396 lines
78 KiB
XML
396 lines
78 KiB
XML
<document id="4BB6EC44C5C2FB040902AB05228BA322" ID-CLB-Dataset="268663" ID-DOI="10.1038/s41586-023-06538-y" ID-GBIF-Dataset="abb5ff8a-a81d-41ce-ac42-c8d88e09a0c4" ID-Zenodo-Dep="8368403" IM.bibliography_approvedBy="julia" IM.illustrations_approvedBy="julia" IM.materialsCitations_approvedBy="julia" IM.metadata_approvedBy="julia" IM.taxonomicNames_approvedBy="julia" IM.treatments_approvedBy="julia" checkinTime="1695322723319" checkinUser="julia" docAuthor="Dearden, Richard P., Lanzetti, Agnese, Giles, Sam, Johanson, Zerina, Jones, Andy S., Lautenschlager, Stephan, Randle, Emma & Sansom, Ivan J." docDate="2023" docId="03C0340FFFD5FF90FCC0F8D39BE94655" docLanguage="en" docName="NatureCLXVI.1-19..pdf" docOrigin="Nature 2023" docTitle="Eriptychius americanus , PF 1795" docType="treatment" docVersion="3" lastPageNumber="4" masterDocId="FFF94C77FFD5FF93FFAEFFD79A554161" masterDocTitle="The oldest three-dimensionally preserved vertebrate neurocranium" masterLastPageNumber="19" masterPageNumber="1" pageNumber="1" updateTime="1695324630334" updateUser="ExternalLinkService">
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<mods:title id="B3BB3F6CCDE06A57E14BD7111059E7E7">The oldest three-dimensionally preserved vertebrate neurocranium</mods:title>
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<mods:namePart id="1CB505C8B5B9D2ACDBF4C92350CA3061">Dearden, Richard P.</mods:namePart>
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<mods:namePart id="D02DD016307B631B756AD5B8B087285C">Lanzetti, Agnese</mods:namePart>
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<mods:namePart id="8EF4E017B42F495C6433191EBF20A348">Giles, Sam</mods:namePart>
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<mods:namePart id="A7CB2A4FC7F93DFED1DA50AFC8396E4F">Johanson, Zerina</mods:namePart>
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<mods:namePart id="36E42C652462F347D08C7CCA66FC83DF">Jones, Andy S.</mods:namePart>
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<mods:namePart id="4E65F87564B4412649BB07CABA8DCC31">Lautenschlager, Stephan</mods:namePart>
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<mods:namePart id="87E8F574C9448E19E5C3F206BB79EAD6">Randle, Emma</mods:namePart>
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<mods:namePart id="7F00D4808AC8F5176C8682C632289940">Sansom, Ivan J.</mods:namePart>
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<mods:title id="B4B83B403EA8A87A1F16FE3FCE530E4B">Nature</mods:title>
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<mods:date id="EFE088378D595ADDDE4B96F40B279D36">2023</mods:date>
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<mods:number id="5C18A33FCC4F11EB8A4EE08FDF72F2C8">2023-09-20</mods:number>
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<mods:number id="6BF7361CD8F180DB279B66EDB666E049">2023</mods:number>
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<mods:identifier id="382F2DB2F613D6048E829F1CE0F1D548" type="CLB-Dataset">268663</mods:identifier>
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<mods:identifier id="6FDE2F9D63A3AC9D7801E67E76D67D67" type="DOI">10.1038/s41586-023-06538-y</mods:identifier>
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<mods:identifier id="E760D0B0A454956CFD7FBC4C0AE3CE83" type="GBIF-Dataset">abb5ff8a-a81d-41ce-ac42-c8d88e09a0c4</mods:identifier>
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<mods:identifier id="926D5E1BD46D24B19A8DA87623DDE09E" type="Zenodo-Dep">8368403</mods:identifier>
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<treatment id="03C0340FFFD5FF90FCC0F8D39BE94655" ID-DOI="http://doi.org/10.5281/zenodo.8368398" ID-Zenodo-Dep="8368398" LSID="urn:lsid:plazi:treatment:03C0340FFFD5FF90FCC0F8D39BE94655" httpUri="http://treatment.plazi.org/id/03C0340FFFD5FF90FCC0F8D39BE94655" lastPageId="3" lastPageNumber="4" pageId="0" pageNumber="1">
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<subSubSection id="C373D692FFD5FF93FCC0F8D39FE74676" box="[878,1458,1795,1816]" pageId="0" pageNumber="1" type="nomenclature">
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<paragraph id="8BD68519FFD5FF93FCC0F8D39FE74676" blockId="0.[815,1497,1193,1903]" box="[878,1458,1795,1816]" pageId="0" pageNumber="1">
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<taxonomicName id="4C69FE9AFFD5FF93FCC0F8D39E034676" authorityName=", PF" authorityYear="1795" box="[878,1110,1796,1815]" family="Eriptychiidae" genus="Eriptychius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="americanus">
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<emphasis id="B91D590BFFD5FF93FCC0F8D39E034676" bold="true" box="[878,1110,1796,1815]" italics="true" pageId="0" pageNumber="1">Eriptychius americanus</emphasis>
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</taxonomicName>
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walcott,p. 167,plate 4,
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<figureCitation id="1352999CFFD5FF93FA93F8D49FC64676" box="[1341,1427,1795,1816]" captionStart="cartilages B" captionStartId="11.[816,902,1768,1786]" captionTargetBox="[125,1477,131,1744]" captionTargetId="figure-7@11.[125,1477,131,1744]" captionText="cartilages B;i,j bothsurfaces of mediolateral cartilages C; k-m,left orbital cartilage in(k) medial,(l) posterior and(m) latero-posterior view.Colour scheme andabbreviations as inFig.2." pageId="0" pageNumber="1">figures 5</figureCitation>
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–11
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</paragraph>
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</subSubSection>
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<subSubSection id="C373D692FFD5FF92FC9EF8E9995E44C3" lastPageId="1" lastPageNumber="2" pageId="0" pageNumber="1" type="materials_examined">
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<paragraph id="8BD68519FFD5FF92FC9EF8E9995E44C3" blockId="0.[815,1497,1193,1903]" lastBlockId="1.[105,787,1394,1987]" lastPageId="1" lastPageNumber="2" pageId="0" pageNumber="1">
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Type material.
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<materialsCitation id="3B018F44FFD5FF92FC67F8EB995E44C3" collectionCode="USNM" collectorName="C. D. walcott" country="United States of America" lastPageId="1" lastPageNumber="2" location="Harding Quarry" municipality="Canon City" pageId="0" pageNumber="1" specimenCode="USNM V2350" specimenCount="1" stateProvince="Colorado" typeStatus="syntype">
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<specimenCount id="9D6F4E90FFD5FF93FC67F8EB9E514631" box="[969,1028,1852,1872]" pageId="0" pageNumber="1">Seven</specimenCount>
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isolated fragments of dermal bone embedded in matrix,collected by
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<collectorName id="269CE0CFFFD5FF93FC5CF88E9E3B460C" box="[1010,1134,1881,1901]" pageId="0" pageNumber="1">C.D. walcott</collectorName>
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from the
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<location id="8EB6D3C2FFD5FF93FB69F88E9F37460C" LSID="urn:lsid:plazi:treatment:03C0340FFFD5FF90FCC0F8D39BE94655:8EB6D3C2FFD5FF93FB69F88E9F37460C" box="[1223,1378,1881,1902]" country="United States of America" municipality="Canon City" name="Harding Quarry" pageId="0" pageNumber="1" stateProvince="Colorado">Harding Quarry</location>
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,
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<collectingMunicipality id="6BB21F63FFD5FF93FAC9F88E9F80460C" box="[1383,1493,1881,1903]" pageId="0" pageNumber="1">Cañon City</collectingMunicipality>
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,
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<collectingRegion id="49AD4BFBFFD4FF92FFC4FAA59A9D44E7" box="[106,200,1394,1414]" country="United States of America" name="Colorado" pageId="1" pageNumber="2">Colorado</collectingRegion>
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,form the
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<typeStatus id="54D23BBBFFD4FF92FE8CFAA59B3A44E7" box="[290,367,1394,1414]" pageId="1" pageNumber="2">syntype</typeStatus>
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<specimenCode id="DBCF2D62FFD4FF92FEDFFAA59BA444E7" box="[369,497,1394,1414]" pageId="1" pageNumber="2">
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USNM V
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<quantity id="4C9128FCFFD4FF92FE6FFAA5985244E7" box="[449,519,1394,1414]" metricMagnitude="1" metricUnit="m" metricValue="5.968999999999999" pageId="1" pageNumber="2" unit="in" value="2350.0">2350 in</quantity>
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</specimenCode>
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the collections of the Smithsonian National Museum of Natural History in washington,DC,
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<collectingCountry id="F37EC589FFD4FF92FD74FA59995244C3" box="[730,775,1422,1442]" name="United States of America" pageId="1" pageNumber="2">USA</collectingCountry>
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.
