparent de Broin and Murelaga 1999</bibRefCitation>
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and
|
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<bibRefCitation id="878E54F40D844C896EDD956367B88F93" author="Joyce, WG" journalOrPublisher="Special Papers in Paleontology" pageId="0" pageNumber="117" pagination="75 - 97" refId="B15" refString="Joyce, WG, Chapman, SD, Moody, RTJ, Walker, CA, 2011. The skull of the solemydid turtle Helochelydra nopcsai from the Early Cretaceous (Barremian) of the Isle of Wight (UK) and a review of Solemydidae. Special Papers in Paleontology 86: 75 - 97" title="The skull of the solemydid turtle Helochelydra nopcsai from the Early Cretaceous (Barremian) of the Isle of Wight (UK) and a review of Solemydidae." volume="86" year="2011">Joyce et al. 2011</bibRefCitation>
|
|
;
|
|
<taxonomicName id="CBC30D5BB03000975AF70E97B6611E7F" authorityName="Cope" authorityYear="1868" class="Reptilia" family="Pleurosternidae" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="family">Pleurosternidae</taxonomicName>
|
|
of Sweetman and Insole 2010); Early Cretaceous (Hauterivian-Barremian), Nouvelle-Aquitaine,
|
|
<collectingCountry id="EE9C0B779CAFC11385ACB6C192294CC6" name="France">France</collectingCountry>
|
|
(
|
|
<taxonomicName id="383F26F7E0D3F5E1D04571AE52E59BCC" class="Reptilia" family="Solemydidae" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="family">Solemydidae</taxonomicName>
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|
of
|
|
<bibRefCitation id="66A6C25550650442A92903D4B8798856" DOI="https://doi.org/10.1016/j.cretres.2012.01.006" author="Neraudeau, D" journalOrPublisher="Cretaceous Research" pageId="0" pageNumber="117" pagination="1 - 14" refId="B26" refString="Neraudeau, D, Allain, R, Ballevre, M, Batten, DJ, Buffetaut, E, Colin, JP, Dabard, MP, 2012. The Hauterivian-Barremian lignitic bone bed of Angeac (Charente, south-west France): stratigraphical, palaeobiological and palaeogeographical implications. Cretaceous Research 37: 1 - 14, DOI: https://doi.org/10.1016/j.cretres.2012.01.006" title="The Hauterivian-Barremian lignitic bone bed of Angeac (Charente, south-west France): stratigraphical, palaeobiological and palaeogeographical implications." url="https://doi.org/10.1016/j.cretres.2012.01.006" volume="37" year="2012">
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<normalizedToken id="6090422288FBD8EEDC1B48E62ECA21F5" originalValue="Néraudeau">Neraudeau</normalizedToken>
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et al. 2012
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</bibRefCitation>
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); Early Cretaceous (Barremian),
|
|
<collectingRegion id="FD28FC105ED4DB1A49B8F91958D40B35" isEnumeration="true">
|
|
Aragon, Castile and
|
|
<normalizedToken id="F343A21953E4AF71C02645352D88EF28" originalValue="León">Leon</normalizedToken>
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, La Rioja
|
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</collectingRegion>
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, and
|
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<collectingRegion id="E160A79710FA07C66F3D5CBB9C34553C" country="Spain" name="Comunitat Valenciana">Valencia</collectingRegion>
|
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,
|
|
<collectingCountry id="F032F8A5ED6F4A7B0310F86C429A6A84" name="Spain">Spain</collectingCountry>
|
|
(
|
|
<taxonomicName id="0A5667ED766E5E3CBD63C81E7931ED61" class="Reptilia" family="Solemydidae" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="family">Solemydidae</taxonomicName>
|
|
of
|
|
<bibRefCitation id="76B8A12369D27A8B80E72A2FFFAA1E45" DOI="https://doi.org/10.7203/sjp.24.1.20346" author="Perez-Garcia, A" journalOrPublisher="Revista Espanola de Paleontologia" pageId="0" pageNumber="117" pagination="93 - 104" refId="B32" refString="Perez-Garcia, A, 2009. Revision historica y sistematica de las primeras citas de quelonios del Cretacico espanol. Revista Espanola de Paleontologia 24: 93 - 104, DOI: https://doi.org/10.7203/sjp.24.1.20346" title="Revision historica y sistematica de las primeras citas de quelonios del Cretacico espanol." url="https://doi.org/10.7203/sjp.24.1.20346" volume="24" year="2009">
|
|
<normalizedToken id="9046176A02105C5840193BD1CA8C8D98" originalValue="Pérez">Perez</normalizedToken>
|
|
<normalizedToken id="1F6F880CCAAF7822E34841375B80E712" originalValue="García">Garcia</normalizedToken>
|
|
2009
|
|
</bibRefCitation>
|
|
;
|
|
<bibRefCitation id="B554C9CC873BED6B94FE856DD753CD14" DOI="https://doi.org/10.1016/j.cretres.2011.09.011" author="Perez-Garcia, A" journalOrPublisher="Cretaceous Research" pageId="0" pageNumber="117" pagination="146 - 158" refId="B33" refString="Perez-Garcia, A, Murelaga, X, Huerta, P, Torcida Fernandez-Baldor, P, 2012. Turtles from the Lower Cretaceous of the Cameros Basin (Iberian Range, Spain). Cretaceous Research 33: 146 - 158, DOI: https://doi.org/10.1016/j.cretres.2011.09.011" title="Turtles from the Lower Cretaceous of the Cameros Basin (Iberian Range, Spain)." url="https://doi.org/10.1016/j.cretres.2011.09.011" volume="33" year="2012">
|
|
<normalizedToken id="952AF6BB8E91C7A9DB349A41A16528C9" originalValue="Pérez-García">Perez-Garcia</normalizedToken>
|
|
et al. 2012
|
|
</bibRefCitation>
|
|
,
|
|
<bibRefCitation id="AA3B5E4D0A99B6B0B1E4517A7ED6AC2D" DOI="https://doi.org/10.1016/j.cretres.2013.03.011" author="Perez-Garcia, A" journalOrPublisher="Cretaceous Research" pageId="0" pageNumber="117" pagination="64 - 82" refId="B34" refString="Perez-Garcia, A, Scheyer, TM, Murelaga, X, 2013. The turtles from the uppermost Jurassic and Early Cretaceous of Galve (Iberian Range, Spain): anatomical, systematic, biostratigraphic and palaeobiogeographical implications. Cretaceous Research 44: 64 - 82, DOI: https://doi.org/10.1016/j.cretres.2013.03.011" title="The turtles from the uppermost Jurassic and Early Cretaceous of Galve (Iberian Range, Spain): anatomical, systematic, biostratigraphic and palaeobiogeographical implications." url="https://doi.org/10.1016/j.cretres.2013.03.011" volume="44" year="2013">2013</bibRefCitation>
|
|
;
|
|
<bibRefCitation id="FEF51994FD8734A9160DE67DDD921F8B" DOI="https://doi.org/10.1007/s13127-014-0188-0" author="Scheyer, TM" journalOrPublisher="Organisms Diversity and Evolution" pageId="0" pageNumber="117" pagination="199 - 212" refId="B42" refString="Scheyer, TM, Perez-Garcia, A, Murelaga, X, 2015. Shell bone histology of solemydid turtles (stem Testudines): palaeoecological implications. Organisms Diversity and Evolution 15: 199 - 212, DOI: https://doi.org/10.1007/s13127-014-0188-0" title="Shell bone histology of solemydid turtles (stem Testudines): palaeoecological implications." url="https://doi.org/10.1007/s13127-014-0188-0" volume="15" year="2015">Scheyer et al. 2015</bibRefCitation>
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); Early Cretaceous (late Barremian - early Aptian), Balve,
|
|
<collectingCountry id="6C310A2328C2E1E2985617857E56EE26" name="Germany">Germany</collectingCountry>
|
|
(all material described herein)
|
|
</materialsCitation>
|
|
.
|
|
</paragraph>
|
|
</subSubSection>
|
|
<subSubSection id="B9094998D226E7A272E446B68178A1A0" pageId="0" pageNumber="117" type="description">
|
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<paragraph id="4891A0A9380D00E3577686E5404B0776" pageId="0" pageNumber="117">Description.</paragraph>
|
|
<paragraph id="A5831623CB194A8F33F222627A99F82A" pageId="0" pageNumber="117">Although the available sample consists of more than 250 specimens, including more than 150 granicones, only a small number of specimens provide useful anatomical information. This description is therefore focused on the most informative specimens.</paragraph>
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<paragraph id="F4AC5B9FA520A465B59B9236412642F2" pageId="0" pageNumber="117">
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|
<emphasis id="84B6124024F201C7C61D48EC3AC53320" bold="true" pageId="0" pageNumber="117">
|
|
<emphasis id="F2B419D56E374D882C6806E12DD7BDD5" italics="true" pageId="0" pageNumber="117">Surface texture</emphasis>
|
|
.
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|
</emphasis>
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The surface of all available shell material and granicones is evenly covered by distinct tubercles (Figs
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<figureCitation id="ED0809D13F0CD863A880554F7DE7695D" captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Shell remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. Photographs of select specimens highlighting variation in preservation. (a) WMNM P 88818, a right epiplastron (also see Fig. 5 b); (b) WMNM P 88824, a right partial hypoplastron (also see Fig. 5 k); (c) WMNM P 88823, a partial left hypo- and xiphiplastron (also see Fig. 5 i); (d) WMNM P 88808, a partial, anterior neural (also see Fig. 3 a); (e) WMNM P 88813, a partial nuchal (also see Fig. 4 b); (f) WMNM P 88816, a partial left peripheral XI (also see Fig. 4 h). All specimens are to scale." figureDoi="10.3897/fr.26.102128.figure2" httpUri="https://binary.pensoft.net/fig/858747" pageId="0" pageNumber="117">2</figureCitation>
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-
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|
<figureCitation id="5AA92253B972FF1D66EDD863C3251A51" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5</figureCitation>
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|
). Each tubercle is columnar and clearly separated from neighboring tubercles. The tubercles of the shell easily dislocate at their base, where they leave behind a circular scar, which can easily be recognized under the microscope. Although some variation is apparent across the shell (e.g., tubercles are less densely spaced over the neurals), most are about 0.5 to 1 mm wide, about 0.5 to 1.5 mm tall, and separated from neighboring tubercles by about 1.5 to 2 mm. The tubercles are less distinct on the available cranial fragment and all granicones, but similar in dimensions. In all regards, the surface texture closely approximates to that of
|
|
<taxonomicName id="7A72FA3C95F1BD1D27EB294A93FDEE1C" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
|
|
<emphasis id="A414C9D8A7ED85B3D3B01308DEB3870E" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
|
|
</taxonomicName>
|
|
material from the Wessex Formation of England (NHMUK R171, R5195-5199, R5201, R5205, R5238).