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</materialsCitation>
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</paragraph>
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</subSubSection>
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<caption id="DF16D591FFD4FF92FFC4FBB59F534472" ID-DOI="http://doi.org/10.5281/zenodo.8368405" ID-Zenodo-Dep="8368405" httpUri="https://zenodo.org/record/8368405/files/figure.png" pageId="1" pageNumber="2" startId="1.[106,137,1122,1139]" subCaptionStartIDs="1.[1105,1134,1147,1165]" subCaptionStarts="Fig.2" targetBox="[348,1251,134,1092]" targetPageId="1" targetType="figure">
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<paragraph id="8BD68519FFD4FF92FFC4FBB59F534472" blockId="1.[106,787,1121,1299]" lastBlockId="1.[816,1489,1121,1299]" pageId="1" pageNumber="2">
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Fig.1 | Overview of
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<taxonomicName id="4C69FE9AFFD4FF92FEBCFBB698634512" authority="PF 1795" authorityName=", PF" authorityYear="1795" box="[274,566,1121,1139]" family="Eriptychiidae" genus="Eriptychius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="2" phylum="Chordata" rank="species" species="americanus">
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<emphasis id="B91D590BFFD4FF92FEBCFBB69BBB4512" bold="true" box="[274,494,1121,1139]" italics="true" pageId="1" pageNumber="2">Eriptychius americanus</emphasis>
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PF 1795
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</taxonomicName>
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. a,b, Photographs of part PF1795a,which had the split face set in epoxy and was manually prepared (a), and its counterpart PF 1795b,which remains in rock matrix(b). Both are shown in an anatomically ventral view.c,d, Digital model of computed tomographic data of the combined part and counterpart with most of dermal skeleton rendered transparent:anatomical ventral view (corresponding to the visible area of the part in epoxy) (c) and anatomical dorsal view (buried in matrix in the
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<emphasis id="B91D590BFFD4FF92FC9EFBB69F534472" bold="true" pageId="1" pageNumber="2">counterpart) (d). Colour scheme for renders:blue-greys,cranial cartilages (matching the detailed scheme in Fig.2); transparencies,the dermal skeleton; orange,branchial plates;red,orbital plates.Anterior to top in a–d.ant.tess., anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, branchial plate;cran.cart.,cranial cartilages;disp.,displaced;frag.,fragment; L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., vasculature;?,probable.Scale bar applies to all panels.</emphasis>
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</paragraph>
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</caption>
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<subSubSection id="C373D692FFD4FF92FFC4FA7B9BA1479B" pageId="1" pageNumber="2" type="diagnosis">
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<paragraph id="8BD68519FFD4FF92FFC4FA7B9BA1479B" blockId="1.[105,787,1394,1987]" pageId="1" pageNumber="2">
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Emended diagnosis. Agnathan with mesomeric dermal tesserae and scales formed from acellular bone overlain by ornament formed from coarse wide-calibre tubular dentine.Body scales covered in multiple elongate ridges.Antorbital neurocranium comprising symmetrical set of elements containing numerous large canals internally, endoskeleton closely associated with but not fused to the surrounding dermal skeleton. Shared with
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<taxonomicName id="4C69FE9AFFD4FF92FEE4F98F9BFD470A" authorityName="Walcott" authorityYear="1892" box="[330,424,1624,1643]" family="Astraspididae" genus="Astraspis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
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<emphasis id="B91D590BFFD4FF92FEE4F98F9BFD470A" bold="true" box="[330,424,1624,1643]" italics="true" pageId="1" pageNumber="2">Astraspis</emphasis>
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</taxonomicName>
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, arandaspids, other ‘ostracoderms’ excluding heterostracans:multiple branchial openings. Shared with
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<taxonomicName id="4C69FE9AFFD4FF92FFC4F9469A9047C5" authorityName="Walcott" authorityYear="1892" box="[106,197,1681,1700]" family="Astraspididae" genus="Astraspis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
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<emphasis id="B91D590BFFD4FF92FFC4F9469A9047C5" bold="true" box="[106,197,1681,1700]" italics="true" pageId="1" pageNumber="2">Astraspis</emphasis>
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</taxonomicName>
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and tessellate heterostracans:dermal head skeleton formed from dorsal and ventral ‘headshields’ of ornamented tesserae.Differs from
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<taxonomicName id="4C69FE9AFFD4FF92FF31F91C9AAF47BF" authorityName="Walcott" authorityYear="1892" box="[159,250,1739,1758]" family="Astraspididae" genus="Astraspis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
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<emphasis id="B91D590BFFD4FF92FF31F91C9AAF47BF" bold="true" box="[159,250,1739,1758]" italics="true" pageId="1" pageNumber="2">Astraspis</emphasis>
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</taxonomicName>
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in that ornament of dermal skeleton comprises ridges and presence of coarse tubular dentine.
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</paragraph>
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</subSubSection>
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<subSubSection id="C373D692FFD4FF92FFC4F8D39F1246A2" pageId="1" pageNumber="2" type="description">
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<paragraph id="8BD68519FFD4FF92FFC4F8D399ED472E" blockId="1.[105,787,1394,1987]" lastBlockId="1.[815,1498,1394,1987]" pageId="1" pageNumber="2">
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Description. Computed tomography scanning of the part and counterpart of PF 1795 (here termed PF 1795a and b, respectively;Methods) confirms the identity of this material as a partially articulated
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<taxonomicName id="4C69FE9AFFD4FF92FD4CF8EA9AE0460C" authorityName="PF" authorityYear="1795" family="Eriptychiidae" genus="Eriptychius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
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<emphasis id="B91D590BFFD4FF92FD4CF8EA9AE0460C" bold="true" italics="true" pageId="1" pageNumber="2">Eriptychius</emphasis>
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</taxonomicName>
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head
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<bibRefCitation id="EFF8F8E8FFD4FF92FF43F8819AA04602" author="Denison, R. H." box="[237,245,1878,1891]" journalOrPublisher="Fieldiana" pageId="1" pageNumber="2" pagination="131 - 192" part="16" refId="ref4498" refString="8. Denison, R. H. Ordovician vertebrates from Western United States. Fieldiana 16, 131 - 192 (1967" title="Ordovician vertebrates from Western United States" type="journal article" year="1967">
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<emphasis id="B91D590BFFD4FF92FF43F8819AA04602" bold="true" box="[237,245,1878,1891]" pageId="1" pageNumber="2">
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<superScript id="7C1C2851FFD4FF92FF43F8819AA04602" attach="left" box="[237,245,1878,1891]" fontSize="5" pageId="1" pageNumber="2">8</superScript>
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</emphasis>
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</bibRefCitation>
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, including components of both the dermal skeleton and endoskeleton (
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<figureCitation id="1352999CFFD4FF92FE8BF8A19B0D46EB" box="[293,344,1910,1931]" captionStart="Fig" captionStartId="1.[106,137,1122,1139]" captionTargetBox="[348,1251,134,1092]" captionTargetId="graphics-459@1.[389,1192,163,1056]" captionText="Fig.1 | Overviewof Eriptychiusamericanus PF 1795.a,b, Photographsof part counterpart) (d). Colourscheme forrenders:blue-greys,cranialcartilages PF1795a,which hadthe splitface set inepoxyandwas manually prepared (a), and (matchingthe detailed scheme inFig.2); transparencies,thedermal skeleton; itscounterpart PF 1795b,which remainsinrock matrix(b). Both are shown inan orange,branchial plates;red,orbital plates.Anterior totop ina–d.ant.tess., anatomically ventral view.c,d, Digital model of computed tomographic dataof anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, thecombined part and counterpart with most of dermal skeleton rendered branchial plate;cran.cart.,cranialcartilages;disp.,displaced;frag.,fragment; transparent:anatomical ventralview (corresponding tothe visible areaof L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., thepart in epoxy) (c) andanatomical dorsal view (buried in matrixin the vasculature;?,probable.Scale bar applies toall panels." figureDoi="http://doi.org/10.5281/zenodo.8368405" httpUri="https://zenodo.org/record/8368405/files/figure.png" pageId="1" pageNumber="2">Fig.1</figureCitation>
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, Extended