|
|
</paragraph>
|
|
<paragraph id="B673220A5E18F9C7E8145287E196EEE1" pageId="0" pageNumber="117">
|
|
<emphasis id="7AA5BDCD2A2CF61908C00A7191D0FAB5" bold="true" pageId="0" pageNumber="117">
|
|
<emphasis id="FEB2DD878EA05BE25742530BF3E6102D" italics="true" pageId="0" pageNumber="117">Carapace</emphasis>
|
|
.
|
|
</emphasis>
|
|
The available material (Figs
|
|
<figureCitation id="185D4F9637E8A0C552DCF81230431A3E" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3</figureCitation>
|
|
,
|
|
<figureCitation id="55ACA72BD154E09DF743F20F0234E2EC" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4</figureCitation>
|
|
) suggests that the carapace had a length of up to 70 cm (see Costals below), a nuchal notch extending laterally up to peripheral II (see Nuchal below), and that deep grooves decorated the neural-costal contacts along the posterior half of the shell (see Costals and Neurals below). The carapace is 5 to 6 mm thin along the peripherals and costals, but thickens to 8 mm at the neurals. All parts of the carapace are evenly covered by distinct tubercles that disarticulate easily (Figs
|
|
<figureCitation id="EF1D057E57688E7B87BECB2F27E7F87D" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3</figureCitation>
|
|
,
|
|
<figureCitation id="BF2AB46DABAA5DA8B3B3B3DDB5B0771F" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4</figureCitation>
|
|
).
|
|
</paragraph>
|
|
<caption id="C8A7C102B0FE9C9CADE22450F7FD4D15" doi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117" start="Figure 3" startId="F3">
|
|
<paragraph id="EC213BD1C70362A8483F028F21706644" pageId="0" pageNumber="117">
|
|
<emphasis id="EE622E4E138E0F9AA43B5606B3DBDDB8" bold="true" pageId="0" pageNumber="117">Figure 3.</emphasis>
|
|
Neural and costal remains of
|
|
<taxonomicName id="3BDA0614DC7A2D0CFDCBB3B8D41A2B16" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
|
|
<emphasis id="BD99D668DC2F58BC0972D88019AB1B55" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
|
|
</taxonomicName>
|
|
from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3D models. (
|
|
<emphasis id="0EB324837216501A5CC7DDCB52E1B64C" bold="true" pageId="0" pageNumber="117">a</emphasis>
|
|
) WMNM P88808, a partial, anterior neural; (
|
|
<emphasis id="1BF5553EB7FED9A9CC49E5E126CD1845" bold="true" pageId="0" pageNumber="117">b</emphasis>
|
|
) WMNM P49476, a partial, posterior neural; (
|
|
<emphasis id="B11CDEA6B91DA9215160E43834726A54" bold="true" pageId="0" pageNumber="117">c</emphasis>
|
|
) WMNM P49351, a partial, posterior neural; (
|
|
<emphasis id="02DAEC7BC1798016C8FB12323C96DF39" bold="true" pageId="0" pageNumber="117">d</emphasis>
|
|
) WMNM P88809, a partial, posterior neural; (
|
|
<emphasis id="A379AC7B7D252DD8B46E2932AA75822C" bold="true" pageId="0" pageNumber="117">e</emphasis>
|
|
) WMNM P88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (
|
|
<emphasis id="38F2E11C462EF97CACEADB3C4FA8B484" bold="true" pageId="0" pageNumber="117">f</emphasis>
|
|
) WMNM P88810, a partial left costal I; (
|
|
<emphasis id="129680013523CB96E362FF65EB55574A" bold="true" pageId="0" pageNumber="117">g</emphasis>
|
|
) WMNM P88812, a partial costal; (
|
|
<emphasis id="C8E27511CAD1D341B61C1514E2D64381" bold="true" pageId="0" pageNumber="117">h</emphasis>
|
|
) WMNM P88811, a partial costal; (
|
|
<emphasis id="193D390883188236A0417B2E5F5F9800" bold="true" pageId="0" pageNumber="117">i</emphasis>
|
|
) WMNM P49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale.
|
|
<emphasis id="DEB108E801F1A4182B327A7F39529019" italics="true" pageId="0" pageNumber="117">Abbreviations</emphasis>
|
|
: co = costal; ne = neural.
|
|
</paragraph>
|
|
</caption>
|
|
<caption id="29F6123DF4F195698E4B1E722BDAA425" doi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117" start="Figure 4" startId="F4">
|
|
<paragraph id="5FE931B2D4EF0539146D0D15B7420A81" pageId="0" pageNumber="117">
|
|
<emphasis id="56B12B174CBE33B35CE5DA99EB274EE2" bold="true" pageId="0" pageNumber="117">Figure 4.</emphasis>
|
|
Nuchal and peripheral remains of
|
|
<taxonomicName id="FE68EB0D6D519FFCE4FA366B6F5C6922" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
|
|
<emphasis id="6CBBD2D08AC857FE93E08480E4F99853" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
|
|
</taxonomicName>
|
|
from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3D models. (
|
|
<emphasis id="B78BD47570336BE7E0D942DF51480BF8" bold="true" pageId="0" pageNumber="117">a</emphasis>
|
|
) WMNM P88813, a partial nuchal; (
|
|
<emphasis id="C0B440612930A00D1629BADD01CD5F0E" bold="true" pageId="0" pageNumber="117">b</emphasis>
|
|
) WMNM P88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (
|
|
<emphasis id="7BD52282C425EF8095E3A56E77FD4C40" bold="true" pageId="0" pageNumber="117">c</emphasis>
|
|
) WMNM P88815, a partial right peripheral I; (
|
|
<emphasis id="7B7DC920FF49AEA5BC1CB05CF18974C5" bold="true" pageId="0" pageNumber="117">d</emphasis>
|
|
) WMNM P49051, a partial bridge peripheral; (
|
|
<emphasis id="D6D6846DB202911CE2122FA0666390B6" bold="true" pageId="0" pageNumber="117">e</emphasis>
|
|
) WMNM P88817, a partial bridge peripheral; (
|
|
<emphasis id="7446A2BCFB287E63C5ED2A1924FC7099" bold="true" pageId="0" pageNumber="117">f</emphasis>
|
|
) WMNM P41690, a partial right peripheral XI; (
|
|
<emphasis id="20E12F681C32FCD554B01D404DA4FAB3" bold="true" pageId="0" pageNumber="117">g</emphasis>
|
|
) WMNM P47259, a partial peripheral; (
|
|
<emphasis id="5A68C765250952EC0EDEB8CB74832F17" bold="true" pageId="0" pageNumber="117">h</emphasis>
|
|
) WMNM P88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale.
|
|
<emphasis id="E0DB74C475B4E6C7237915D266E4E99C" italics="true" pageId="0" pageNumber="117">Abbreviations</emphasis>
|
|
: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale.
|
|
</paragraph>
|
|
</caption>
|
|
<paragraph id="FBF5A8ABE41E9452683736E82DCB0D36" pageId="0" pageNumber="117">
|
|
<emphasis id="E00D198005ADAA988926F137ED130060" bold="true" pageId="0" pageNumber="117">
|
|
<emphasis id="C53A8147B565011C2527B8992B8BE67E" italics="true" pageId="0" pageNumber="117">Neurals</emphasis>
|
|
.
|
|
</emphasis>
|
|
We are able to identify six elements as neurals: WMNM P88808 (Fig.
|
|
<figureCitation id="12A8F68170CF1E420FE02B3165CDCE83" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3a</figureCitation>
|
|
), P49476 (Fig.
|
|
<figureCitation id="7785DFFE8EDEB81B9D1BA3108ACE232C" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3b</figureCitation>
|
|
), P49351 (Fig.
|
|
<figureCitation id="E9FC526EA591F12AACB286D1265205A2" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3c</figureCitation>
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), P88809 (Fig.
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<figureCitation id="4C11FCB0AC0792F5297F8C4B8AD5DF5D" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3h</figureCitation>
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), and P88807 (Fig.
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<figureCitation id="E565A627883B306DD0E7A2C4418B6486" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3e</figureCitation>
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). WMNM P88807 is a larger shell fragment that consists of two neural elements in addition to costal remains including two neural elements that are up to 8 mm thick (Fig.
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<figureCitation id="F82FA888C0B68A8263FFC43264877443" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3e</figureCitation>
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). The posteriorly angled orientation of the associated rib heads relative to the neural column suggest that the elements represent neurals V, VI, or VII. Their outline is hexagonal with short anterolateral sides. The anterior of the two neural shows faint traces of a midline keel, as has also been reported for neurals of
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<taxonomicName id="47A3EA4ECF441DE1AA32A81B267FEC54" authorityName="Hay" authorityYear="1908" class="Reptilia" family="Pleurosternidae" genus="Naomichelys" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Naomichelys speciosa" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="species" species="speciosa">
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<emphasis id="76EEBCF2FBC9C97CD6627F894C2EC0AF" italics="true" pageId="0" pageNumber="117">Naomichelys speciosa</emphasis>
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</taxonomicName>
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(
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<bibRefCitation id="6F8E06EB02092BDCE65070FC26B0CF89" author="Joyce, WG" journalOrPublisher="Special Papers in Paleontology" pageId="0" pageNumber="117" pagination="75 - 97" refId="B15" refString="Joyce, WG, Chapman, SD, Moody, RTJ, Walker, CA, 2011. The skull of the solemydid turtle Helochelydra nopcsai from the Early Cretaceous (Barremian) of the Isle of Wight (UK) and a review of Solemydidae. Special Papers in Paleontology 86: 75 - 97" title="The skull of the solemydid turtle Helochelydra nopcsai from the Early Cretaceous (Barremian) of the Isle of Wight (UK) and a review of Solemydidae." volume="86" year="2011">Joyce et al. 2011</bibRefCitation>
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). WMNM P88808 (Fig.
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<figureCitation id="114A7DD510393A0407FFF24FBA4CD471" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3a</figureCitation>
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) is the largest available neural, flat, about 5 mm thick, and has a hexagonal outline (i.e., contacts with six surrounding bones are apparent). Its dorsal surface is badly eroded: there are faint traces of an intervertebral sulcus, but only the bases remain of the surface tubercles. The orientation of this element is not clear, but its size suggests that it originates from the anterior half of the shell. WMNM P49351 (Fig.
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<figureCitation id="799224AABBAC64CA9BD235FA870E9439" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3c</figureCitation>
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) is only half as long anteroposteriorly as the previous element, but about 7 mm thick. A sharp median depression crosses its surface. WMNM P49476 (Fig.
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<figureCitation id="1B39E62CC9F85B203FEAAB98962333CA" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3b</figureCitation>
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) has similar dimensions to WMNM P49351, but it is only about 4 mm thick and flat. The equilateral, hexagonal outline of the last two elements suggests an origin from the back of the shell. WMNM P88809 (Fig.
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<figureCitation id="42BB2C593FA5DFCAAE0F560AAA9B72EE" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3d</figureCitation>
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) is the smallest available neural and only about 2 mm thin. Only the right two thirds are preserved. Its small size and equilateral, hexagonal outline suggest that it originates from the posterior portions of the shell of a juvenile.