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<taxonomicName id="4C69FE9AFFD4FF92FE6AF8A19BA646EB" authorityName="Walker" authorityYear="1862" box="[452,499,1910,1930]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD4FF92FE58F8A1981F46EB" box="[502,586,1910,1931]" captionStart-0="Fig" captionStart-1="Fig" captionStartId-0="1.[106,137,1122,1139]" captionStartId-1="3.[106,137,701,718]" captionTargetBox-0="[348,1251,134,1092]" captionTargetBox-1="[313,1277,120,680]" captionTargetId-0="graphics-459@1.[389,1192,163,1056]" captionTargetId-1="graphics-692@3.[373,1255,157,437]" captionTargetPageId-0="1" captionTargetPageId-1="3" captionText-0="Fig.1 | Overviewof Eriptychiusamericanus PF 1795.a,b, Photographsof part PF1795a,which hadthe splitface set inepoxyandwas manually prepared (a), and itscounterpart PF 1795b,which remainsinrock matrix(b). Both are shown inan anatomically ventral view.c,d, Digital model of computed tomographic dataof thecombined part and counterpart with most of dermal skeleton rendered transparent:anatomical ventralview (corresponding tothe visible areaof thepart in epoxy) (c) andanatomical dorsal view (buried in matrixin the counterpart) (d). Colourscheme forrenders:blue-greys,cranialcartilages (matchingthe detailed scheme inFig.2); transparencies,thedermal skeleton; orange,branchial plates;red,orbital plates.Anterior totop ina–d.ant.tess., anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, branchial plate;cran.cart.,cranialcartilages;disp.,displaced;frag.,fragment; L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., vasculature;?,probable.Scale bar applies toall panels." captionText-1="Fig. 3| The neurocrania of vertebrates in a phylogenetic framework. a, Haikouichthys, adapted fromref.3, CC-BY 4.0.b, Petromyzon.c, Generalized cyathaspid,adapted from ref.20, Springer Nature.d, Sacabambaspis, reproduced fromref.31, copyright ©1993 Elsevier Masson SAS,all rights reserved (but see ref.53 for alternative interpretation).e, Eriptychius. f, Shuyu,adapted from ref.7, SpringerNature.g, Dicksonosteus adapted with permission from ref.14,Muséum national d’histoire naturelle,Paris. Phylogenetic schemeis the Adams consensus tree described inthe text,bold taxon namesrepresent thosedepicted above.Coloured regions illustrate positions of key sensory organs:yellow,eyes;blue,pineal.Light grey lines represent body outlines,black lines represent dermalbody armour,lightgrey" figureDoi-0="http://doi.org/10.5281/zenodo.8368405" figureDoi-1="http://doi.org/10.5281/zenodo.8368409" httpUri-0="https://zenodo.org/record/8368405/files/figure.png" httpUri-1="https://zenodo.org/record/8368409/files/figure.png" pageId="1" pageNumber="2">Figs.1–3</figureCitation>
|
||
and Supplementary Video;see Supplementary Information for comments on histology). The articulated individual is confined to the near surface of the matrix; below it is a mash of isolated elements typical of the Harding Sandstone bone beds including additional tesserae referable to
|
||
<taxonomicName id="4C69FE9AFFD4FF92FA94FA589FFE44C3" authorityName="PF" authorityYear="1795" box="[1338,1451,1423,1442]" family="Eriptychiidae" genus="Eriptychius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD4FF92FA94FA589FFE44C3" bold="true" box="[1338,1451,1423,1442]" italics="true" pageId="1" pageNumber="2">Eriptychius</emphasis>
|
||
</taxonomicName>
|
||
that do not seem to be associated with the articulated specimen (Extended
|
||
<taxonomicName id="4C69FE9AFFD4FF92FC81FA1F990A44BD" authorityName="Walker" authorityYear="1862" box="[815,863,1480,1500]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD4FF92FCCAFA1F99CC44BD" box="[868,921,1480,1501]" captionStart="Fig" captionStartId="1.[106,137,1122,1139]" captionTargetBox="[348,1251,134,1092]" captionTargetId="graphics-459@1.[389,1192,163,1056]" captionText="Fig.1 | Overviewof Eriptychiusamericanus PF 1795.a,b, Photographsof part counterpart) (d). Colourscheme forrenders:blue-greys,cranialcartilages PF1795a,which hadthe splitface set inepoxyandwas manually prepared (a), and (matchingthe detailed scheme inFig.2); transparencies,thedermal skeleton; itscounterpart PF 1795b,which remainsinrock matrix(b). Both are shown inan orange,branchial plates;red,orbital plates.Anterior totop ina–d.ant.tess., anatomically ventral view.c,d, Digital model of computed tomographic dataof anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, thecombined part and counterpart with most of dermal skeleton rendered branchial plate;cran.cart.,cranialcartilages;disp.,displaced;frag.,fragment; transparent:anatomical ventralview (corresponding tothe visible areaof L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., thepart in epoxy) (c) andanatomical dorsal view (buried in matrixin the vasculature;?,probable.Scale bar applies toall panels." figureDoi="http://doi.org/10.5281/zenodo.8368405" httpUri="https://zenodo.org/record/8368405/files/figure.png" pageId="1" pageNumber="2">Fig. 1</figureCitation>
|
||
). Denison
|
||
<bibRefCitation id="EFF8F8E8FFD4FF92FBADFA139E5E44B0" author="Denison, R. H." box="[1027,1035,1476,1489]" journalOrPublisher="Fieldiana" pageId="1" pageNumber="2" pagination="131 - 192" part="16" refId="ref4498" refString="8. Denison, R. H. Ordovician vertebrates from Western United States. Fieldiana 16, 131 - 192 (1967" title="Ordovician vertebrates from Western United States" type="journal article" year="1967">
|
||
<emphasis id="B91D590BFFD4FF92FBADFA139E5E44B0" bold="true" box="[1027,1035,1476,1489]" pageId="1" pageNumber="2">
|
||
<superScript id="7C1C2851FFD4FF92FBADFA139E5E44B0" attach="none" box="[1027,1035,1476,1489]" fontSize="5" pageId="1" pageNumber="2">8</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
described several large elements of globular calcified cartilage as part of the internal skeleton of
|
||
<taxonomicName id="4C69FE9AFFD4FF92FA93FA329FFB4499" authorityName="PF" authorityYear="1795" box="[1341,1454,1509,1528]" family="Eriptychiidae" genus="Eriptychius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD4FF92FA93FA329FFB4499" bold="true" box="[1341,1454,1509,1528]" italics="true" pageId="1" pageNumber="2">Eriptychius</emphasis>
|
||
</taxonomicName>
|
||
and we have been able to distinguish ten separate cartilages in total comprising the endoskeletal cranium (
|
||
<figureCitation id="1352999CFFD4FF92FB3CF9C99F454753" box="[1170,1296,1566,1587]" captionStart="Fig" captionStartId="1.[106,137,1122,1139]" captionTargetBox="[348,1251,134,1092]" captionTargetId="graphics-459@1.[389,1192,163,1056]" captionText="Fig.1 | Overviewof Eriptychiusamericanus PF 1795.a,b, Photographsof part counterpart) (d). Colourscheme forrenders:blue-greys,cranialcartilages PF1795a,which hadthe splitface set inepoxyandwas manually prepared (a), and (matchingthe detailed scheme inFig.2); transparencies,thedermal skeleton; itscounterpart PF 1795b,which remainsinrock matrix(b). Both are shown inan orange,branchial plates;red,orbital plates.Anterior totop ina–d.ant.tess., anatomically ventral view.c,d, Digital model of computed tomographic dataof anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, thecombined part and counterpart with most of dermal skeleton rendered branchial plate;cran.cart.,cranialcartilages;disp.,displaced;frag.,fragment; transparent:anatomical ventralview (corresponding tothe visible areaof L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., thepart in epoxy) (c) andanatomical dorsal view (buried in matrixin the vasculature;?,probable.Scale bar applies toall panels." figureDoi="http://doi.org/10.5281/zenodo.8368405" httpUri="https://zenodo.org/record/8368405/files/figure.png" pageId="1" pageNumber="2">Figs. 1 and 2</figureCitation>
|
||
and Extended
|
||
<taxonomicName id="4C69FE9AFFD4FF92FA06F9C99F8C4753" authorityName="Walker" authorityYear="1862" box="[1448,1497,1566,1586]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD4FF92FC9FF9ED9925472E" box="[817,880,1594,1615]" captionStart="Fig" captionStartId="1.[106,137,1122,1139]" captionTargetBox="[348,1251,134,1092]" captionTargetId="graphics-459@1.[389,1192,163,1056]" captionText="Fig.1 | Overviewof Eriptychiusamericanus PF 1795.a,b, Photographsof part counterpart) (d). Colourscheme forrenders:blue-greys,cranialcartilages PF1795a,which hadthe splitface set inepoxyandwas manually prepared (a), and (matchingthe detailed scheme inFig.2); transparencies,thedermal skeleton; itscounterpart PF 1795b,which remainsinrock matrix(b). Both are shown inan orange,branchial plates;red,orbital plates.Anterior totop ina–d.ant.tess., anatomically ventral view.c,d, Digital model of computed tomographic dataof anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, thecombined part and counterpart with most of dermal skeleton rendered branchial plate;cran.cart.,cranialcartilages;disp.,displaced;frag.,fragment; transparent:anatomical ventralview (corresponding tothe visible areaof L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., thepart in epoxy) (c) andanatomical dorsal view (buried in matrixin the vasculature;?,probable.Scale bar applies toall panels." figureDoi="http://doi.org/10.5281/zenodo.8368405" httpUri="https://zenodo.org/record/8368405/files/figure.png" pageId="1" pageNumber="2">Figs. 2</figureCitation>
|
||
and
|
||
<figureCitation id="1352999CFFD4FF92FC30F9EC99F8472E" box="[926,941,1595,1615]" captionStart="fragment" captionStartId="10.[816,898,1768,1786]" captionTargetBox="[127,1483,120,1744]" captionTargetId="figure-3@10.[226,1377,131,1744]" captionText="fragment inlateral view.i, rightbranchialplate inlateral view.Colour scheme indorsal view.c,chain of orbital plates in visceral view,showing curved visceral and abbreviations as in Figs.1and 2,except:grey,cartilages;blue-green, surfaces.d,e,the best-visualizedorbital plate in(d) lateral and(e) visceral view. dermal skeleton.Lighter shades denote material from PF 1795a,darker shades f,g,left branchial plate in(f) lateral and (g)anterior view.h,?R.branchial plate PF 1795b." pageId="1" pageNumber="2">4</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8BD68519FFD4FF92FCE9F9809F1246A2" blockId="1.[815,1498,1394,1987]" pageId="1" pageNumber="2">
|
||
Six cartilages were identified on the split surface of PF1795a by ref.8 (Extended
|
||
<taxonomicName id="4C69FE9AFFD4FF92FC31F9A3999A47E9" authorityName="Walker" authorityYear="1862" box="[927,975,1652,1672]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD4FF92FC7DF9A39E5F47E9" box="[979,1034,1652,1673]" captionStart="Fig" captionStartId="1.[106,137,1122,1139]" captionTargetBox="[348,1251,134,1092]" captionTargetId="graphics-459@1.[389,1192,163,1056]" captionText="Fig.1 | Overviewof Eriptychiusamericanus PF 1795.a,b, Photographsof part counterpart) (d). Colourscheme forrenders:blue-greys,cranialcartilages PF1795a,which hadthe splitface set inepoxyandwas manually prepared (a), and (matchingthe detailed scheme inFig.2); transparencies,thedermal skeleton; itscounterpart PF 1795b,which remainsinrock matrix(b). Both are shown inan orange,branchial plates;red,orbital plates.Anterior totop ina–d.ant.tess., anatomically ventral view.c,d, Digital model of computed tomographic dataof anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, thecombined part and counterpart with most of dermal skeleton rendered branchial plate;cran.cart.,cranialcartilages;disp.,displaced;frag.,fragment; transparent:anatomical ventralview (corresponding tothe visible areaof L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., thepart in epoxy) (c) andanatomical dorsal view (buried in matrixin the vasculature;?,probable.Scale bar applies toall panels." figureDoi="http://doi.org/10.5281/zenodo.8368405" httpUri="https://zenodo.org/record/8368405/files/figure.png" pageId="1" pageNumber="2">Fig. 2</figureCitation>
|
||
) and the remaining four are buried within the matrix of PF 1795b. The cartilages are closely wrapped by articulated squamation anteriorly,dorsally and ventrally and to one side (Extended
|
||
<taxonomicName id="4C69FE9AFFD4FF92FC81F91D990847BF" authorityName="Walker" authorityYear="1862" box="[815,861,1738,1758]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD4FF92FCCEF91D99C847BF" box="[864,925,1738,1759]" captionStart="Fig" captionStartId="3.[106,137,701,718]" captionTargetBox="[313,1277,120,680]" captionTargetId="graphics-692@3.[373,1255,157,437]" captionTargetPageId="3" captionText="Fig. 3| The neurocrania of vertebrates in a phylogenetic framework. a, Haikouichthys, adapted fromref.3, CC-BY 4.0.b, Petromyzon.c, Generalized cyathaspid,adapted from ref.20, Springer Nature.d, Sacabambaspis, reproduced fromref.31, copyright ©1993 Elsevier Masson SAS,all rights reserved (but see ref.53 for alternative interpretation).e, Eriptychius. f, Shuyu,adapted from ref.7, SpringerNature.g, Dicksonosteus adapted with permission from ref.14,Muséum national d’histoire naturelle,Paris. Phylogenetic schemeis the Adams consensus tree described inthe text,bold taxon namesrepresent thosedepicted above.Coloured regions illustrate positions of key sensory organs:yellow,eyes;blue,pineal.Light grey lines represent body outlines,black lines represent dermalbody armour,lightgrey" figureDoi="http://doi.org/10.5281/zenodo.8368409" httpUri="https://zenodo.org/record/8368409/files/figure.png" pageId="1" pageNumber="2">Figs.3</figureCitation>
|
||
and
|
||
<figureCitation id="1352999CFFD4FF92FC66F91D998247BF" box="[968,983,1738,1758]" captionStart="fragment" captionStartId="10.[816,898,1768,1786]" captionTargetBox="[127,1483,120,1744]" captionTargetId="figure-3@10.[226,1377,131,1744]" captionText="fragment inlateral view.i, rightbranchialplate inlateral view.Colour scheme indorsal view.c,chain of orbital plates in visceral view,showing curved visceral and abbreviations as in Figs.1and 2,except:grey,cartilages;blue-green, surfaces.d,e,the best-visualizedorbital plate in(d) lateral and(e) visceral view. dermal skeleton.Lighter shades denote material from PF 1795a,darker shades f,g,left branchial plate in(f) lateral and (g)anterior view.h,?R.branchial plate PF 1795b." pageId="1" pageNumber="2">4</figureCitation>
|
||
); however,there is a clear separation between dermal and endoskeletal elements, unlike in galeaspids and osteostracans
|
||
<bibRefCitation id="EFF8F8E8FFD4FF92FA6AF9349F864791" author="Wang, N. & Donoghue, P. C. J. & Smith, M. M. & Sansom, I. J." box="[1476,1491,1763,1776]" journalOrPublisher="Histology of the galeaspid dermoskeleton and endoskeleton and the origin and early evolution of the vertebrate cranial endoskeleton. J. Vertebr. Paleontol." pageId="1" pageNumber="2" pagination="745 - 756" part="25" refId="ref5533" refString="36. Wang, N., Donoghue, P. C. J., Smith, M. M., Sansom, I. J. & Nian-zhong, W. Histology of the galeaspid dermoskeleton and endoskeleton and the origin and early evolution of the vertebrate cranial endoskeleton. J. Vertebr. Paleontol. 25, 745 - 756 (2005)." title="Nian-zhong, W" type="journal article" year="2005">
|
||
<emphasis id="B91D590BFFD4FF92FA6AF9349F864791" bold="true" box="[1476,1491,1763,1776]" pageId="1" pageNumber="2">
|
||
<superScript id="7C1C2851FFD4FF92FA6AF9349F864791" attach="right" box="[1476,1491,1763,1776]" fontSize="5" pageId="1" pageNumber="2">36</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
. This squamation comprises a range of dermal element types including the scale types identified by ref.8 and scales with an anteroposteriorly oriented ornament from farther back on the head.It also includes small, curved orbital plates(
|
||
<figureCitation id="1352999CFFD4FF92FBAEF88E9E04460C" box="[1024,1105,1881,1902]" captionStart="Fig" captionStartId="1.[106,137,1122,1139]" captionTargetBox="[348,1251,134,1092]" captionTargetId="graphics-459@1.[389,1192,163,1056]" captionText="Fig.1 | Overviewof Eriptychiusamericanus PF 1795.a,b, Photographsof part counterpart) (d). Colourscheme forrenders:blue-greys,cranialcartilages PF1795a,which hadthe splitface set inepoxyandwas manually prepared (a), and (matchingthe detailed scheme inFig.2); transparencies,thedermal skeleton; itscounterpart PF 1795b,which remainsinrock matrix(b). Both are shown inan orange,branchial plates;red,orbital plates.Anterior totop ina–d.ant.tess., anatomically ventral view.c,d, Digital model of computed tomographic dataof anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, thecombined part and counterpart with most of dermal skeleton rendered branchial plate;cran.cart.,cranialcartilages;disp.,displaced;frag.,fragment; transparent:anatomical ventralview (corresponding tothe visible areaof L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., thepart in epoxy) (c) andanatomical dorsal view (buried in matrixin the vasculature;?,probable.Scale bar applies toall panels." figureDoi="http://doi.org/10.5281/zenodo.8368405" httpUri="https://zenodo.org/record/8368405/files/figure.png" pageId="1" pageNumber="2">Fig. 1c,d</figureCitation>
|
||
and Extended
|
||
<taxonomicName id="4C69FE9AFFD4FF92FB75F88E9F5C460C" authorityName="Walker" authorityYear="1862" box="[1243,1289,1881,1901]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD4FF92FAA2F88E9F36460C" box="[1292,1379,1881,1902]" captionStart="Fig" captionStartId="3.[106,137,701,718]" captionTargetBox="[313,1277,120,680]" captionTargetId="graphics-692@3.[373,1255,157,437]" captionTargetPageId="3" captionText="Fig. 3| The neurocrania of vertebrates in a phylogenetic framework. a, Haikouichthys, adapted fromref.3, CC-BY 4.0.b, Petromyzon.c, Generalized cyathaspid,adapted from ref.20, Springer Nature.d, Sacabambaspis, reproduced fromref.31, copyright ©1993 Elsevier Masson SAS,all rights reserved (but see ref.53 for alternative interpretation).e, Eriptychius. f, Shuyu,adapted from ref.7, SpringerNature.g, Dicksonosteus adapted with permission from ref.14,Muséum national d’histoire naturelle,Paris. Phylogenetic schemeis the Adams consensus tree described inthe text,bold taxon namesrepresent thosedepicted above.Coloured regions illustrate positions of key sensory organs:yellow,eyes;blue,pineal.Light grey lines represent body outlines,black lines represent dermalbody armour,lightgrey" figureDoi="http://doi.org/10.5281/zenodo.8368409" httpUri="https://zenodo.org/record/8368409/files/figure.png" pageId="1" pageNumber="2">Fig.3c–e</figureCitation>
|
||
) and several plates similar in morphology(
|
||
<figureCitation id="1352999CFFD4FF92FBFBF8A19EFD46EB" box="[1109,1192,1910,1931]" captionStart="Fig" captionStartId="1.[106,137,1122,1139]" captionTargetBox="[348,1251,134,1092]" captionTargetId="graphics-459@1.[389,1192,163,1056]" captionText="Fig.1 | Overviewof Eriptychiusamericanus PF 1795.a,b, Photographsof part counterpart) (d). Colourscheme forrenders:blue-greys,cranialcartilages PF1795a,which hadthe splitface set inepoxyandwas manually prepared (a), and (matchingthe detailed scheme inFig.2); transparencies,thedermal skeleton; itscounterpart PF 1795b,which remainsinrock matrix(b). Both are shown inan orange,branchial plates;red,orbital plates.Anterior totop ina–d.ant.tess., anatomically ventral view.c,d, Digital model of computed tomographic dataof anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, thecombined part and counterpart with most of dermal skeleton rendered branchial plate;cran.cart.,cranialcartilages;disp.,displaced;frag.,fragment; transparent:anatomical ventralview (corresponding tothe visible areaof L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., thepart in epoxy) (c) andanatomical dorsal view (buried in matrixin the vasculature;?,probable.Scale bar applies toall panels." figureDoi="http://doi.org/10.5281/zenodo.8368405" httpUri="https://zenodo.org/record/8368405/files/figure.png" pageId="1" pageNumber="2">Fig. 1c,d</figureCitation>
|
||
and Extended
|
||