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</paragraph>
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<paragraph id="08118F43FFE53A5181BA76A3965EBA68" pageId="0" pageNumber="117">
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<emphasis id="0F7154850E932C30E8356FE4BF77A463" bold="true" pageId="0" pageNumber="117">
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<emphasis id="A62C63D24BE5F78212C80A6A88B7CFA9" italics="true" pageId="0" pageNumber="117">Costals</emphasis>
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.
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</emphasis>
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A number of elements can be diagnosed as costals, of which the following seven are the most informative: WMNM P88807 (Fig.
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<figureCitation id="6850A35D52709CC9961038F6F5E3AA4B" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3e</figureCitation>
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), a shell fragment that includes remains of three costals, WMNM P88810, a partial left costal I (Fig.
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<figureCitation id="E2B78520D942D59C5CFC72A8F0ED04BC" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3f</figureCitation>
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), P88811 (Fig.
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<figureCitation id="94B2EC4B9D427A527907BCD40AECAA7F" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3h</figureCitation>
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) and P88812 (Fig.
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<figureCitation id="4B76784B5E24F0F9E4A7881970D5F1EC" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3g</figureCitation>
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), two large costal fragments, and P49470, the median portion of a midsized costal (Fig.
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<figureCitation id="10F1B13DEC93FC304012968B112730AF" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3i</figureCitation>
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). The dermal portion of these costals is about 4 mm thick. The ribs, when present, are broad, about a quarter of the anteroposterior width of the costal, but only add about 2-3 mm to the thickness of each element. The three costal remains preserved in WMNM P88807 (Fig.
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<figureCitation id="CEBF87C3A05EC0A6ABCD3B249D0546F8" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3e</figureCitation>
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) are in articulation with two neurals and, therefore, best document the medial aspect of these elements. The slight posteriorly angled orientation of the ribs suggest that they originate from the mid-posterior part of the shell. All elements have two contacts with the adjacent neurals. The free, medial portion of one costal rib is well preserved. It is relatively narrow, about 2 mm across, and delimits a relatively low costovertebral tunnel. WMNM P88810 (Fig.
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<figureCitation id="55581885CED0206E27A364D62482FF98" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3f</figureCitation>
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) can be identified as a partial left costal I by the presence of the articular scar for dorsal rib I. The sulcus between vertebral I and pleural I diagonally traverses the dorsal surface of this element. A rounded anteromedial facet for articulation with the nuchal suggests that the nuchal deeply inserted between the anterior costals. WMNM P88811 (Fig.
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<figureCitation id="AEACA9DC02889B5947F311DD8B0BD766" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3h</figureCitation>
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) is significant, as it possesses intact anterior and posterior margins. The greatest anteroposterior
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<normalizedToken id="26372B3FAC9C830D4762EC451E19B548" originalValue="“width”">"width"</normalizedToken>
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of this element is 56 mm, which is a little greater than the greatest costal width that can be observed for the only known complete skeleton of the helochelydrid
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<taxonomicName id="DC787D8FD676B9392F4A7A78A63FABE1" authorityName="Hay" authorityYear="1908" class="Reptilia" family="Pleurosternidae" genus="Naomichelys" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Naomichelys speciosa" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="species" species="speciosa">
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<emphasis id="82D26866C767BBFEA685FA3CF62DDABC" italics="true" pageId="0" pageNumber="117">Naomichelys speciosa</emphasis>
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</taxonomicName>
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(
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<bibRefCitation id="A383CF6D0B51D2AA38FB53465B5AD59A" DOI="https://doi.org/10.1666/14-002" author="Joyce, WG" journalOrPublisher="Journal of Paleontology" pageId="0" pageNumber="117" pagination="1257 - 1287" refId="B16" refString="Joyce, WG, Sterli, J, Chapman, SD, 2014. The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas. Journal of Paleontology 88: 1257 - 1287, DOI: https://doi.org/10.1666/14-002" title="The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas." url="https://doi.org/10.1666/14-002" volume="88" year="2014">Joyce et al. 2014</bibRefCitation>
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). Assuming similar proportions, the full carapace may therefore have reached a length of over 70 cm. WMNM P49470 (Fig.
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<figureCitation id="3807503943118230D7524801517960A1" captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Neural and costal remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88808, a partial, anterior neural; (b) WMNM P 49476, a partial, posterior neural; (c) WMNM P 49351, a partial, posterior neural; (d) WMNM P 88809, a partial, posterior neural; (e) WMNM P 88807, a carapacial fragment consisting of two neurals and remnants of three adjacent costals; (f) WMNM P 88810, a partial left costal I; (g) WMNM P 88812, a partial costal; (h) WMNM P 88811, a partial costal; (i) WMNM P 49470, a partial right costal. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: co = costal; ne = neural." figureDoi="10.3897/fr.26.102128.figure3" httpUri="https://binary.pensoft.net/fig/858748" pageId="0" pageNumber="117">3i</figureCitation>
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) only has an anteroposterior
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<normalizedToken id="14EDFE1080F1BE553F71EAF7D0118DA7" originalValue="“width”">"width"</normalizedToken>
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of 3 cm and, therefore, likely originates from the back of the shell of a smaller individual. It is notable for exhibiting a concavity just above the rib head, near its medial articulation with the neurals. Together with the grooved neurals, this suggests some meso-scale sculpturing of the shell surface towards the back of the shell.
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</paragraph>
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<paragraph id="29594D9E1FAD8421CAFB8A583BA722F7" pageId="0" pageNumber="117">
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<emphasis id="813E04E6602C1B50CB2DE937808AB264" bold="true" pageId="0" pageNumber="117">
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<emphasis id="2EBAB110AC98D3F8758C82D4C42E46E7" italics="true" pageId="0" pageNumber="117">Nuchal</emphasis>
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.
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</emphasis>
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Two specimens preserve remains of the nuchal: WMNM P88813, the right half of a large nuchal (Fig.
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<figureCitation id="AF6FFB0B521A508F5F2D987702918101" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4a</figureCitation>
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), and WMNM P88814, a smaller nuchal fragment, which remains articulated with remnants of the adjacent right costal I and peripherals I and II (Fig.
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<figureCitation id="E312C83CE20444B0D78459F79B920AB8" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4b</figureCitation>
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). The two specimens suggest that a broad nuchal notch was present that was framed by peripherals II, as is typical for helochelydrids (
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<bibRefCitation id="B76418C114088074661AB0971C4188C5" DOI="https://doi.org/10.1666/14-002" author="Joyce, WG" journalOrPublisher="Journal of Paleontology" pageId="0" pageNumber="117" pagination="1257 - 1287" refId="B16" refString="Joyce, WG, Sterli, J, Chapman, SD, 2014. The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas. Journal of Paleontology 88: 1257 - 1287, DOI: https://doi.org/10.1666/14-002" title="The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas." url="https://doi.org/10.1666/14-002" volume="88" year="2014">Joyce et al. 2014</bibRefCitation>
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;
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<bibRefCitation id="1A1D953C3D4326C77F45A31F56BBCDAB" DOI="https://doi.org/10.1016/j.cretres.2019.104291" author="Perez-Garcia, A" journalOrPublisher="Cretaceous Research" pageId="0" pageNumber="117" refId="B35" refString="Perez-Garcia, A, Espilez, E, Mampel, L, Alcala, L, 2020a. A new basal turtle represented by the two most complete skeletons of Helochelydridae in Europe. Cretaceous Research 107: 104291. https://doi.org/10.1016/j.cretres.2019.104291" title="A new basal turtle represented by the two most complete skeletons of Helochelydridae in Europe. Cretaceous Research 107: 104291." url="https://doi.org/10.1016/j.cretres.2019.104291" year="2020 a">
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<normalizedToken id="60388AC71918B803E6443DB8EA3FA98E" originalValue="Pérez-García">Perez-Garcia</normalizedToken>
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et al. 2020a
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</bibRefCitation>
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). The oblique anterolateral contact of the nuchal with peripheral I is oriented more to the anterior than the side. There is no evidence for costiform processes. The right portion of a cervical scute is preserved in WMNM P88813 (Fig.
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<figureCitation id="059FE22086F62996873D875187200498" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4a</figureCitation>
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). As this specimen broke near the midline, we estimate that the complete cervical was at least twice as wide than long. Lateral to the cervical scute, a sulcus is apparent that delimits the contact between marginal I and vertebral I. There is no evidence for the presence of a prepleural on the nuchal. The skin-scute sulcus runs on the underside of the nuchal just behind the anterior margin of the bone.
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</paragraph>
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<paragraph id="8C7FCCD0D0FDEED3FE45D725D94C964C" pageId="0" pageNumber="117">
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<emphasis id="7F23ECF9D57091BE9BC52A65A05ADE7A" bold="true" pageId="0" pageNumber="117">
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<emphasis id="A609630AB3F38BE6318D8791F6236449" italics="true" pageId="0" pageNumber="117">Peripherals</emphasis>
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.
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</emphasis>
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A number of peripherals are preserved, of which we present the six most informative. WMNM P88814 includes a nearly complete right peripheral I articulated with remnants of the adjacent element (Fig.
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<figureCitation id="927AF2AE84A3B1CC6B9EF92A46342CFB" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4b</figureCitation>
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). WMNM P88815 is a first peripheral I as well, but lacks much of its dorsal surface (Fig.
|
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<figureCitation id="B6B7D69BF3A5572FE42B16C27CF9EBD0" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4c</figureCitation>
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). These two elements document that peripheral series articulated with the nuchal laterally along an oblique (WMNM P88814) to straight (WMNM P88815) suture, costal I posteriorly, and peripheral II laterally. The scute-skin sulcus runs diagonally across the underside of the peripheral, thereby bridging the location of this sulcus near the margin of the shell at the nuchal with the deep location on the second peripheral. Although damaged, these bones also document that the visceral cavity extends all the way from the bridge region to the medial fifth of peripheral I, as had previously been documented for other helochelydrids (
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<bibRefCitation id="CDEA51362FC71F568EBCFB7E311246BB" DOI="https://doi.org/10.1666/14-002" author="Joyce, WG" journalOrPublisher="Journal of Paleontology" pageId="0" pageNumber="117" pagination="1257 - 1287" refId="B16" refString="Joyce, WG, Sterli, J, Chapman, SD, 2014. The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas. Journal of Paleontology 88: 1257 - 1287, DOI: https://doi.org/10.1666/14-002" title="The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas." url="https://doi.org/10.1666/14-002" volume="88" year="2014">Joyce et al, 2014</bibRefCitation>
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;
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<normalizedToken id="9BE5BB21D8D24A3F7C852834235D7199" originalValue="Pérez-Garcia">Perez-Garcia</normalizedToken>
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et al, 2020a). WMNM P88816 is a relatively large peripheral element that we tentatively interpret as the right peripheral XI, as it displays a small, but elongate facet on its visceral side best interpreted by reference to
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<taxonomicName id="AFE00C3F2AD2ED1977D9171DD509F4F1" authorityName="Hay" authorityYear="1908" class="Reptilia" family="Pleurosternidae" genus="Naomichelys" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Naomichelys speciosa" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="species" species="speciosa">
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<emphasis id="9B602BC5A7BA75E53095F5E7220E2372" italics="true" pageId="0" pageNumber="117">Naomichelys speciosa</emphasis>
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</taxonomicName>
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(
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<bibRefCitation id="36529EE980F7B1E676BC18501B266615" DOI="https://doi.org/10.1666/14-002" author="Joyce, WG" journalOrPublisher="Journal of Paleontology" pageId="0" pageNumber="117" pagination="1257 - 1287" refId="B16" refString="Joyce, WG, Sterli, J, Chapman, SD, 2014. The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas. Journal of Paleontology 88: 1257 - 1287, DOI: https://doi.org/10.1666/14-002" title="The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas." url="https://doi.org/10.1666/14-002" volume="88" year="2014">Joyce et al. 2014</bibRefCitation>
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) as the articulation site for thoracic rib IX (Fig.