<taxonomicName id="4C69FE9AFFD4FF92FA99F8A19F3346EB" authorityName="Walker" authorityYear="1862" box="[1335,1382,1910,1930]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="1" pageNumber="2" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD4FF92FAC7F8A19FEF46EB" box="[1385,1466,1910,1931]" captionStart="Fig" captionStartId="3.[106,137,701,718]" captionTargetBox="[313,1277,120,680]" captionTargetId="graphics-692@3.[373,1255,157,437]" captionTargetPageId="3" captionText="Fig. 3| The neurocrania of vertebrates in a phylogenetic framework. a, Haikouichthys, adapted fromref.3, CC-BY 4.0.b, Petromyzon.c, Generalized cyathaspid,adapted from ref.20, Springer Nature.d, Sacabambaspis, reproduced fromref.31, copyright ©1993 Elsevier Masson SAS,all rights reserved (but see ref.53 for alternative interpretation).e, Eriptychius. f, Shuyu,adapted from ref.7, SpringerNature.g, Dicksonosteus adapted with permission from ref.14,Muséum national d’histoire naturelle,Paris. Phylogenetic schemeis the Adams consensus tree described inthe text,bold taxon namesrepresent thosedepicted above.Coloured regions illustrate positions of key sensory organs:yellow,eyes;blue,pineal.Light grey lines represent body outlines,black lines represent dermalbody armour,lightgrey" figureDoi="http://doi.org/10.5281/zenodo.8368409" httpUri="https://zenodo.org/record/8368409/files/figure.png" pageId="1" pageNumber="2">Fig.3f–i</figureCitation>
|
||
) to an isolated
|
||
<taxonomicName id="4C69FE9AFFD4FF92FC0DF8449E4046C7" authorityName="PF" authorityYear="1795" box="[931,1045,1939,1958]" family="Eriptychiidae" genus="Eriptychius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="1" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD4FF92FC0DF8449E4046C7" bold="true" box="[931,1045,1939,1958]" italics="true" pageId="1" pageNumber="2">Eriptychius</emphasis>
|
||
</taxonomicName>
|
||
‘branchio-cornual plate’ identified by ref. 37 (plate 2, 4–7). There is no obvious pineal dermal plate.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<caption id="DF16D591FFD7FF91FFC4FC5499C0450B" ID-DOI="http://doi.org/10.5281/zenodo.8368407" ID-Zenodo-Dep="8368407" httpUri="https://zenodo.org/record/8368407/files/figure.png" pageId="2" pageNumber="3" startId="2.[106,137,899,916]" subCaptionStartIDs="2.[552,581,1138,1156]" subCaptionStarts="Fig.1 w" targetBox="[290,1301,124,880]" targetPageId="2">
|
||
<paragraph id="8BD68519FFD7FF91FFC4FC5498BA45E5" blockId="2.[106,780,898,1156]" pageId="2" pageNumber="3">
|
||
Fig. 2| The neurocranial cartilages of
|
||
<taxonomicName id="4C69FE9AFFD7FF91FE1BFC55982F42F5" authority="PF 1795" authorityName=", PF" authorityYear="1795" box="[437,634,898,916]" family="Eriptychiidae" genus="Eriptychius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="2" pageNumber="3" phylum="Chordata" rank="species" species="americanus">
|
||
<emphasis id="B91D590BFFD7FF91FE1BFC55986342F5" bold="true" box="[437,566,898,916]" italics="true" pageId="2" pageNumber="3">E.americanus</emphasis>
|
||
PF 1795
|
||
</taxonomicName>
|
||
based on computed tomography data. a–c, Cranial cartilages in estimated life position, with cartilages coloured in pairs in dorsal (a), ventral (b)and anterior(c)view. d,e, Mediolateral cartilages A in dorsal view(d) and median dorsal cartilage in ventral view(e)rendered with a vertical height map texture to emphasize the surface topology.f, Reconstruction of the forebrain relative to the cranial cartilages using a lamprey as a model
|
||
<emphasis id="B91D590BFFD7FF91FE0DFBF79BED454B" bold="true" box="[419,440,1056,1066]" pageId="2" pageNumber="3">
|
||
<bibRefCitation id="EFF8F8E8FFD7FF91FE0DFBF79BFC454B" author="Marinelli, W. & Strenger, A." box="[419,425,1056,1066]" journalOrPublisher="Lieferung. Lampetra fluviatilis (Deuticke" pageId="2" pageNumber="3" refId="ref4523" refString="9. Marinelli, W. & Strenger, A. Vergleichende Anatomie und Morphologie der Wirbeltiere. I. Lieferung. Lampetra fluviatilis (Deuticke, 1954)." title="Vergleichende Anatomie und Morphologie der Wirbeltiere. I" type="book" year="1954">9</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFF8F8E8FFD7FF91FE02FBF79BED454B" author="Sugahara, F. & Murakami, Y. & Pascual-Anaya, J. & Kuratani, S." box="[428,440,1056,1066]" journalOrPublisher="Brain Behav. Evol." pageId="2" pageNumber="3" pagination="305 - 317" part="96" refId="ref6113" refString="52. Sugahara, F., Murakami, Y., Pascual-Anaya, J. & Kuratani, S. Forebrain architecture and development in cyclostomes, with reference to the early morphology and evolution of the vertebrate head. Brain Behav. Evol. 96, 305 - 317 (2022)." title="Forebrain architecture and development in cyclostomes, with reference to the early morphology and evolution of the vertebrate head" type="journal article" year="2022">52</bibRefCitation>
|
||
</emphasis>
|
||
,shown in dorsal view.g, Cartilages in dorsal view,rendered transparent to show internal vasculature(red). h,i, Cartilages in preserved position in anterior view with dermal skeleton shown(h)and removed (i). Colours in a,b,c,f,h,i as in Fig.1 with the following
|
||
</paragraph>
|
||
<paragraph id="8BD68519FFD7FF91FC9EFC5599C0450B" blockId="2.[816,1494,898,1130]" pageId="2" pageNumber="3">additions.Green,dermal skeleton.Red dashed line represents inferred position of mouth in c,h,i. In d and e lighter colours denote areas closer to the camera. Abbreviations as in Fig.1with the following additions:antorb.proc,antorbital process;ext.vasc.op.,external vascular openings;forebr.,forebrain;lat., lateral;medlat.cart,mediolateral cartilage;med.dors.cart,median dorsal cartilage;med.,medial;med.vent.cart.,median ventral cartilage;med.vent. ridge,median ventral ridge;olf.bulb,olfactory bulb;pin.,pineal organ;pin.op., pineal opening;vent.,ventral.Scale bar in a is shared by b,c; scale bar in d is shared by e.</paragraph>
|
||
</caption>
|
||
<subSubSection id="C373D692FFD7FF90FF2FFB359BE94655" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3" type="discussion">
|
||
<paragraph id="8BD68519FFD7FF91FF2FFB359B974676" blockId="2.[105,787,1250,1988]" pageId="2" pageNumber="3">
|
||
Denison identified two cartilages as the orbital cartilages on the basis of concave posterior faces
|
||
<bibRefCitation id="EFF8F8E8FFD7FF91FEDAFB2B9B294468" author="Denison, R. H." box="[372,380,1276,1289]" journalOrPublisher="Fieldiana" pageId="2" pageNumber="3" pagination="131 - 192" part="16" refId="ref4498" refString="8. Denison, R. H. Ordovician vertebrates from Western United States. Fieldiana 16, 131 - 192 (1967" title="Ordovician vertebrates from Western United States" type="journal article" year="1967">
|
||
<emphasis id="B91D590BFFD7FF91FEDAFB2B9B294468" bold="true" box="[372,380,1276,1289]" pageId="2" pageNumber="3">8</emphasis>
|
||
</bibRefCitation>
|
||
. Our scan data confirm that these represent cross-sections through large fossae that we interpret as forming the anterior wall of the orbits.These fossae are flanked laterally by an antorbital process (Fig. 2a,b and Extended
|
||
<taxonomicName id="4C69FE9AFFD7FF91FDA0FA8298684408" authorityName="Walker" authorityYear="1862" box="[526,573,1365,1385]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD7FF91FDEEFA8298204408" box="[576,629,1365,1386]" captionStart="cartilages B" captionStartId="11.[816,902,1768,1786]" captionTargetBox="[125,1477,131,1744]" captionTargetId="figure-7@11.[125,1477,131,1744]" captionText="cartilages B;i,j bothsurfaces of mediolateral cartilages C; k-m,left orbital cartilage in(k) medial,(l) posterior and(m) latero-posterior view.Colour scheme andabbreviations as inFig.2." pageId="2" pageNumber="3">Fig.5</figureCitation>
|
||
) that suggests a dorsolateral orientation of the orbit. A smaller ventral fossa on each orbital cartilage below the orbit may have provided a location for muscle attachment (Fig. 2). One orbital cartilage is posteriorly displaced, along with elements of anterior squamation including a rostral scale (Fig. 1 and Extended
|
||
<taxonomicName id="4C69FE9AFFD7FF91FE98FA329B304498" authorityName="Walker" authorityYear="1862" box="[310,357,1509,1529]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD7FF91FEC7FA339BCA4498" box="[361,415,1508,1529]" captionStart="Fig" captionStartId="3.[106,137,701,718]" captionTargetBox="[313,1277,120,680]" captionTargetId="graphics-692@3.[373,1255,157,437]" captionTargetPageId="3" captionText="Fig. 3| The neurocrania of vertebrates in a phylogenetic framework. a, Haikouichthys, adapted fromref.3, CC-BY 4.0.b, Petromyzon.c, Generalized cyathaspid,adapted from ref.20, Springer Nature.d, Sacabambaspis, reproduced fromref.31, copyright ©1993 Elsevier Masson SAS,all rights reserved (but see ref.53 for alternative interpretation).e, Eriptychius. f, Shuyu,adapted from ref.7, SpringerNature.g, Dicksonosteus adapted with permission from ref.14,Muséum national d’histoire naturelle,Paris. Phylogenetic schemeis the Adams consensus tree described inthe text,bold taxon namesrepresent thosedepicted above.Coloured regions illustrate positions of key sensory organs:yellow,eyes;blue,pineal.Light grey lines represent body outlines,black lines represent dermalbody armour,lightgrey" figureDoi="http://doi.org/10.5281/zenodo.8368409" httpUri="https://zenodo.org/record/8368409/files/figure.png" pageId="2" pageNumber="3">Fig. 3</figureCitation>
|
||
) and has rotated by about 180° in the sagittal axis;when rotated back into position it aligns with the orbital plates (Fig. 1). The anteriormost branchial plate lies posterolateral to the other orbital cartilage (Fig. 1 and Extended
|
||