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<figureCitation id="73B932FD47FB6FD37788232C70C300A6" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4h</figureCitation>
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). This element shows faint traces of an intermarginal scute sulcus and documents that the visceral cavity extends posteriorly from the bridge region to reach the pygal. The visceral cavity of
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<taxonomicName id="69CB02E291005274828691D625A71A4D" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
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<emphasis id="449A62B886D47AA8B45BDCB6A4791E47" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
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</taxonomicName>
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, therefore, extended all the way from the medial fifth of peripheral I to the pygal, much as in
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<taxonomicName id="33DC476BFDA7F86E9EB3273DA8008DAC" genus="Aragochersis" lsidName="Aragochersis lignitesta" pageId="0" pageNumber="117" rank="species" species="lignitesta">
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<emphasis id="3E0529EEA31C39898C74EDA5703606F8" italics="true" pageId="0" pageNumber="117">Aragochersis lignitesta</emphasis>
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</taxonomicName>
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(
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<normalizedToken id="E103529CE45AFB6043CD5C6CA51B9425" originalValue="Pérez-Garcia">Perez-Garcia</normalizedToken>
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et al, 2020a), but also
|
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<taxonomicName id="C6F7B2B126FB61C2E435E1089B632818" baseAuthorityName="Owen" baseAuthorityYear="1842" class="Reptilia" family="Pleurosternidae" genus="Pleurosternon" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Pleurosternon bullockii" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="species" species="bullockii">
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<emphasis id="0D502C24204A73CFAE27BE762EB2BCFA" italics="true" pageId="0" pageNumber="117">Pleurosternon bullockii</emphasis>
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</taxonomicName>
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(
|
|
<bibRefCitation id="33DF3708242D79866ED17DAF968DE99B" DOI="https://doi.org/10.3897/fr.25.85334" author="Joyce, WG" journalOrPublisher="Fossil Record" pageId="0" pageNumber="117" pagination="263 - 274" refId="B14" refString="Joyce, WG, Bourque, JR, Fernandez, V, Rollot, Y, 2022. An alternative interpretation of small bodied turtles from the "Middle Purbeck" of England as a new species of compsemydid turtle. Fossil Record 25: 263 - 274, DOI: https://doi.org/10.3897/fr.25.85334" title="An alternative interpretation of small bodied turtles from the " Middle Purbeck " of England as a new species of compsemydid turtle." url="https://doi.org/10.3897/fr.25.85334" volume="25" year="2022">Joyce et al. 2022</bibRefCitation>
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). However, three finger-like pockets within the cavity of WMNM P88816 suggests that the visceral cavity is not developed evenly across the shell (see arrows in Fig.
|
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<figureCitation id="82E065B0EFF9912AEBD470CD70696B95" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4h</figureCitation>
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). The waning visceral cavity combined with minor asymmetries suggest that WMNM P41690 is a damaged right peripheral X as well (Fig.
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<figureCitation id="B1A33EB0B511C0C4A1ED5DF5128E6DB9" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4f</figureCitation>
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). As the most anterior and most posterior peripherals show evidence of a visceral cavity, it is not possible to identify the serial identity of the damaged, remaining peripheral elements. Three notable remains, WMNM P47259 (Fig.
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<figureCitation id="650C986803666AED2CB7D962E0DAAFE5" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4g</figureCitation>
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), P49051 (Fig.
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<figureCitation id="ABD4F3E321983F5B493EC9418171B28A" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4d</figureCitation>
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), and P88817 (Fig.
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<figureCitation id="E96EC78B87C03500484BB221FD3980C8" captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Nuchal and peripheral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88813, a partial nuchal; (b) WMNM P 88814, a carapacial fragment consisting of remnants of a nuchal, rights peripheral I, remnants of right peripheral II, and remnants of right costal I; (c) WMNM P 88815, a partial right peripheral I; (d) WMNM P 49051, a partial bridge peripheral; (e) WMNM P 88817, a partial bridge peripheral; (f) WMNM P 41690, a partial right peripheral XI; (g) WMNM P 47259, a partial peripheral; (h) WMNM P 88816, a partial left peripheral XI. Anatomical direction are placed in quotes for elements with uncertain orientation. With the exception of the illustrations, all specimens are to scale. Abbreviations: CE = cervical scale; co = costal; MA = marginal scale; nu = nuchal; pe = peripheral; VE = vertebral scale." figureDoi="10.3897/fr.26.102128.figure4" httpUri="https://binary.pensoft.net/fig/858749" pageId="0" pageNumber="117">4e</figureCitation>
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) nevertheless document variable development of a low gutter. WMNM P49051 and P88817 furthermore highlight that the dorsal and ventral branches that make up each peripheral are thin-walled and meet each other just medial to the edge of the carapace. The peripherals of the bridge region are therefore extremely fragile.
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</paragraph>
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<paragraph id="2C35D5A4B63C0561F0F67CE12A23369E" pageId="0" pageNumber="117">
|
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<emphasis id="2B95EC913B7BCC03D2DAB23345C6012D" bold="true" pageId="0" pageNumber="117">
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<emphasis id="3A516D1A7CB734F4CA90CA2F3C076349" italics="true" pageId="0" pageNumber="117">Epiplastron</emphasis>
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.
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</emphasis>
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The only available epiplastron, WMNM P88818, is near complete, but mineral precipitations on the external surface partially conceal its sulci (Fig.
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|
<figureCitation id="301979497F2635D1666AD87B12C3DCB1" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5b</figureCitation>
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). The epiplastron formed the anterior aspects of the transversely oriented anterior plastral lobe. The epiplastron has a convex posteromedial contact with the entoplastron. The epiplastron hereby slightly underlaps the entoplastron, which, as a result, would have appeared smaller in ventral than in dorsal view. An equally broad contact is present posterolaterally with the hyoplastron. Though damaged, deep pockets along the posterolateral margin of the epiplastron suggest that at least two peg-like processes of the hyoplastron inserted into the epiplastron. A short median contact is furthermore apparent with its counterpart. A notch along the anterior margin of the epiplastron signifies the location of the gular-extragular sulcus. A faint trace of this sulcus can be gleaned on the ventral surface that runs posteriorly from the notch (Fig.
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<figureCitation id="AD09BC81800EA4D1EC8D900858BBAEBB" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5b</figureCitation>
|
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). A similar notch for the extragular-humeral sulcus is apparent on the lateral margin of the epiplastron (Fig.
|
|
<figureCitation id="20DB53F8A3BF21A2FED45365432D5D58" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5b</figureCitation>
|
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). On the dorsal side, the extragular-humeral sulcus veers posteriorly to cross the lateral side of the epiplastron (see arrows in Fig.
|
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<figureCitation id="AECA7CB3646DAC3E3C2BC92C5AF2B79E" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5b</figureCitation>
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). Its posterior end is not preserved, as this was likely located on the hyoplastron. As a result of this arrangement, the extragular covered only part of the epiplastron in ventral view, but most of it in dorsal view. The dorsal side of the epiplastron is characterized by a thickened epiplastral lip.
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</paragraph>
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<caption id="09841A5CEFD6F1326600B083C764C334" doi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117" start="Figure 5" startId="F5">
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<paragraph id="5E7F5298B358F5F0CA4B361236F7A754" pageId="0" pageNumber="117">
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<emphasis id="1C14463BF113652573601BF421E6A3C2" bold="true" pageId="0" pageNumber="117">Figure 5.</emphasis>
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Plastral remains of
|
|
<taxonomicName id="79F4E7938D70C20629721BC203FF6375" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
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<emphasis id="8F49A3D260AA51C18A9FBC4194CD4704" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
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</taxonomicName>
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from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3D models. (
|
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<emphasis id="B1BCCB4415CB35EFE9934D6B211E911F" bold="true" pageId="0" pageNumber="117">a</emphasis>
|
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) WMNM P88819, a partial entoplastron; (
|
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<emphasis id="3E07177E170FF2A3E1FE51ECAC7C573F" bold="true" pageId="0" pageNumber="117">b</emphasis>
|
|
) WMNM P88818, a right epiplastron; (
|
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<emphasis id="BEE4C7B180EBF4CE3599509916BF9FC1" bold="true" pageId="0" pageNumber="117">c</emphasis>
|
|
) WMNM P48296, a partial entoplastron; (
|
|
<emphasis id="5206525ABF883654B5AA544FFDFCF0C1" bold="true" pageId="0" pageNumber="117">d</emphasis>
|
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) WMNM P47721, a partial right hyoplastron; (
|
|
<emphasis id="423E6CF6BCF0E77AB4ABDA3D1C562B2F" bold="true" pageId="0" pageNumber="117">e</emphasis>
|
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) WMNM P88820, a partial left hyoplastron; (
|
|
<emphasis id="E20F248C63FA3ED24ECEAA5A24BE3602" bold="true" pageId="0" pageNumber="117">f</emphasis>
|
|
) WMNM P48226, a plastral fragment; (
|
|
<emphasis id="02CFADA985CDC23F2085E785D08FF3EF" bold="true" pageId="0" pageNumber="117">g</emphasis>
|
|
) WMNM P88822, a plastral fragment; (
|
|
<emphasis id="260E16CD3DD7C0FA5B73F067DD86A69B" bold="true" pageId="0" pageNumber="117">h</emphasis>
|
|
) WMNM P88821, a plastral fragment; (
|
|
<emphasis id="B65DA680D1531EA30D77EC8870C98E7D" bold="true" pageId="0" pageNumber="117">i</emphasis>
|
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) WMNM P88823, a partial left hypo- and xiphiplastron; (
|
|
<emphasis id="6BB3D49FD0B0BFAD78AE2B2270FE8ADA" bold="true" pageId="0" pageNumber="117">j</emphasis>
|
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) WMNM P88825, a partial right hypoplastron; (
|
|
<emphasis id="7F027CA2F282DFB48EAACBAEEEA5B98F" bold="true" pageId="0" pageNumber="117">k</emphasis>
|
|
) WMNM P88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale.
|
|
<emphasis id="77BD1CCCB8C85AA0FC36823B9F329AA5" italics="true" pageId="0" pageNumber="117">Abbreviations</emphasis>
|
|
: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron.
|
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</paragraph>
|
|
</caption>
|
|
<paragraph id="2D6CB4A172AA798EB659ABF4C4A707D4" pageId="0" pageNumber="117">
|
|
<emphasis id="0E771B0E960824E8A481528298996467" bold="true" pageId="0" pageNumber="117">
|
|
<emphasis id="DCD97310BD6B7530D6FE2F1976EDC644" italics="true" pageId="0" pageNumber="117">Entoplastron</emphasis>
|
|
.
|
|
</emphasis>
|
|
Two partial entoplastra are available, WMNM P48296 (Fig.
|
|
<figureCitation id="D868BBDDD0437C51B441386491B57CF4" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5a</figureCitation>
|
|
) and P88819 (Fig.
|
|
<figureCitation id="BDC557C5FC7BC990AA73C8AAECF747EB" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5c</figureCitation>
|
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), of which the former is better preserved. The posterior half of the entoplastron is lacking in either specimen, but comparison with
|
|
<taxonomicName id="2443E4B6C04AE2F3C40836F91187003A" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
|
|
<emphasis id="1841CB5F31C99A6C3E69F3CABE48C17F" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
|
|
</taxonomicName>
|
|
(NHMUK R171) suggests that the entoplastron was slightly longer than wide. The entoplastron slightly overlaps the epiplastron along its slightly convex anterolateral contact. The posterolateral contact, as far as preserved, broadly underlapped the hyoplastron. Traces of the embryonic interclavicle are apparent on the dorsal side of both available elements, much as in other helochelydrids. The gulars only cover a small triangular surface on the ventral side of WMNM P48296, but about a third of the ventral surface of P88819. The extragulars minutely overlapped both elements. Only a minute entoplastral scute is apparent in WMNM P48296. An entoplastral scute seems to be absent in WMNM P88819, in which the humeral-gular junction is just about preserved.
|
|
</paragraph>
|
|
<paragraph id="EAE8D5122442DEE2A1F32D38DD5E3EDD" pageId="0" pageNumber="117">
|
|
<emphasis id="9C836CA3CF17584D774B0BD37341501A" bold="true" pageId="0" pageNumber="117">
|
|
<emphasis id="BFBC0C59E234B9DA289516E5768B5D87" italics="true" pageId="0" pageNumber="117">Hyoplastron</emphasis>
|
|
.
|
|
</emphasis>
|
|
Two partial hypoplastral remains can be identified with confidence. WMNM P47721 (Fig.
|
|
<figureCitation id="A1B4AB2861D5552C36FF53D24E179C9F" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5d</figureCitation>
|
|
), the more complete of the two, documents the lateral margin of the anterior plastral lobe, just posterior to the articulation of the hyoplastron with the epiplastron, while WMNM P88820 (Fig.
|
|
<figureCitation id="82BB885D82E80B483F40F6DEABFB85D6" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5e</figureCitation>
|
|
) documents a segment slightly further to the back. The two fragments combined indicate that the skin-scute sulcus is barely located on the visceral side of the hyoplastron, but converges with its margin towards the back. Indeed, the sulcus terminates in WMNM P88820 near the margin, which suggests that this element broke at the humeral-pectoral sulcus. The resulting lip is only lightly developed towards the back, as seen in WMNM P88820, but becomes pronounced anteriorly to form a finger-like process that inserted anteromedially into the epiplastron, as seen in WMNM P47721.
|
|
</paragraph>
|
|
<paragraph id="DA03483943DE3B6D49B3A29C8D1273F7" pageId="0" pageNumber="117">
|
|
Two additional, fragmentary specimens, WMNM P48226 (Fig.
|
|
<figureCitation id="39E6AFA2C779D3D27BAB57571CF2C861" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5f</figureCitation>
|
|
) and P88821 (Fig.
|
|
<figureCitation id="A2CFB52957ABD95A8EA1AABA6BF3E862" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5h</figureCitation>
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|
), exhibit radiating, pin-like tubercles on their visceral sides that are reminiscent of those found on the hyoplastron of
|
|
<taxonomicName id="791A5B2B7F96E23628FABEE000913345" authorityName="Hay" authorityYear="1908" class="Reptilia" family="Pleurosternidae" genus="Naomichelys" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Naomichelys speciosa" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="species" species="speciosa">
|
|
<emphasis id="0A0D63F0DFED8B839D2FCA7E224FFC0A" italics="true" pageId="0" pageNumber="117">Naomichelys speciosa</emphasis>
|
|
</taxonomicName>
|
|
, but they may also represent remains of the mesoplastra or hypoplastra. A third fragment, WMNM P88822 (Fig.
|
|
<figureCitation id="F9BDEBB806DDC7EFB888E47626CC3415" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5g</figureCitation>
|
|
), is a fragment of the bridge that may similarly represent part of the hyo- or hypoplastron. A rib-like thickening on the visceral side of this fragment is the remnants of a buttress. A series of clearly delineated sulci form the outlines of at least two inframarginals, two marginals, and the neighboring plastral scutes.
|
|
</paragraph>
|
|
<paragraph id="05D55C7FDFFF1C10F217ADE2597273CA" pageId="0" pageNumber="117">
|
|
<emphasis id="1CD96C09CE9D809644EE5DEFB0D0AD3F" bold="true" pageId="0" pageNumber="117">
|
|
<emphasis id="1AA32C2D2395558F7FAFF50FC8F8D4D2" italics="true" pageId="0" pageNumber="117">Mesoplastron</emphasis>
|
|
.
|
|
</emphasis>
|
|
No mesoplastral remains could be identified as such.
|
|
</paragraph>
|
|
<paragraph id="46C1089E0E33FFF540249D98DF649F42" pageId="0" pageNumber="117">
|
|
<emphasis id="90C6AC94AA780CBB48A33ABA8BFDAF8E" bold="true" pageId="0" pageNumber="117">
|
|
<emphasis id="34310F2C1D1C031675191075ADBA5485" italics="true" pageId="0" pageNumber="117">Hypoplastron</emphasis>
|
|
.
|
|
</emphasis>
|
|
We are able to identify at least three remains as hypoplastra, WMNM P88823 (Fig.
|
|
<figureCitation id="78EEB7AC625E8E090B6B27F92794AD4D" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5i</figureCitation>
|
|
), P88825 (Fig.
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|
<figureCitation id="1CB0714C3DF9D5982E195E1127C5003C" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5j</figureCitation>
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), and P88824 (Fig.
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<figureCitation id="70C554206BDAD81E6B495B7F388D4BCE" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5k</figureCitation>
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), all of which preserve the thickened inguinal notch, but can be recognized as hypoplastra by remnants of the xiphiplastral articulation. WMNM P88823, the best of the three preserved elements, documents the transverse contact with the mesoplastron and the slightly oblique posterior contact with the xiphiplastron, which is dorsally stabilized by two xiphiplastral processes that dorsally cover the hypoplastron and a hypoplastral process that dorsally covers the xiphiplastron (Fig.
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<figureCitation id="5F163F46CA7254B3BDAA116846F89D90" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5i</figureCitation>
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). The relatively intact medial margin suggests that central and posterior plastral fontanelles are present (see arrows in Fig.
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<figureCitation id="586F9E85B8BE5F97D7CAB629215B8749" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5i</figureCitation>
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) and that the hypoplastra only contacted one another along rib-like processes that radiate from the inguinal notch, features otherwise found in
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<taxonomicName id="B1C5AC344D549F1D852B1763340450F1" authorityName="Hay" authorityYear="1908" class="Reptilia" family="Pleurosternidae" genus="Naomichelys" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Naomichelys speciosa" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="species" species="speciosa">
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<emphasis id="A54D9BEF4B5D0A0B3DE2A94AAC8C1EC3" italics="true" pageId="0" pageNumber="117">Naomichelys speciosa</emphasis>
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</taxonomicName>
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(
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<bibRefCitation id="5D0FB8F6694FEB42BF8DC2CC2B873C58" DOI="https://doi.org/10.1666/14-002" author="Joyce, WG" journalOrPublisher="Journal of Paleontology" pageId="0" pageNumber="117" pagination="1257 - 1287" refId="B16" refString="Joyce, WG, Sterli, J, Chapman, SD, 2014. The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas. Journal of Paleontology 88: 1257 - 1287, DOI: https://doi.org/10.1666/14-002" title="The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas." url="https://doi.org/10.1666/14-002" volume="88" year="2014">Joyce et al. 2014</bibRefCitation>
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), but not in the holotype of
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<taxonomicName id="444D62FD4E3CFA74B33C93B23D110D5F" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
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<emphasis id="B0942F3CDE434D68FC4D25D5D54A0373" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
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</taxonomicName>
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(NHMUK R171). The relatively small size of this specimen opens the possibility, however, that these are juvenile features. As preserved, the abdominal/femoral sulcus crosses the hypoplastron diagonally from the inguinal notch to the central plastral fontanelle, but otherwise runs posterior from the notch to meet with the skin-scute sulcus on the dorsal side (Fig.
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<figureCitation id="63ACF304D30CB84D3D01C6A787C2DDE0" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5i</figureCitation>
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). In this area, the hypoplastron is thickened to nearly 2 cm in the smallest available specimen (WMNM P88823), but reaches 2.5 cm in the largest (WMNM P88825). In the area of the notch, the skin-scute sulcus runs onto the dorsal side of the hypoplastron to form a semi-lunate lip, but runs near parallel to the margin once it reaches the xiphiplastron (Fig.
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<figureCitation id="F1AE34B665EAA3078315ABD22AE361C3" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5j</figureCitation>
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).
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</paragraph>
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<paragraph id="F69E2C72D8EF036FC316178C70C8C414" pageId="0" pageNumber="117">
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<emphasis id="BFA24ECD4DF213F4BD833BAAE987DEC2" bold="true" pageId="0" pageNumber="117">
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<emphasis id="D0FA4BB838F88E8D824F532D22CA6BBC" italics="true" pageId="0" pageNumber="117">Xiphiplastron</emphasis>
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.
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</emphasis>
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The remnants of a xiphiplastron are preserved in WMNM P88823 attached to the hypoplastron (Fig.
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<figureCitation id="15CC8922741382BB6B550DB28B69C091" captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Plastral remains of Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. (a) WMNM P 88819, a partial entoplastron; (b) WMNM P 88818, a right epiplastron; (c) WMNM P 48296, a partial entoplastron; (d) WMNM P 47721, a partial right hyoplastron; (e) WMNM P 88820, a partial left hyoplastron; (f) WMNM P 48226, a plastral fragment; (g) WMNM P 88822, a plastral fragment; (h) WMNM P 88821, a plastral fragment; (i) WMNM P 88823, a partial left hypo- and xiphiplastron; (j) WMNM P 88825, a partial right hypoplastron; (k) WMNM P 88824, a right partial hypoplastron. With exception of the illustrations, all specimens are to scale. Abbreviations: AB = abdominal scale; EG = extragular scale; EN = entoplastral scale; FE = femoral scale; GU = gular scale; HU = humeral scale; hyp = hypoplastron; xi = xiphiplastron." figureDoi="10.3897/fr.26.102128.figure5" httpUri="https://binary.pensoft.net/fig/858750" pageId="0" pageNumber="117">5i</figureCitation>
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). Its contact with the hypoplastron is described above, as is the development of the skin-scute sulcus. A comparison with
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<taxonomicName id="AFC41AA22AAB3F3B43320C9E4212EFB6" authorityName="Hay" authorityYear="1908" class="Reptilia" family="Pleurosternidae" genus="Naomichelys" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Naomichelys speciosa" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="species" species="speciosa">
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<emphasis id="B3BD1F34528748BB25DEF37AB45E2D67" italics="true" pageId="0" pageNumber="117">Naomichelys speciosa</emphasis>
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</taxonomicName>
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suggests that the xiphiplastron broke close to the femoral-anal sulcus. The anal scute, therefore, did not cross the hypoplastra-xiphiplastral suture.