<taxonomicName id="4C69FE9AFFD7FF91FDEFF9EC9824472E" authorityName="Walker" authorityYear="1862" box="[577,625,1595,1615]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD7FF91FDDAF9ED98F9472E" box="[628,684,1594,1615]" captionStart="Fig" captionStartId="3.[106,137,701,718]" captionTargetBox="[313,1277,120,680]" captionTargetId="graphics-692@3.[373,1255,157,437]" captionTargetPageId="3" captionText="Fig. 3| The neurocrania of vertebrates in a phylogenetic framework. a, Haikouichthys, adapted fromref.3, CC-BY 4.0.b, Petromyzon.c, Generalized cyathaspid,adapted from ref.20, Springer Nature.d, Sacabambaspis, reproduced fromref.31, copyright ©1993 Elsevier Masson SAS,all rights reserved (but see ref.53 for alternative interpretation).e, Eriptychius. f, Shuyu,adapted from ref.7, SpringerNature.g, Dicksonosteus adapted with permission from ref.14,Muséum national d’histoire naturelle,Paris. Phylogenetic schemeis the Adams consensus tree described inthe text,bold taxon namesrepresent thosedepicted above.Coloured regions illustrate positions of key sensory organs:yellow,eyes;blue,pineal.Light grey lines represent body outlines,black lines represent dermalbody armour,lightgrey" figureDoi="http://doi.org/10.5281/zenodo.8368409" httpUri="https://zenodo.org/record/8368409/files/figure.png" pageId="2" pageNumber="3">Fig. 3</figureCitation>
|
||
), suggesting that the relative positions of the orbits,otic region and branchial arches were similar to those inferred in heterostracans
|
||
<emphasis id="B91D590BFFD7FF91FD3DF9A798FE471C" bold="true" box="[659,683,1648,1661]" pageId="2" pageNumber="3">
|
||
<bibRefCitation id="EFF8F8E8FFD7FF91FD3DF9A798CF471C" author="Janvier, P." box="[659,666,1648,1661]" journalOrPublisher="Clarendon" pageId="2" pageNumber="3" refId="ref4201" refString="2. Janvier, P. Early Vertebrates (Clarendon, 1996)" title="Early Vertebrates" type="book" year="1996">2</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFF8F8E8FFD7FF91FD30F9A798FE471C" author="Stensio, E." box="[670,683,1648,1661]" editor="Piveteau, J." journalOrPublisher="Masson" pageId="2" pageNumber="3" pagination="96 - 383" refId="ref4915" refString="19. Stensio, E. in Traite de Paleontologie (ed. Piveteau, J.) 96 - 383 (Masson, 1964)." title="Traite de Paleontologie" type="book chapter" year="1964">19</bibRefCitation>
|
||
</emphasis>
|
||
, although this is impossible to judge exactly because of the collapse of the dermal skeleton.The absence of anything assignable to the branchial skeleton suggests that the branchial arches were not incorporated into a single mineralization with the neurocranium,a major difference with respect to galeaspids and osteostracans
|
||
<emphasis id="B91D590BFFD7FF91FE09F8D79BE9466C" bold="true" box="[423,444,1792,1805]" pageId="2" pageNumber="3">
|
||
<bibRefCitation id="EFF8F8E8FFD7FF91FE09F8D79BF9466C" author="Gai, Z. & Donoghue, P. C. J. & Zhu, M. & Janvier, P. & Stampanoni, M." box="[423,428,1792,1805]" journalOrPublisher="Nature" pageId="2" pageNumber="3" pagination="324 - 327" part="476" refId="ref4346" refString="7. Gai, Z., Donoghue, P. C. J., Zhu, M., Janvier, P. & Stampanoni, M. Fossil jawless fish from China foreshadows early jawed vertebrate anatomy. Nature 476, 324 - 327 (2011)" title="Fossil jawless fish from China foreshadows early jawed vertebrate anatomy" type="journal article" year="2011">7</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFF8F8E8FFD7FF91FE1EF8D79BE9466C" author="Janvier, P." box="[432,444,1792,1805]" journalOrPublisher="Revision des Osteostraces de la Formation de Wood Bay (Devonien Inferieur du Spitsberg) (Cahiers d e Paleontologie) (Editions du CNRS" pageId="2" pageNumber="3" refId="ref4714" refString="15. Janvier, P. Les Cephalaspides du Spitsberg. Anatomie, Phylogenie et Systematique des Osteostraces Siluro-devoniens. Revision des Osteostraces de la Formation de Wood Bay (Devonien Inferieur du Spitsberg) (Cahiers d e Paleontologie) (Editions du CNRS, 1985)." title="Les Cephalaspides du Spitsberg. Anatomie, Phylogenie et Systematique des Osteostraces Siluro-devoniens" type="book" year="1985">15</bibRefCitation>
|
||
</emphasis>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="8BD68519FFD7FF91FF2FF8F79E1047A3" blockId="2.[105,787,1250,1988]" lastBlockId="2.[815,1498,1250,1987]" pageId="2" pageNumber="3">
|
||
The remaining cartilages lie between the two orbital cartilages, although they have slumped slightly and been pulled posteriorly on one side with the displaced orbital cartilage (Figs. 1 and 2a–c and Extended
|
||
<taxonomicName id="4C69FE9AFFD7FF91FF62F8A19AAE46EB" authorityName="Walker" authorityYear="1862" box="[204,251,1910,1930]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD7FF91FF50F8A19B6046EB" box="[254,309,1910,1931]" captionStart="fragment" captionStartId="10.[816,898,1768,1786]" captionTargetBox="[127,1483,120,1744]" captionTargetId="figure-3@10.[226,1377,131,1744]" captionText="fragment inlateral view.i, rightbranchialplate inlateral view.Colour scheme indorsal view.c,chain of orbital plates in visceral view,showing curved visceral and abbreviations as in Figs.1and 2,except:grey,cartilages;blue-green, surfaces.d,e,the best-visualizedorbital plate in(d) lateral and(e) visceral view. dermal skeleton.Lighter shades denote material from PF 1795a,darker shades f,g,left branchial plate in(f) lateral and (g)anterior view.h,?R.branchial plate PF 1795b." pageId="2" pageNumber="3">Fig. 4</figureCitation>
|
||
). There are three paired cartilages arranged symmetrically across the midline—two dorsal (termed mediolateral A, B) and one ventral (termed mediolateral C)—along with one unpaired midline cartilage dorsally and one ventrally.Of the two unpaired cartilages, the smaller dorsal element is kite-shaped and has both the dorsal and ventral surfaces punctured by large medial foramina that we consider likely to be the pineal opening (Fig. 2a,e). These dorsal and ventral foramina do not exactly line up anteroposteriorly but appear to communicate with a common large space inside the cartilage (Fig. 2g and Extended
|
||
<taxonomicName id="4C69FE9AFFD7FF91FBFFFA589ED444C2" authorityName="Walker" authorityYear="1862" box="[1105,1153,1423,1443]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD7FF91FB2BFA599EEB44C2" box="[1157,1214,1422,1443]" captionStart="cartilages B" captionStartId="12.[816,900,1607,1625]" captionTargetBox="[110,1479,128,1591]" captionTargetId="figure-4@12.[121,1481,131,1584]" captionText="cartilages B;h,mediolateralcartilages C;i,jleftorbitalcartilage in (i)dorsal and endocranium ofEriptychiusPF1795.a,b,mediandorsalcartilage in (a)dorsal (j) ventralview;k,l rightorbitalcartilage in(k) dorsaland (l)ventralview.Colour and(b)ventralview;c,d,medianventralcartilage in (c)ventraland(d)dorsalview; scheme as inFig.2. " pageId="2" pageNumber="3">Fig. 6</figureCitation>
|
||
). This kite-shaped cartilage is preserved overlying the mediolateral cartilages A and its ventral surface bears a median ridge with a shallow depression on either side (Fig. 2a,e). Together with shallow depressions on the dorsal surface of mediolateral cartilages A, these depressions frame paired fossae which we interpret as having accommodated part of the forebrain, possibly the olfactory bulbs (Fig.2d–f), an interpretation consistent with their position relative to the orbits and putative pineal opening. Thus,we infer this to be the dorsal side of the animal and term this the median dorsal cartilage.The larger median ventral cartilage underlies the mediolateral cartilages.
|
||
</paragraph>
|
||
<paragraph id="8BD68519FFD7FF90FCE9F91D994744E7" blockId="2.[815,1498,1250,1987]" lastBlockId="3.[105,788,1078,1844]" lastPageId="3" lastPageNumber="4" pageId="2" pageNumber="3">
|
||
All cartilages are pervasively penetrated by canals (Fig. 2g and Extended
|
||
<taxonomicName id="4C69FE9AFFD7FF91FC3AF9309996479A" authorityName="Walker" authorityYear="1862" box="[916,963,1767,1787]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="2" pageNumber="3" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD7FF91FC69F93199AB479A" box="[967,1022,1766,1787]" captionStart="cartilages B" captionStartId="12.[816,900,1607,1625]" captionTargetBox="[110,1479,128,1591]" captionTargetId="figure-4@12.[121,1481,131,1584]" captionText="cartilages B;h,mediolateralcartilages C;i,jleftorbitalcartilage in (i)dorsal and endocranium ofEriptychiusPF1795.a,b,mediandorsalcartilage in (a)dorsal (j) ventralview;k,l rightorbitalcartilage in(k) dorsaland (l)ventralview.Colour and(b)ventralview;c,d,medianventralcartilage in (c)ventraland(d)dorsalview; scheme as inFig.2. " pageId="2" pageNumber="3">Fig. 6</figureCitation>
|
||
). In the larger cartilages,that is,the orbital cartilages and the mediolateral cartilages A,this tends to follow the pattern of a larger trunk entering the cartilage from the posterior side before splitting into smaller branches that open to the surface.The pattern is notexactlybilaterallysymmetricalinthepairedandunpairedelements but does follow a similar organization with the trunk canal entering at equivalent points. These canals could plausibly have carried sensory rami onto the surface of the head; for example, the superficial ophthalmic nerve in the case of the canals in the orbital cartilages. However,the canal openings are not restricted to surfaces where the cartilages contact the dermal skeleton.On the basis of this and the lack of sensory canal openings in the head tesserae,they may have carried vasculature instead or as well.In living chondrichthyans,canals carry vasculatureandtransportprechondrocytesintothecartilagefromthe perichondral surface.This could implicate the canals in
|
||
<taxonomicName id="4C69FE9AFFD6FF90FD22FB3498AF4597" authorityName="PF" authorityYear="1795" box="[652,762,1251,1270]" family="Eriptychiidae" genus="Eriptychius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FD22FB3498AF4597" bold="true" box="[652,762,1251,1270]" italics="true" pageId="3" pageNumber="4">Eriptychius</emphasis>
|
||
</taxonomicName>
|
||
in both cartilage maintenance and interstitial cartilage growth,although in modern chondrichthyans the width of these canals are an order of magnitudesmallerthanin
|
||