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</paragraph>
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<paragraph id="71D5101205B102A7A4E8D02FB2A8F078" pageId="0" pageNumber="117">
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<emphasis id="DDC0B36A383690FF7EDC07CE4B50485F" bold="true" pageId="0" pageNumber="117">
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<emphasis id="3CE93580B3E1CD41ADE889FCFBAD6D80" italics="true" pageId="0" pageNumber="117">Granicones</emphasis>
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.
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</emphasis>
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More than 150 granicones are available for study (Fig.
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<figureCitation id="F3C07436301C5746493B8B7E0C30A12A" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A selection of granicones, Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. All specimens are shown to scale in lateral (top), apical (bottom left), and basal view (bottom right) and associated with their WMNM catalog number. Specimens were in part chosen to highlight the morphological extremely highlighted by the PC plots (see Fig. 7)." figureDoi="10.3897/fr.26.102128.figure6" httpUri="https://binary.pensoft.net/fig/858751" pageId="0" pageNumber="117">6</figureCitation>
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). With few exceptions (see below), all granicones are singular, conical elements that consist of a flat base, which anchored the element into the skin, and an apex, which protruded from the body (
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<bibRefCitation id="871748625796CF550826C2C9E37450B8" DOI="https://doi.org/10.1666/14-002" author="Joyce, WG" journalOrPublisher="Journal of Paleontology" pageId="0" pageNumber="117" pagination="1257 - 1287" refId="B16" refString="Joyce, WG, Sterli, J, Chapman, SD, 2014. The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas. Journal of Paleontology 88: 1257 - 1287, DOI: https://doi.org/10.1666/14-002" title="The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas." url="https://doi.org/10.1666/14-002" volume="88" year="2014">Joyce et al. 2014</bibRefCitation>
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). The external surface of all elements is covered by surface texture that closely mirrors that of the shell. A qualitative assessment suggests that three morphotypes are present: cones (ca. 35%), hooks (ca. 43%), and plates (ca. 22%). As the name implies, cones are conical elements where the apex is situated above the base (e.g., Fig.
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<figureCitation id="9EADD9443648B585BDD6C597EB55C484" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A selection of granicones, Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. All specimens are shown to scale in lateral (top), apical (bottom left), and basal view (bottom right) and associated with their WMNM catalog number. Specimens were in part chosen to highlight the morphological extremely highlighted by the PC plots (see Fig. 7)." figureDoi="10.3897/fr.26.102128.figure6" httpUri="https://binary.pensoft.net/fig/858751" pageId="0" pageNumber="117">6</figureCitation>
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: P88864, P88873). These elements tend to be smaller. Hooks greatly resemble cones, but the apex is offset beyond the surface of the base (e.g., Fig.
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<figureCitation id="2EBF9FE0D886AECBF58F9EABDD634DE9" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A selection of granicones, Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. All specimens are shown to scale in lateral (top), apical (bottom left), and basal view (bottom right) and associated with their WMNM catalog number. Specimens were in part chosen to highlight the morphological extremely highlighted by the PC plots (see Fig. 7)." figureDoi="10.3897/fr.26.102128.figure6" httpUri="https://binary.pensoft.net/fig/858751" pageId="0" pageNumber="117">6</figureCitation>
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: P88922, P88856). Plates, finally, resemble hooks in that the base is offset from the base, but that are notably flat (e.g., Fig.
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<figureCitation id="B665D550870DF7896FD0C4DB862B2733" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A selection of granicones, Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. All specimens are shown to scale in lateral (top), apical (bottom left), and basal view (bottom right) and associated with their WMNM catalog number. Specimens were in part chosen to highlight the morphological extremely highlighted by the PC plots (see Fig. 7)." figureDoi="10.3897/fr.26.102128.figure6" httpUri="https://binary.pensoft.net/fig/858751" pageId="0" pageNumber="117">6</figureCitation>
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: P88939, P88921). These tend to be the largest available elements. Our morphometric analysis (see below) suggests that these morphotypes do not populate discrete parts of the morphospace, but rather grade into each other. We nonetheless retain this terminology for convenience. As all elements were found in isolation, we do not know which parts of the body they once covered. A complete skeleton of
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<taxonomicName id="2692A4213EF2FC3B75FE08BEA3C04AE1" authorityName="Hay" authorityYear="1908" class="Reptilia" family="Pleurosternidae" genus="Naomichelys" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Naomichelys speciosa" order="Testudines" pageId="0" pageNumber="117" phylum="Chordata" rank="species" species="speciosa">
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<emphasis id="AFCCA541ED1C33972411DEC2C412E4C6" italics="true" pageId="0" pageNumber="117">Naomichelys speciosa</emphasis>
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</taxonomicName>
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that includes numerous articulated granicones, however, suggests that plates covered the external surfaces of the lower limbs while the spur-like hooks were located on the walking surfaces of the limbs (
|
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<bibRefCitation id="3E9CA3165D41673F8499378D155095E0" DOI="https://doi.org/10.1666/14-002" author="Joyce, WG" journalOrPublisher="Journal of Paleontology" pageId="0" pageNumber="117" pagination="1257 - 1287" refId="B16" refString="Joyce, WG, Sterli, J, Chapman, SD, 2014. The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas. Journal of Paleontology 88: 1257 - 1287, DOI: https://doi.org/10.1666/14-002" title="The skeletal morphology of the solemydid turtle Naomichelys speciosa from the Early Cretaceous of Texas." url="https://doi.org/10.1666/14-002" volume="88" year="2014">Joyce et al. 2014</bibRefCitation>
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).
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</paragraph>
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<caption id="C9092E5F75B9DCADE81E423769D0C26B" doi="10.3897/fr.26.102128.figure6" httpUri="https://binary.pensoft.net/fig/858751" pageId="0" pageNumber="117" start="Figure 6" startId="F6">
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<paragraph id="B7EFBD86AA3D9C51A3EA9257D4F488D1" pageId="0" pageNumber="117">
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<emphasis id="E29B58A03451867B3A25514575742288" bold="true" pageId="0" pageNumber="117">Figure 6.</emphasis>
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A selection of granicones,
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<taxonomicName id="604CA348C6BEAA47860F8EA2264E6F06" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
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<emphasis id="39D852111872B276D5095B3C9E724054" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
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</taxonomicName>
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from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3D models. All specimens are shown to scale in lateral (top), apical (bottom left), and basal view (bottom right) and associated with their WMNM catalog number. Specimens were in part chosen to highlight the morphological extremely highlighted by the PC plots (see Fig.
|
|
<figureCitation id="E5C8B3B204ECDC0984F4C622F52D74F0" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Morphospaces of four granicone measurements from PCA ordination (N = 165). (a, b), bivariate morphospaces including size variation. (c, d), morphospaces corrected for size and allometry. A, PC 1 vs. PC 2. B, PC 2 vs. PC 3. C, PC 2 vs. PC 3. D, PC 2 vs. PC 4. The proportion of variance explained by PC axes are indicated in the axis labels, whereby values in A, B represent total variance (including size) and values in B, C represent shape variance (re-scaled to exclude PC 1). An identical plot with unique point labels is provided in Suppl. material 2." figureDoi="10.3897/fr.26.102128.figure7" httpUri="https://binary.pensoft.net/fig/858752" pageId="0" pageNumber="117">7</figureCitation>
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).
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</paragraph>
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</caption>
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<paragraph id="C1BF7355E7383E7E34EF1AEF9366EF38" pageId="0" pageNumber="117">
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A total of five elements deviate strongly from the three morphotypes described above (Fig.
|
|
<figureCitation id="C8FAF577872B0702B429D679F588E1C7" captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. A selection of granicones, Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. All images were obtained from CT generated 3 D models. All specimens are shown to scale in lateral (top), apical (bottom left), and basal view (bottom right) and associated with their WMNM catalog number. Specimens were in part chosen to highlight the morphological extremely highlighted by the PC plots (see Fig. 7)." figureDoi="10.3897/fr.26.102128.figure6" httpUri="https://binary.pensoft.net/fig/858751" pageId="0" pageNumber="117">6</figureCitation>
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). Two of these (WMNM P88935 and P88940) resemble large hooks by having an apex that is offset relative to the base, but the base in itself is not flat and round, but rather folded along the long axis of the element. We speculate that this type of element may have covered folds on the outside of the limbs. The three other unusual granicones are remnants of compound granicones. WMNM P88974 consists of a single, conical apex, but open sutures on at least one side of its base suggest that it was laterally sutured to at least one more granicone. WMNM P88964 consists of two conical apices that are broadly sutured to one another at their bases. Although the outline of the jointly formed base is somewhat angular, the element appears to be intact. WMNM P88962 also consists of two conical apices, but numerous angularly arranged sutures on both sides of the jointly formed base suggest that it formally articulated with at least two, but likely up to five additional granicones, to form a compound element with at least four to seven apices.
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</paragraph>
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<paragraph id="B87E94340609FDFCDBC8CC91300ADA89" pageId="0" pageNumber="117">
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PCA of all measurements indicates that 65.7% of the overall variance can be explained by size or allometric differences between specimens, explained by PC1 (Table
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<tableCitation id="3BF20148500E28BD2A1FCF72CAA3DB7A" captionStart="Table 1" captionStartId="T1" captionText="Table 1. PCA results (n = 136)." httpUri="http://table.plazi.org/id/5D016B16A480C122AEB8591A24810488" pageId="0" pageNumber="117" tableUuid="5D016B16A480C122AEB8591A24810488">1</tableCitation>
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). No differentiation of morphotypes is apparent based on size differences alone. Our morphospace plot of size-adjusted principal components (Fig.
|
|
<figureCitation id="6976C07145314CCD983C64839E447AD7" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Morphospaces of four granicone measurements from PCA ordination (N = 165). (a, b), bivariate morphospaces including size variation. (c, d), morphospaces corrected for size and allometry. A, PC 1 vs. PC 2. B, PC 2 vs. PC 3. C, PC 2 vs. PC 3. D, PC 2 vs. PC 4. The proportion of variance explained by PC axes are indicated in the axis labels, whereby values in A, B represent total variance (including size) and values in B, C represent shape variance (re-scaled to exclude PC 1). An identical plot with unique point labels is provided in Suppl. material 2." figureDoi="10.3897/fr.26.102128.figure7" httpUri="https://binary.pensoft.net/fig/858752" pageId="0" pageNumber="117">7</figureCitation>
|
|
) shows that PC2, which explains 71.9% of shape variation after correcting for size, is the only principal component along which separate clusters of specimens may be determined, as plated granicones plot at low PC2 values with minimal overlap of specimens pre-assigned to other morphotypes. Cones and hooks have broad overlap across PC3 (20% of variance; Fig.
|
|
<figureCitation id="29C80173CD8A74CE61B8B372F57EF4C7" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Morphospaces of four granicone measurements from PCA ordination (N = 165). (a, b), bivariate morphospaces including size variation. (c, d), morphospaces corrected for size and allometry. A, PC 1 vs. PC 2. B, PC 2 vs. PC 3. C, PC 2 vs. PC 3. D, PC 2 vs. PC 4. The proportion of variance explained by PC axes are indicated in the axis labels, whereby values in A, B represent total variance (including size) and values in B, C represent shape variance (re-scaled to exclude PC 1). An identical plot with unique point labels is provided in Suppl. material 2." figureDoi="10.3897/fr.26.102128.figure7" httpUri="https://binary.pensoft.net/fig/858752" pageId="0" pageNumber="117">7a</figureCitation>
|
|
) and PC4 (8.0% of variance; Fig.
|
|
<figureCitation id="DB7EE1A90E31D3BCF03DF5B657C74658" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Morphospaces of four granicone measurements from PCA ordination (N = 165). (a, b), bivariate morphospaces including size variation. (c, d), morphospaces corrected for size and allometry. A, PC 1 vs. PC 2. B, PC 2 vs. PC 3. C, PC 2 vs. PC 3. D, PC 2 vs. PC 4. The proportion of variance explained by PC axes are indicated in the axis labels, whereby values in A, B represent total variance (including size) and values in B, C represent shape variance (re-scaled to exclude PC 1). An identical plot with unique point labels is provided in Suppl. material 2." figureDoi="10.3897/fr.26.102128.figure7" httpUri="https://binary.pensoft.net/fig/858752" pageId="0" pageNumber="117">7b</figureCitation>
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|
). The absence of separation for these morphotypes and the broad spread of values across PC3 can be explained by shape variation within the pre-assigned hook or cone condition. For example, hooks exist that have large basal measurements but low tip height (Table
|
|
<tableCitation id="CF645B71F4B86BA21F6C8EE23D56B61B" captionStart="Table 1" captionStartId="T1" captionText="Table 1. PCA results (n = 136)." httpUri="http://table.plazi.org/id/5D016B16A480C122AEB8591A24810488" pageId="0" pageNumber="117" tableUuid="5D016B16A480C122AEB8591A24810488">1</tableCitation>
|
|
), such as WMNM P88873 (Fig.
|
|
<figureCitation id="3343EFFA388DAD0F390AE074269AC2C1" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Morphospaces of four granicone measurements from PCA ordination (N = 165). (a, b), bivariate morphospaces including size variation. (c, d), morphospaces corrected for size and allometry. A, PC 1 vs. PC 2. B, PC 2 vs. PC 3. C, PC 2 vs. PC 3. D, PC 2 vs. PC 4. The proportion of variance explained by PC axes are indicated in the axis labels, whereby values in A, B represent total variance (including size) and values in B, C represent shape variance (re-scaled to exclude PC 1). An identical plot with unique point labels is provided in Suppl. material 2." figureDoi="10.3897/fr.26.102128.figure7" httpUri="https://binary.pensoft.net/fig/858752" pageId="0" pageNumber="117">7a</figureCitation>
|
|
), the positive extreme point on PC3, but simultaneously hooks exist that have high tip heights compared to relatively small bases (e.g., WMNM P88864, which plots at low PC3 values for the hooked morphotype; Fig.
|
|
<figureCitation id="9F9C6621C2ECDCEE5C9ADDF237784C74" captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Morphospaces of four granicone measurements from PCA ordination (N = 165). (a, b), bivariate morphospaces including size variation. (c, d), morphospaces corrected for size and allometry. A, PC 1 vs. PC 2. B, PC 2 vs. PC 3. C, PC 2 vs. PC 3. D, PC 2 vs. PC 4. The proportion of variance explained by PC axes are indicated in the axis labels, whereby values in A, B represent total variance (including size) and values in B, C represent shape variance (re-scaled to exclude PC 1). An identical plot with unique point labels is provided in Suppl. material 2." figureDoi="10.3897/fr.26.102128.figure7" httpUri="https://binary.pensoft.net/fig/858752" pageId="0" pageNumber="117">7a</figureCitation>
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). Similarly, cones with strongly asymmetrical bases plot at extreme negative PC3 values (e.g., WMNM P88940), whereas cones with similar tip offset but symmetrical bases plot at extreme positive PC3 values (e.g., WMNM P88921). The relatively close association in morphospace of hooks like WMNM P88873 and cones like WMNM P88921 is thus primarily caused by similar base dimensions paired with low tip heights (Table
|
|
<tableCitation id="C421CF7627306E7580DA607F83D7F1D4" captionStart="Table 1" captionStartId="T1" captionText="Table 1. PCA results (n = 136)." httpUri="http://table.plazi.org/id/5D016B16A480C122AEB8591A24810488" pageId="0" pageNumber="117" tableUuid="5D016B16A480C122AEB8591A24810488">1</tableCitation>
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), regardless of their difference in tip offset which is the primary distinction of cones and hooks based on qualitative comparisons.
|
|
</paragraph>
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|
<caption id="3A24A0AC4426D4BC70F31C8F660037ED" doi="10.3897/fr.26.102128.figure7" httpUri="https://binary.pensoft.net/fig/858752" pageId="0" pageNumber="117" start="Figure 7" startId="F7">
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|
<paragraph id="9DFD62A376A3A763E0C23B4FB02B05D5" pageId="0" pageNumber="117">
|
|
<emphasis id="454CA8DE1D470EAAEDC158B361B40436" bold="true" pageId="0" pageNumber="117">Figure 7.</emphasis>
|
|
Morphospaces of four granicone measurements from PCA ordination (N = 165). (
|
|
<emphasis id="9243955E17A50A4F2D3DD83A8BE94596" bold="true" pageId="0" pageNumber="117">a, b)</emphasis>
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|
, bivariate morphospaces including size variation. (
|
|
<emphasis id="4EBB38A452BDC09BA3F6825DA65FCFF7" bold="true" pageId="0" pageNumber="117">c, d</emphasis>
|
|
), morphospaces corrected for size and allometry. A, PC1 vs. PC2. B, PC2 vs. PC3. C, PC2 vs. PC3. D, PC2 vs. PC4. The proportion of variance explained by PC axes are indicated in the axis labels, whereby values in
|
|
<emphasis id="D5CF9A5D776CC1727644A93A9130CD21" bold="true" pageId="0" pageNumber="117">A, B</emphasis>
|
|
represent total variance (including size) and values in
|
|
<emphasis id="EECF207E4FD4E2C203622E6B12ABB2DB" bold="true" pageId="0" pageNumber="117">B, C</emphasis>
|
|
represent shape variance (re-scaled to exclude PC1). An identical plot with unique point labels is provided in Suppl. material 2.
|
|
</paragraph>
|
|
</caption>
|
|
<caption id="5D016B16A480C122AEB8591A24810488" ID-Table-UUID="5D016B16A480C122AEB8591A24810488" httpUri="http://table.plazi.org/id/5D016B16A480C122AEB8591A24810488" pageId="0" pageNumber="117" start="Table 1" startId="T1">
|
|
<paragraph id="CE1767808B739B4692AD511D594D0D41" pageId="0" pageNumber="117">
|
|
<emphasis id="253AC6C7A4F8948EC5259C549E3979A8" bold="true" pageId="0" pageNumber="117">Table 1.</emphasis>
|
|
PCA results (n = 136).