<taxonomicName id="4C69FE9AFFD6FF90FECDFAEE9B98442D" authorityName="PF" authorityYear="1795" box="[355,461,1337,1356]" family="Eriptychiidae" genus="Eriptychius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FECDFAEE9B98442D" bold="true" box="[355,461,1337,1356]" italics="true" pageId="3" pageNumber="4">Eriptychius</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="B91D590BFFD6FF90FE62FAE29BBE4423" bold="true" box="[460,491,1333,1346]" pageId="3" pageNumber="4">
|
||
<superScript id="7C1C2851FFD6FF90FE62FAE29BBE4423" attach="left" box="[460,491,1333,1346]" fontSize="5" pageId="3" pageNumber="4">
|
||
<bibRefCitation id="EFF8F8E8FFD6FF90FE62FAE29B8F4423" author="Dean, M. N. & Socha, J. J. & Hall, B. K. & Summers, A. P." box="[460,474,1333,1346]" journalOrPublisher="J. Appl. Ichthyol." pageId="3" pageNumber="4" pagination="263 - 267" part="26" refId="ref5642" refString="38. Dean, M. N., Socha, J. J., Hall, B. K. & Summers, A. P. Canaliculi in the tessellated skeleton of cartilaginous fishes. J. Appl. Ichthyol. 26, 263 - 267 (2010)." title="Canaliculi in the tessellated skeleton of cartilaginous fishes" type="journal article" year="2010">38</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="EFF8F8E8FFD6FF90FE70FAE29BBE4423" author="Marconi, A. & Hancock-Ronemus, A. & Gillis, J. A." box="[478,491,1333,1346]" journalOrPublisher="Leucoraja erinacea. eLife" pageId="3" pageNumber="4" pagination="53414" part="9" refId="ref5695" refString="39. Marconi, A., Hancock-Ronemus, A. & Gillis, J. A. Adult chondrogenesis and spontaneous cartilage repair in the skate, Leucoraja erinacea. eLife 9, e 53414 (2020)." title="Adult chondrogenesis and spontaneous cartilage repair in the skate" type="journal article" year="2020">39</bibRefCitation>
|
||
</superScript>
|
||
</emphasis>
|
||
.Althoughcanalshavenotbeen explicitly reported in other early vertebrate cartilages,sections through galeaspid cartilage suggest that this tissue has a degree of vascularity
|
||
<bibRefCitation id="EFF8F8E8FFD6FF90FD50FAB99958441A" author="Wang, N. & Donoghue, P. C. J. & Smith, M. M. & Sansom, I. J." box="[766,781,1390,1403]" journalOrPublisher="Histology of the galeaspid dermoskeleton and endoskeleton and the origin and early evolution of the vertebrate cranial endoskeleton. J. Vertebr. Paleontol." pageId="3" pageNumber="4" pagination="745 - 756" part="25" refId="ref5533" refString="36. Wang, N., Donoghue, P. C. J., Smith, M. M., Sansom, I. J. & Nian-zhong, W. Histology of the galeaspid dermoskeleton and endoskeleton and the origin and early evolution of the vertebrate cranial endoskeleton. J. Vertebr. Paleontol. 25, 745 - 756 (2005)." title="Nian-zhong, W" type="journal article" year="2005">
|
||
<emphasis id="B91D590BFFD6FF90FD50FAB99958441A" bold="true" box="[766,781,1390,1403]" pageId="3" pageNumber="4">
|
||
<superScript id="7C1C2851FFD6FF90FD50FAB99958441A" attach="right" box="[766,781,1390,1403]" fontSize="5" pageId="3" pageNumber="4">36</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
<emphasis id="B91D590BFFD6FF90FCA2FAA5994744E7" bold="true" box="[780,786,1394,1414]" pageId="3" pageNumber="4">.</emphasis>
|
||
</paragraph>
|
||
<caption id="DF16D591FFD6FF90FFC4FD6A9E0A42B8" ID-DOI="http://doi.org/10.5281/zenodo.8368409" ID-Zenodo-Dep="8368409" httpUri="https://zenodo.org/record/8368409/files/figure.png" pageId="3" pageNumber="4" startId="3.[106,137,701,718]" targetBox="[313,1277,120,680]" targetPageId="3">
|
||
<paragraph id="8BD68519FFD6FF90FFC4FD6A995D42B9" blockId="3.[106,778,701,985]" pageId="3" pageNumber="4">
|
||
Fig. 3| The neurocrania of vertebrates in a phylogenetic framework. a,
|
||
<taxonomicName id="4C69FE9AFFD6FF90FFD2FD009AAD4389" authorityName="Luo, Hu & Shu" authorityYear="1999" box="[124,248,727,744]" family="Myllokunmingiidae" genus="Haikouichthys" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FFD2FD009AAD4389" bold="true" box="[124,248,727,744]" italics="true" pageId="3" pageNumber="4">Haikouichthys</emphasis>
|
||
</taxonomicName>
|
||
, adapted from ref.3, CC-BY 4.0.b,
|
||
<emphasis id="B91D590BFFD6FF90FDB1FD0098D94389" bold="true" box="[543,652,727,744]" italics="true" pageId="3" pageNumber="4">
|
||
<taxonomicName id="4C69FE9AFFD6FF90FDB1FD0098DD4389" authorityName="Linnaeus" authorityYear="1758" box="[543,648,727,744]" class="Petromyzonti" family="Petromyzontidae" genus="Petromyzon" kingdom="Animalia" order="Petromyzontiformes" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Petromyzon</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
c, Generalized cyathaspid,adapted from ref.20, Springer Nature.d,
|
||
<taxonomicName id="4C69FE9AFFD6FF90FD99FD2598944262" authorityName="Gagnier, Blieck & Rodrigo" authorityYear="1986" box="[567,705,754,771]" family="Arandaspididae" genus="Sacabambaspis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FD99FD2598944262" bold="true" box="[567,705,754,771]" italics="true" pageId="3" pageNumber="4">Sacabambaspis</emphasis>
|
||
</taxonomicName>
|
||
, reproduced from ref.31, copyright ©1993 Elsevier Masson SAS,all rights reserved (but see ref.53 for alternative interpretation).e,
|
||
<emphasis id="B91D590BFFD6FF90FDF2FCF098904259" bold="true" box="[604,709,807,824]" italics="true" pageId="3" pageNumber="4">
|
||
<taxonomicName id="4C69FE9AFFD6FF90FDF2FCF098944259" authorityName="PF" authorityYear="1795" box="[604,705,807,824]" family="Eriptychiidae" genus="Eriptychius" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">Eriptychius</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
f,
|
||
<taxonomicName id="4C69FE9AFFD6FF90FFD7FC959AFB4232" authorityName="Gai, Donoghue, Zhu, Janvier & Stampanoni" authorityYear="2011" box="[121,174,834,851]" genus="Shuyu" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FFD7FC959AFB4232" bold="true" box="[121,174,834,851]" italics="true" pageId="3" pageNumber="4">Shuyu</emphasis>
|
||
</taxonomicName>
|
||
,adapted from ref.7, Springer Nature.g,
|
||
<taxonomicName id="4C69FE9AFFD6FF90FDA6FC9598D14232" authorityName="Goujet" authorityYear="1975" box="[520,644,834,851]" family="Phlyctaeniidae" genus="Dicksonosteus" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FDA6FC9598D14232" bold="true" box="[520,644,834,851]" italics="true" pageId="3" pageNumber="4">Dicksonosteus</emphasis>
|
||
</taxonomicName>
|
||
adapted with permission from ref.14,Muséum national d’histoire naturelle,Paris. Phylogenetic scheme is the Adams consensus tree described in the text,bold taxon names represent those depicted above.Coloured regions illustrate positions of key sensory organs:yellow,eyes;blue,pineal.Light grey lines represent body outlines,black lines represent dermal body armour,light grey
|
||
</paragraph>
|
||
<paragraph id="8BD68519FFD6FF90FC9EFD6B9E0A42B8" blockId="3.[816,1496,700,985]" pageId="3" pageNumber="4">
|
||
regions with dark grey borders represent cranial mineralizations,brown regions represent the brain (b), imprints of the brain in the dorsal headshield (c) or the endocast (f,g). Square brackets represent the preorbital part of the head.
|
||
<taxonomicName id="4C69FE9AFFD6FF90FC2AFCDA99BF427F" authorityName="Linnaeus" authorityYear="1758" box="[900,1002,781,798]" class="Petromyzonti" family="Petromyzontidae" genus="Petromyzon" kingdom="Animalia" order="Petromyzontiformes" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FC2AFCDA99BF427F" bold="true" box="[900,1002,781,798]" italics="true" pageId="3" pageNumber="4">Petromyzon</emphasis>
|
||
</taxonomicName>
|
||
redrawn from Sketchfab (https://sketchfab.com/ 3d-models/lamprey-inside-c9f05e3f00c94d929e7ed018fac6d782) under a CC BY4.0 licence.Abbreviations:CG,crown group;mand.gnath.,mandibulate gnathostomes;TG,total group.*‘Crown-group cyclostomes’ represents the
|
||
<taxonomicName id="4C69FE9AFFD6FF90FC9EFCA1999C42E9" box="[816,969,886,904]" class="Petromyzonti" family="Petromyzontidae" kingdom="Animalia" order="Petromyzontiformes" pageId="3" pageNumber="4" phylum="Chordata" rank="family">Petromyzontidae</taxonomicName>
|
||
and Myxinoidea total groups and
|
||
<taxonomicName id="4C69FE9AFFD6FF90FB40FCA09F0542E9" authorityName="Bardack & Richardson" authorityYear="1977" box="[1262,1360,887,904]" class="Petromyzonti" family="Petromyzontidae" genus="Gilpichthys" higherTaxonomySource="GBIF" kingdom="Animalia" order="Petromyzontiformes" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FB40FCA09F0542E9" bold="true" box="[1262,1360,887,904]" italics="true" pageId="3" pageNumber="4">Gilpichthys</emphasis>
|
||
</taxonomicName>
|
||
, which was recovered in a polytomy with those two groups.**Cyclostome and gnathostome total groups in this topology recovered in a polytomy with
|
||
<taxonomicName id="4C69FE9AFFD6FF90FA87FC7A9FFC42DF" authorityName="Simonetta & Insom" authorityYear="1993" box="[1321,1449,941,958]" genus="Metaspriggina" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FA87FC7A9FFC42DF" bold="true" box="[1321,1449,941,958]" italics="true" pageId="3" pageNumber="4">Metaspriggina</emphasis>
|
||
</taxonomicName>
|
||
and (
|
||
<taxonomicName id="4C69FE9AFFD6FF90FC96FC1099E142B9" authorityName="Luo, Hu & Shu" authorityYear="1999" box="[824,948,967,984]" family="Myllokunmingiidae" genus="Haikouichthys" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FC96FC1099E142B9" bold="true" box="[824,948,967,984]" italics="true" pageId="3" pageNumber="4">Haikouichthys</emphasis>
|
||
</taxonomicName>
|
||
+
|
||
<taxonomicName id="4C69FE9AFFD6FF90FC68FC109E0142B9" authorityName="Shu, Zhang & Han" authorityYear="1999" box="[966,1108,967,984]" family="Myllokunmingiidae" genus="Myllokunmingia" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FC68FC109E0142B9" bold="true" box="[966,1108,967,984]" italics="true" pageId="3" pageNumber="4">Myllokunmingia</emphasis>
|
||
</taxonomicName>
|
||
).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BD68519FFD6FF90FF2FFA589BE94655" blockId="3.[105,788,1078,1844]" pageId="3" pageNumber="4">
|
||