|
|
</paragraph>
|
|
</caption>
|
|
<paragraph id="BB07844D3AE7F73DD563AEE97483D185" pageId="0" pageNumber="117">
|
|
<table id="A938239D24D3713C53C0E7AAAFA40AAE" pageId="0" pageNumber="117">
|
|
<tr id="A627B343EE2A5FBB65C72D1AAFC0FCFA" pageId="0" pageNumber="117">
|
|
<th id="528C2413D0C8BC4106C0E539158B839D" colspan="1" isEmpty="true" pageId="0" pageNumber="117" rowspan="1">-</th>
|
|
<th id="8CB0C75922714D093D07390FDAA20A26" colspan="1" pageId="0" pageNumber="117" rowspan="1">PC1</th>
|
|
<th id="246D150A82CAE4B12F6521B53A45D079" colspan="1" pageId="0" pageNumber="117" rowspan="1">PC2</th>
|
|
<th id="59AC1B9023C2A2848F88E904A5376378" colspan="1" pageId="0" pageNumber="117" rowspan="1">PC3</th>
|
|
<th id="537549122F8B51DCC3FAC6C785B83FC5" colspan="1" pageId="0" pageNumber="117" rowspan="1">PC4</th>
|
|
</tr>
|
|
<tr id="74052F7D1A87668F06BC948AC0C7F78D" pageId="0" pageNumber="117">
|
|
<td id="75F4BF941535218EDB2B5670106EC599" colspan="1" pageId="0" pageNumber="117" rowspan="1">Eigenvalues</td>
|
|
<td id="F5BEFD917B7BF8E8292301AD5017CC17" colspan="1" pageId="0" pageNumber="117" rowspan="1">2.628</td>
|
|
<td id="6A22400DAFF1B6DE5E7E9B7C391DD9CB" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.987</td>
|
|
<td id="953EA0384AF8E3CE336385C9DB147496" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.275</td>
|
|
<td id="275A60A0E6C2F70A110890DB50861490" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.110</td>
|
|
</tr>
|
|
<tr id="A3BCB6121AA37AE8A5D433B1CE1AEF18" pageId="0" pageNumber="117">
|
|
<td id="97088C45693AC8A8E50E1EE82F626D9B" colspan="1" pageId="0" pageNumber="117" rowspan="1">Proportion of total variance explained</td>
|
|
<td id="D5F29662FE6440F08B7D384E91CA6FF9" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.657</td>
|
|
<td id="CA5639A11AF7EE0B45A3EF906361C6C2" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.247</td>
|
|
<td id="CBF7A689F5B64D1FBF89C6918876E5B3" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.069</td>
|
|
<td id="2179C37FAA0593FC3930DA5DE044AB2D" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.028</td>
|
|
</tr>
|
|
<tr id="8D5061BF86492960B62C33D27E233D9F" pageId="0" pageNumber="117">
|
|
<td id="39C593EEAD022DA681F9A96262026BFD" colspan="1" pageId="0" pageNumber="117" rowspan="1">Proportion of shape variance explained</td>
|
|
<td id="6E3CE3D784F9E24C8E0425925817B2CF" colspan="1" pageId="0" pageNumber="117" rowspan="1">-</td>
|
|
<td id="B76C9304E62576D421923813ED090380" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.719</td>
|
|
<td id="1329083E4A1941DB6EE1F1A11109AFD8" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.200</td>
|
|
<td id="3FF00A428813B56CE4452CFA199267E6" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.080</td>
|
|
</tr>
|
|
<tr id="87626E0CCE126DEC7B0E91A34ED60CB2" pageId="0" pageNumber="117">
|
|
<td id="3A4C6DA29981932FD977CF4E1700367F" colspan="5" pageId="0" pageNumber="117" rowspan="1">Eigenvector coefficients</td>
|
|
</tr>
|
|
<tr id="9BF7F2416B566381D803CB031ED5B18C" pageId="0" pageNumber="117">
|
|
<td id="29D5BB40D4BA28BFDDEEB97A9460E072" colspan="1" pageId="0" pageNumber="117" rowspan="1">height</td>
|
|
<td id="03CE712642A407C993986C3DA5B7E7B1" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.170</td>
|
|
<td id="B8C162B19FE812E59D216979FAF5C202" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.963</td>
|
|
<td id="E1EFDFD57EA9C7E76A9BC09BD004E16B" colspan="1" pageId="0" pageNumber="117" rowspan="1">-0.155</td>
|
|
<td id="C774E4F5FB3EAB22AE6CF022A7D8CD52" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.140</td>
|
|
</tr>
|
|
<tr id="BF010ED8F3E80088B5EB9B45502241E6" pageId="0" pageNumber="117">
|
|
<td id="15749E1481EEBC7AB33D404E11AFB1B1" colspan="1" pageId="0" pageNumber="117" rowspan="1">minimum base</td>
|
|
<td id="1B01A6FEDA01A62B02FB9AC876F6AE23" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.575</td>
|
|
<td id="59166959DAF74D8E9B37DEED077AA866" colspan="1" pageId="0" pageNumber="117" rowspan="1">-0.010</td>
|
|
<td id="102231469B46E7411CD085EEB41AAFDF" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.551</td>
|
|
<td id="DAE0113836C658EF7347F4F4756A5002" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.597</td>
|
|
</tr>
|
|
<tr id="BBEA3AF4F7A8C1028A6EE309CAA2738E" pageId="0" pageNumber="117">
|
|
<td id="E99ABBD1B6AA8CBF64AD04385CD93F35" colspan="1" pageId="0" pageNumber="117" rowspan="1">maximum base</td>
|
|
<td id="6384E6CBF7F2C00FC7BB2A9A077B2A4E" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.591</td>
|
|
<td id="BC1AC51BDB1C6A92C8F9C025EC4C3826" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.045</td>
|
|
<td id="883B7C666E5F65D0BDDA365E065718E7" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.229</td>
|
|
<td id="95BDB7B355DF608D77FFCB60E4922555" colspan="1" pageId="0" pageNumber="117" rowspan="1">-0.772</td>
|
|
</tr>
|
|
<tr id="6137C6BA0182EBA03FF3CDC0D2339CC3" pageId="0" pageNumber="117">
|
|
<td id="429A554ADCEDB4D4B4B65B4AB7E848F9" colspan="1" pageId="0" pageNumber="117" rowspan="1">tip offset</td>
|
|
<td id="6F6252BBFE7A5CB3917270AB6E6FAFFA" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.540</td>
|
|
<td id="4064DFC2DB3F5C365E180266104747E5" colspan="1" pageId="0" pageNumber="117" rowspan="1">-0.24</td>
|
|
<td id="208ADF206F8B3FC83ABDC1C71579B554" colspan="1" pageId="0" pageNumber="117" rowspan="1">-0.788</td>
|
|
<td id="290413E41C64151A48EE22A4A9AFF8F7" colspan="1" pageId="0" pageNumber="117" rowspan="1">0.166</td>
|
|
</tr>
|
|
</table>
|
|
</paragraph>
|
|
<paragraph id="B6B38E5533F6499BB15A3116BA0E9558" pageId="0" pageNumber="117">
|
|
<emphasis id="33E586BBCFEE5FB6E78CC891AD9921FD" bold="true" pageId="0" pageNumber="117">
|
|
<emphasis id="E89CAB86163AD1179889C7F1980C7557" italics="true" pageId="0" pageNumber="117">Cranium</emphasis>
|
|
.
|
|
</emphasis>
|
|
A three-dimensionally preserved, partial, left parietal (WMNM P88806) is the only skull bone in the sample (Fig.
|
|
<figureCitation id="E2F5569D81B32C8AA48EF069FC708B2E" captionStart="Figure 8" captionStartId="F8" captionText="Figure 8. Helochelydra nopcsai from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. WMNM P 88806, a partial parietal in dorsal (a), ventral (b), medial (c), and lateral view (d). All images were obtained from CT generated 3 D models." figureDoi="10.3897/fr.26.102128.figure8" httpUri="https://binary.pensoft.net/fig/858753" pageId="0" pageNumber="117">8</figureCitation>
|
|
). The anterolateral and medial margins of its dorsal plate, which formerly contacted the left postorbital and the right parietal, are intact, but the anterior and posterior margins show signs of damage. Striations on the posteromedial side of the element, contained within the area that constituted the cavum cranii, could plausibly represent the articulation area with the supraoccipital, which is usually overlain by the parietal in this area. A short posterolateral facet may represent the former contact with the left squamosal. A comparison with IWCMS 1998.21, a complete
|
|
<taxonomicName id="305760CBE3E15A18957B76101222121D" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
|
|
<emphasis id="F07913D41D96FE589F543771E1DB9185" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
|
|
</taxonomicName>
|
|
skull from the Isle of Wight (
|
|
<bibRefCitation id="3FF5D05698FC3DAEED681A92D3B77B31" author="Joyce, WG" journalOrPublisher="Special Papers in Paleontology" pageId="0" pageNumber="117" pagination="75 - 97" refId="B15" refString="Joyce, WG, Chapman, SD, Moody, RTJ, Walker, CA, 2011. The skull of the solemydid turtle Helochelydra nopcsai from the Early Cretaceous (Barremian) of the Isle of Wight (UK) and a review of Solemydidae. Special Papers in Paleontology 86: 75 - 97" title="The skull of the solemydid turtle Helochelydra nopcsai from the Early Cretaceous (Barremian) of the Isle of Wight (UK) and a review of Solemydidae." volume="86" year="2011">Joyce et al. 2011</bibRefCitation>
|
|
), suggests that only the anterior half of the element is preserved. The two specimens likely originated from individuals of the same size, as WMNM P88806 has an anteroposterior length of about 2.7 cm, while the equivalent area measures about 2.8 cm in IWCMS 1998.21. The two skulls differ notably in that WMNM P88806 exhibits clear cranial sulci which outline at least 12 polygonal cranial scutes. Their great number, however, obscures homology with the cranial scutes of other Cretaceous turtles, as these typically exhibit only four scutes in the equivalent area (
|
|
<bibRefCitation id="66D2FE18E33D9CC5BF81FC9ACC6F50A2" DOI="https://doi.org/10.1080/14772019.2012.708674" author="Sterli, J" journalOrPublisher="Journal of Systematic Palaeontology" pageId="0" pageNumber="117" pagination="835 - 852" refId="B45" refString="Sterli, J, de la Fuente, MS, 2013. New evidence from the Palaeocene of Patagonia (Argentina) on the evolution and palaeo-biogeography of Meiolaniformes (Testudinata, new taxon name). Journal of Systematic Palaeontology 11: 835 - 852, DOI: https://doi.org/10.1080/14772019.2012.708674" title="New evidence from the Palaeocene of Patagonia (Argentina) on the evolution and palaeo-biogeography of Meiolaniformes (Testudinata, new taxon name)." url="https://doi.org/10.1080/14772019.2012.708674" volume="11" year="2013">Sterli and de la Fuente 2013</bibRefCitation>
|
|
;
|
|
<bibRefCitation id="FCB07969EBD4C7625C9DAE231E043C5F" DOI="https://doi.org/10.7717/peerj.11495" author="Evers, S" journalOrPublisher="Nauchnaya Sessiya Instituta Paleobiologii, Tezisy Dokladov" pageId="0" pageNumber="117" refId="B4" refString="Evers, S, Rollot, Y, Joyce, WG, 2021. New interpretation of the cranial osteology of the Early Cretaceous turtle Arundelemys dardeni (Paracryptodira) based on a CT-based re-evaluation of the holotype. PeerJ 9: e11495. https://doi.org/10.7717/peerj.11495" title="New interpretation of the cranial osteology of the Early Cretaceous turtle Arundelemys dardeni (Paracryptodira) based on a CT-based re-evaluation of the holotype. PeerJ 9: e 11495." url="https://doi.org/10.7717/peerj.11495" year="2021">Evers et al. 2021</bibRefCitation>
|
|
). On the ventral side, traces remain of the heavily damaged descending process. What remains is an outwardly convex crest of bone that documents the greatest width of the cavum cranii.
|
|
</paragraph>
|
|
<caption id="6A74F6BBB3C36F681591D7BE8B7CDA61" doi="10.3897/fr.26.102128.figure8" httpUri="https://binary.pensoft.net/fig/858753" pageId="0" pageNumber="117" start="Figure 8" startId="F8">
|
|
<paragraph id="A671BE2D3CD78A55359AAFAC3ABCC602" pageId="0" pageNumber="117">
|
|
<emphasis id="E1C2AE4360AC3FD8F69D411359F90738" bold="true" pageId="0" pageNumber="117">Figure 8.</emphasis>
|
|
<taxonomicName id="7024936535DB4C3C8AFE9D9915F81E12" authorityName="Lapparent de Broin & Murelaga" authorityYear="1999" family="Solemydidae" genus="Helochelydra" kingdom="Animalia" lsidName="Helochelydra nopcsai" order="Testudines" pageId="0" pageNumber="117" rank="species" species="nopcsai">
|
|
<emphasis id="40C8C035397C53376A0500CB3E8A8A7F" italics="true" pageId="0" pageNumber="117">Helochelydra nopcsai</emphasis>
|
|
</taxonomicName>
|
|
from the Early Cretaceous (late Barremian - early Aptian) of Balve, North Rhine-Westphalia, Germany. WMNM P88806, a partial parietal in dorsal (
|
|
<emphasis id="D9E072238977D404B1FF3B99740F76AB" bold="true" pageId="0" pageNumber="117">a</emphasis>
|
|
), ventral (
|
|
<emphasis id="D843067FF9C4C70AC78F501647854827" bold="true" pageId="0" pageNumber="117">b</emphasis>
|
|
), medial (
|
|
<emphasis id="90F617F8C75A51B3BCBC20A9D94F6A5B" bold="true" pageId="0" pageNumber="117">c</emphasis>
|
|
), and lateral view (
|
|
<emphasis id="79FB7719F235BA09EB804009F624AD7B" bold="true" pageId="0" pageNumber="117">d</emphasis>
|
|
). All images were obtained from CT generated 3D models.
|
|
</paragraph>
|
|
</caption>
|
|
</subSubSection>
|
|
</treatment>
|
|
</document> |