In concert with the displacement of the cartilages,the dermal squamation has undergone postmortem collapse.we interpret the articulated area of squamation as having covered the right side of the head, around the region of the right orbit and the right side of the mouth,as well as the areas dorsal and ventral to the mouth (
|
||
<figureCitation id="1352999CFFD6FF90FDFEF9D698D04774" box="[592,645,1537,1558]" captionStart="Fig" captionStartId="1.[106,137,1122,1139]" captionTargetBox="[348,1251,134,1092]" captionTargetId="graphics-459@1.[389,1192,163,1056]" captionText="Fig.1 | Overviewof Eriptychiusamericanus PF 1795.a,b, Photographsof part counterpart) (d). Colourscheme forrenders:blue-greys,cranialcartilages PF1795a,which hadthe splitface set inepoxyandwas manually prepared (a), and (matchingthe detailed scheme inFig.2); transparencies,thedermal skeleton; itscounterpart PF 1795b,which remainsinrock matrix(b). Both are shown inan orange,branchial plates;red,orbital plates.Anterior totop ina–d.ant.tess., anatomically ventral view.c,d, Digital model of computed tomographic dataof anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, thecombined part and counterpart with most of dermal skeleton rendered branchial plate;cran.cart.,cranialcartilages;disp.,displaced;frag.,fragment; transparent:anatomical ventralview (corresponding tothe visible areaof L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., thepart in epoxy) (c) andanatomical dorsal view (buried in matrixin the vasculature;?,probable.Scale bar applies toall panels." figureDoi="http://doi.org/10.5281/zenodo.8368405" httpUri="https://zenodo.org/record/8368405/files/figure.png" pageId="3" pageNumber="4">Fig. 1</figureCitation>
|
||
and Extended
|
||
<taxonomicName id="4C69FE9AFFD6FF90FFC4F9C99AC24753" authorityName="Walker" authorityYear="1862" box="[106,151,1566,1586]" class="Insecta" family="Noctuidae" genus="Data" kingdom="Animalia" order="Lepidoptera" pageId="3" pageNumber="4" phylum="Arthropoda" rank="genus">Data</taxonomicName>
|
||
<figureCitation id="1352999CFFD6FF90FF34F9C99A9B4753" box="[154,206,1566,1587]" captionStart="Fig" captionStartId="3.[106,137,701,718]" captionTargetBox="[313,1277,120,680]" captionTargetId="graphics-692@3.[373,1255,157,437]" captionTargetPageId="3" captionText="Fig. 3| The neurocrania of vertebrates in a phylogenetic framework. a, Haikouichthys, adapted fromref.3, CC-BY 4.0.b, Petromyzon.c, Generalized cyathaspid,adapted from ref.20, Springer Nature.d, Sacabambaspis, reproduced fromref.31, copyright ©1993 Elsevier Masson SAS,all rights reserved (but see ref.53 for alternative interpretation).e, Eriptychius. f, Shuyu,adapted from ref.7, SpringerNature.g, Dicksonosteus adapted with permission from ref.14,Muséum national d’histoire naturelle,Paris. Phylogenetic schemeis the Adams consensus tree described inthe text,bold taxon namesrepresent thosedepicted above.Coloured regions illustrate positions of key sensory organs:yellow,eyes;blue,pineal.Light grey lines represent body outlines,black lines represent dermalbody armour,lightgrey" figureDoi="http://doi.org/10.5281/zenodo.8368409" httpUri="https://zenodo.org/record/8368409/files/figure.png" pageId="3" pageNumber="4">Fig.3</figureCitation>
|
||
).On the basis of this interpretation,the cartilages would have comprised the preorbital region of the head,surrounding the mouth (
|
||
<figureCitation id="1352999CFFD6FF90FFDEF9809A80470A" box="[112,213,1623,1644]" captionStart="Fig" captionStartId="1.[106,137,1122,1139]" captionTargetBox="[348,1251,134,1092]" captionTargetId="graphics-459@1.[389,1192,163,1056]" captionText="Fig.1 | Overviewof Eriptychiusamericanus PF 1795.a,b, Photographsof part counterpart) (d). Colourscheme forrenders:blue-greys,cranialcartilages PF1795a,which hadthe splitface set inepoxyandwas manually prepared (a), and (matchingthe detailed scheme inFig.2); transparencies,thedermal skeleton; itscounterpart PF 1795b,which remainsinrock matrix(b). Both are shown inan orange,branchial plates;red,orbital plates.Anterior totop ina–d.ant.tess., anatomically ventral view.c,d, Digital model of computed tomographic dataof anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, thecombined part and counterpart with most of dermal skeleton rendered branchial plate;cran.cart.,cranialcartilages;disp.,displaced;frag.,fragment; transparent:anatomical ventralview (corresponding tothe visible areaof L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., thepart in epoxy) (c) andanatomical dorsal view (buried in matrixin the vasculature;?,probable.Scale bar applies toall panels." figureDoi="http://doi.org/10.5281/zenodo.8368405" httpUri="https://zenodo.org/record/8368405/files/figure.png" pageId="3" pageNumber="4">Fig. 2c,h,i</figureCitation>
|
||
). The articulated dorsal and ventral patches of squamation indicate that the mouth opened between the cartilages,bordered by the ‘rostral’ plates
|
||
<bibRefCitation id="EFF8F8E8FFD6FF90FE95F95A9B1647FB" author="Denison, R. H." box="[315,323,1677,1690]" journalOrPublisher="Fieldiana" pageId="3" pageNumber="4" pagination="131 - 192" part="16" refId="ref4498" refString="8. Denison, R. H. Ordovician vertebrates from Western United States. Fieldiana 16, 131 - 192 (1967" title="Ordovician vertebrates from Western United States" type="journal article" year="1967">
|
||
<emphasis id="B91D590BFFD6FF90FE95F95A9B1647FB" bold="true" box="[315,323,1677,1690]" pageId="3" pageNumber="4">
|
||
<superScript id="7C1C2851FFD6FF90FE95F95A9B1647FB" attach="none" box="[315,323,1677,1690]" fontSize="5" pageId="3" pageNumber="4">8</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
and was oriented supraterminally (
|
||
<figureCitation id="1352999CFFD6FF90FD0CF947995147C5" box="[674,772,1680,1701]" captionStart="Fig" captionStartId="1.[106,137,1122,1139]" captionTargetBox="[348,1251,134,1092]" captionTargetId="graphics-459@1.[389,1192,163,1056]" captionText="Fig.1 | Overviewof Eriptychiusamericanus PF 1795.a,b, Photographsof part counterpart) (d). Colourscheme forrenders:blue-greys,cranialcartilages PF1795a,which hadthe splitface set inepoxyandwas manually prepared (a), and (matchingthe detailed scheme inFig.2); transparencies,thedermal skeleton; itscounterpart PF 1795b,which remainsinrock matrix(b). Both are shown inan orange,branchial plates;red,orbital plates.Anterior totop ina–d.ant.tess., anatomically ventral view.c,d, Digital model of computed tomographic dataof anterior tesserae;artic.vent.tess.,articulated ventral tesserae;branch.plate, thecombined part and counterpart with most of dermal skeleton rendered branchial plate;cran.cart.,cranialcartilages;disp.,displaced;frag.,fragment; transparent:anatomical ventralview (corresponding tothe visible areaof L., left;orb.cart.,orbital cartilage;orb.plates,orbital plates;R.,right;vasc., thepart in epoxy) (c) andanatomical dorsal view (buried in matrixin the vasculature;?,probable.Scale bar applies toall panels." figureDoi="http://doi.org/10.5281/zenodo.8368405" httpUri="https://zenodo.org/record/8368405/files/figure.png" pageId="3" pageNumber="4">Fig. 2c,h,i</figureCitation>
|
||
), unlike most jawless stem-gnathostomes (notable exceptions being
|
||
<taxonomicName id="4C69FE9AFFD6FF90FFC4F91C9A9E47BF" box="[106,203,1739,1758]" family="Doryaspidae" genus="Doryaspis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FFC4F91C9A9E47BF" bold="true" box="[106,203,1739,1758]" italics="true" pageId="3" pageNumber="4">Doryaspis</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation id="EFF8F8E8FFD6FF90FF65F9119A8E47B2" author="Pernegre, V. N." box="[203,219,1734,1747]" journalOrPublisher="Svalbard. J. Vertebr. Paleontol." pageId="3" pageNumber="4" pagination="735 - 746" part="22" refId="ref5735" refString="40. Pernegre, V. N. The genus Doryaspis White (Heterostraci) from the Lower Devonian of Vestspitsbergen, Svalbard. J. Vertebr. Paleontol. 22, 735 - 746 (2003)." title="The genus Doryaspis White (Heterostraci) from the Lower Devonian of Vestspitsbergen" type="journal article" year="2003">
|
||
<emphasis id="B91D590BFFD6FF90FF65F9119A8E47B2" bold="true" box="[203,219,1734,1747]" pageId="3" pageNumber="4">
|
||
<superScript id="7C1C2851FFD6FF90FF65F9119A8E47B2" attach="both" box="[203,219,1734,1747]" fontSize="5" pageId="3" pageNumber="4">40</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
<emphasis id="B91D590BFFD6FF90FF73F91D9B2B47BF" bold="true" box="[221,382,1738,1758]" pageId="3" pageNumber="4">
|
||
and
|
||
<taxonomicName id="4C69FE9AFFD6FF90FEABF91C9B2B47BF" authorityName="Schlüter" authorityYear="1887" box="[261,382,1739,1758]" family="Psammosteidae" genus="Drepanaspis" higherTaxonomySource="GBIF" kingdom="Animalia" pageId="3" pageNumber="4" phylum="Chordata" rank="genus">
|
||
<emphasis id="B91D590BFFD6FF90FEABF91C9B2B47BF" bold="true" box="[261,382,1739,1758]" italics="true" pageId="3" pageNumber="4">Drepanaspis</emphasis>
|
||
</taxonomicName>
|
||
</emphasis>
|
||
<bibRefCitation id="EFF8F8E8FFD6FF90FED0F9119BDE47B2" author="Gross, W." box="[382,395,1734,1747]" journalOrPublisher="Palaeontographica A" pageId="3" pageNumber="4" pagination="133 - 155" part="121" refId="ref5774" refString="41. Gross, W. Drepanaspis gemuendenensis Schluter. Neuuntersuchung. Palaeontographica A 121, 133 - 155 (1963)." title="Drepanaspis gemuendenensis Schluter. Neuuntersuchung" type="journal article" year="1963">
|
||
<emphasis id="B91D590BFFD6FF90FED0F9119BDE47B2" bold="true" box="[382,395,1734,1747]" pageId="3" pageNumber="4">
|
||
<superScript id="7C1C2851FFD6FF90FED0F9119BDE47B2" attach="left" box="[382,395,1734,1747]" fontSize="5" pageId="3" pageNumber="4">41</superScript>
|
||
</emphasis>
|
||
</bibRefCitation>
|
||
).The fact that the cartilages are separate may mean that some movement of the oral endoskeleton was possible, although this would have presumably been limited by their close relationship with the squamation.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |