772 lines
121 KiB
XML
772 lines
121 KiB
XML
<document ID-DOI="http://doi.org/10.5281/zenodo.6315568" ID-GBIF-Dataset="93361143-cbf9-4a9e-a1c3-0271ebeea406" ID-GBIF-Taxon="193586406" ID-Zenodo-Dep="6315568" approvalRequired="4" approvalRequired_for_document="1" approvalRequired_for_textStreams="2" approvalRequired_for_treatments="1" checkinTime="1645629896261" checkinUser="felipe" docAuthor="Jagielska, Natalia, O’Sullivan, Michael, Funston, Gregory F., Butler, Ian B., Challands, Thomas J., Clark, Neil D. L., Fraser, Nicholas C., Penny, Amelia, Ross, Dugald A., Wilkinson, Mark & Brusatte, Stephen L." docDate="2022" docId="03E587D1F10D5A30104A8B0B9DA6E108" docLanguage="en" docName="CurrBiol.32.1-8.pdf.imf" docOrigin="Current Biology 32" docSource="http://dx.doi.org/10.1016/j.cub.2022.01.073" docTitle="Dearc sgiathanach Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte 2022, sp. nov." docType="treatment" docVersion="6" lastPageId="6" lastPageNumber="6" masterDocId="FFDCFFA9F10C5A3613088D1C9E21E42B" masterDocTitle="A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs" masterLastPageNumber="8" masterPageNumber="1" pageId="1" pageNumber="1" updateTime="1646094374445" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-4.0">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>A skeleton from the Middle Jurassic of Scotland illuminates an earlier origin of large pterosaurs</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Jagielska, Natalia</mods:namePart>
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<mods:affiliation>School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, Scotland EH 9 3 FE, UK</mods:affiliation>
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<mods:nameIdentifier type="email">n.jagielska@sms.ed.ac.uk</mods:nameIdentifier>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>O’Sullivan, Michael</mods:namePart>
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<mods:affiliation>Gortroe Rathkeale, Co Limerick, Ireland</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Funston, Gregory F.</mods:namePart>
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<mods:affiliation>School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, Scotland EH 9 3 FE, UK</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Butler, Ian B.</mods:namePart>
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<mods:affiliation>School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, Scotland EH 9 3 FE, UK</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Challands, Thomas J.</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Clark, Neil D. L.</mods:namePart>
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<mods:affiliation>The Hunterian, University of Glasgow, University Avenue, Glasgow, Scotland G 12 8 QQ, UK</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Fraser, Nicholas C.</mods:namePart>
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<mods:affiliation>School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, Scotland EH 9 3 FE, UK & National Museums Scotland, Chambers Street, Edinburgh, Scotland EH 1 1 JF, UK</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Penny, Amelia</mods:namePart>
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<mods:affiliation>School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, Scotland EH 9 3 FE, UK & School of Biology, University of St Andrews, Biomedical Sciences Research Complex, St. Andrews, Scotland KY 16 9 ST, UK</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Ross, Dugald A.</mods:namePart>
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<mods:affiliation>Staffin Museum, 6 Ellishadder, Staffin, Isle of Skye, Scotland IV 51 9 JE, UK</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Wilkinson, Mark</mods:namePart>
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<mods:affiliation>School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, Scotland EH 9 3 FE, UK</mods:affiliation>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Brusatte, Stephen L.</mods:namePart>
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<mods:affiliation>School of GeoSciences, University of Edinburgh, Grant Institute, James Hutton Road, Edinburgh, Scotland EH 9 3 FE, UK & National Museums Scotland, Chambers Street, Edinburgh, Scotland EH 1 1 JF, UK</mods:affiliation>
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<mods:nameIdentifier type="email">stephen.brusatte@ed.ac.uk</mods:nameIdentifier>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:titleInfo>
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<mods:title>Current Biology</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2022</mods:date>
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<mods:detail type="pubDate">
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<mods:number>2022-02-28</mods:number>
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</mods:detail>
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<mods:detail type="volume">
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<mods:number>32</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>1</mods:start>
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<mods:end>8</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:location>
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<mods:url>http://dx.doi.org/10.1016/j.cub.2022.01.073</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1016/j.cub.2022.01.073</mods:identifier>
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<mods:identifier type="GBIF-Dataset">93361143-cbf9-4a9e-a1c3-0271ebeea406</mods:identifier>
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<mods:identifier type="Zenodo-Dep">6251702</mods:identifier>
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</mods:mods>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.6315568" ID-GBIF-Taxon="193586406" ID-Zenodo-Dep="6315568" LSID="urn:lsid:plazi:treatment:03E587D1F10D5A30104A8B0B9DA6E108" httpUri="http://treatment.plazi.org/id/03E587D1F10D5A30104A8B0B9DA6E108" lastPageId="6" lastPageNumber="6" pageId="1" pageNumber="1">
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<subSubSection box="[834,1108,1559,1580]" pageId="1" pageNumber="1" type="nomenclature">
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<paragraph blockId="1.[811,1450,1413,1580]" box="[834,1108,1559,1580]" pageId="1" pageNumber="1">
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<taxonomicName authority="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte, 2022" authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[834,1021,1559,1580]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="1" pageNumber="1" phylum="Chordata" rank="species" species="sgiathanach" status="sp. nov.">
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<emphasis box="[834,1021,1559,1580]" italics="true" pageId="1" pageNumber="1">Dearc sgiathanach</emphasis>
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</taxonomicName>
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<taxonomicNameLabel box="[1027,1108,1559,1580]" pageId="1" pageNumber="1" rank="species">sp. nov.</taxonomicNameLabel>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="1" pageNumber="1" type="etymology">
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<paragraph blockId="1.[811,1450,1634,1769]" box="[811,929,1634,1652]" pageId="1" pageNumber="1">
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<heading bold="true" box="[811,929,1634,1652]" fontSize="8" level="6" pageId="1" pageNumber="1" reason="6">
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<emphasis bold="true" box="[811,929,1634,1652]" pageId="1" pageNumber="1">Etymology</emphasis>
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</heading>
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</paragraph>
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<paragraph blockId="1.[811,1450,1634,1769]" pageId="1" pageNumber="1">Scottish Gaelic, with the double meaning of ‘‘winged reptile’’and ‘‘reptile from Skye,’’ paying homage to pterosaurs (winged reptiles) and the Gaelic name for Skye (An t-Eilean Sgitheanach, the ‘‘winged isle’’). Phonetic pronunciation: jark ski-an-ach.</paragraph>
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</subSubSection>
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<subSubSection lastPageId="2" lastPageNumber="2" pageId="1" pageNumber="1" type="materials_examined">
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<materialsCitation ID-GBIF-Occurrence="3496997301" collectedFrom="embedded in a well-sorted lagoonal bioclastic limestone" collectionCode="NMS" country="United Kingdom" lastPageId="2" lastPageNumber="2" location="Rubha nam Brathairean" municipality="Trotternish" pageId="1" pageNumber="1" specimenCode="G.2021.6.1-4, NMS G.2021.6.1, NMS G.2021.6.3, NMS G.2021.6.4" specimenCount="1" stateProvince="Isle of Skye" typeStatus="holotype">
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<paragraph blockId="1.[811,1450,1823,1987]" lastBlockId="2.[159,798,1470,1666]" lastPageId="2" lastPageNumber="2" pageId="1" pageNumber="1">
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<heading bold="true" box="[811,911,1823,1841]" fontSize="8" level="6" pageId="1" pageNumber="1" reason="6">
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<emphasis bold="true" box="[811,911,1823,1841]" pageId="1" pageNumber="1">
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<typeStatus box="[811,911,1823,1841]" pageId="1" pageNumber="1">Holotype</typeStatus>
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</emphasis>
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</heading>
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<collectionCode box="[811,862,1849,1870]" collectionName="Singapore, National University of Singapore, Raffles Museum of Biodiversity Research, Zoological Reference Collection" pageId="1" pageNumber="1">NMS</collectionCode>
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(National Museums Scotland, Edinburgh)
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<specimenCode box="[1316,1449,1849,1870]" pageId="1" pageNumber="1">G.2021.6.1-4</specimenCode>
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(
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<figureCitation box="[817,932,1879,1900]" captionStart="Figure 1" captionStartId="2.[1056,1116,271,290]" captionTargetBox="[160,1018,273,1413]" captionTargetId="figure-152@2.[159,1020,271,1414]" captionTargetPageId="2" captionText="Figure 1. The new Middle Jurassic pterosaur Dearc sgiathanach (A–C) Photographs of main slab (NMS G.2021.6.1– 2), bones in dorsal view (A); wing phalanges 2–3 (NMS G.2021.6.3–4), in dorsal view (B); and main counterslab (NMS G.2021.6.3), bones in ventral view (C). (D–F) Schematic drawings of (A)–(C). (G–H) Reconstruction of skull in dorsal (G) and ventral (I) views and skeleton in lateral view (H). ar, articular region; cd, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, digit; den, dentary;dors,dorsal vertebrae;dpc,deltopectoral crest; ep, extensor process; fm, femur; gas, gastralia; hdc, humeral distal condyle; hmt, humeral tubercle; isc, ischium; ju, jugal; max, maxilla; mc, metacarpal; mt,metatarsal;po,postorbital;r, ribs; sac, sacral plate; sca, scapula; scv, sacral (?) vertebrae; sm, sesamoid; sq, squamosal; st, sternum; symp, symphysis; uc, ulnar crest; ul, ulna; wp, wing phalanx; r/l, right/left. Blue on reconstructions are missing regions; red line in (E) is location of histological sectioning. Scale bars, 30 mm. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251706" httpUri="https://zenodo.org/record/6251706/files/figure.png" pageId="1" pageNumber="1">Figures 1, 2</figureCitation>
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,
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<figureCitation box="[944,956,1879,1900]" captionStart="Figure 3" captionStartId="4.[159,219,1471,1490]" captionTargetBox="[161,1465,271,1431]" captionTargetId="figure-185@4.[161,1465,271,1431]" captionTargetPageId="4" captionText="Figure 3. Postcranial skeleton and dentition of the new Middle Jurassic pterosaur Dearc sgiathanach Photographs of the right manus (A), cervical series (B), pubic region (C), right humerus (D), left humerus (E), left metacarpal-phalanx articular region (F), right maxilla (G), and left pes (H) of NMS G.2021.6.1–4. ac, anterior caudal vertebrae; ar, articular region; c, condyle; cdv, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, dentary; dc, distal condyle; dpc, deltopectoral crest;dvrt, dorsal vertebrae; en,enamel;exp,extensor process; fem,femur; ft, flexor tubercle; gas,gastralia; h, humerus;ic, intercondylar groove; isc,ischium; max,maxilla; mcp,metacarpal; md, maxillary dentition; mg, medial groove; mt, metatarsal; nss,neural spine scar; pa, preacetabular process; poz, postzygapophysis ppb,prepubis; prez,prezygapophysis; rad,radius;rb,rib; sac,sacrum;sm,sesamoid;uc,ulnar crest; ul,ulna;un, ungual;vert, vertebra;wp1, wing phalanx one. Scale, 10 mm per bar. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251720" httpUri="https://zenodo.org/record/6251720/files/figure.png" pageId="1" pageNumber="1">3</figureCitation>
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, S
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<figureCitation box="[983,995,1879,1900]" captionStart="Figure 1" captionStartId="2.[1056,1116,271,290]" captionTargetBox="[160,1018,273,1413]" captionTargetId="figure-152@2.[159,1020,271,1414]" captionTargetPageId="2" captionText="Figure 1. The new Middle Jurassic pterosaur Dearc sgiathanach (A–C) Photographs of main slab (NMS G.2021.6.1– 2), bones in dorsal view (A); wing phalanges 2–3 (NMS G.2021.6.3–4), in dorsal view (B); and main counterslab (NMS G.2021.6.3), bones in ventral view (C). (D–F) Schematic drawings of (A)–(C). (G–H) Reconstruction of skull in dorsal (G) and ventral (I) views and skeleton in lateral view (H). ar, articular region; cd, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, digit; den, dentary;dors,dorsal vertebrae;dpc,deltopectoral crest; ep, extensor process; fm, femur; gas, gastralia; hdc, humeral distal condyle; hmt, humeral tubercle; isc, ischium; ju, jugal; max, maxilla; mc, metacarpal; mt,metatarsal;po,postorbital;r, ribs; sac, sacral plate; sca, scapula; scv, sacral (?) vertebrae; sm, sesamoid; sq, squamosal; st, sternum; symp, symphysis; uc, ulnar crest; ul, ulna; wp, wing phalanx; r/l, right/left. Blue on reconstructions are missing regions; red line in (E) is location of histological sectioning. Scale bars, 30 mm. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251706" httpUri="https://zenodo.org/record/6251706/files/figure.png" pageId="1" pageNumber="1">2</figureCitation>
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, and S3), a three-dimensionally preserved articulated skeleton, lacking anterior and dorsal portions of the cranium, left manus, portions of the wings, hindlimb elements, and the distal tail. The fossil was separated into four slabs during preparation: the main slab contains the majority of bones, exposed in dorsal view (
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<specimenCode box="[399,570,1499,1520]" pageId="2" pageNumber="2">NMS G.2021.6.1</specimenCode>
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), and the main counter slab contains bones exposed ventrally (
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<specimenCode box="[576,751,1528,1549]" pageId="2" pageNumber="2">NMS G.2021.6.3</specimenCode>
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). An additional block contains a wing phalanx (
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<specimenCode box="[611,789,1557,1578]" pageId="2" pageNumber="2">NMS G.2021.6.4</specimenCode>
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). The skull and anterior cervical vertebrae (NMS G.
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<date box="[699,792,1586,1607]" pageId="2" pageNumber="2" value="2021-06-02">2021.6.2</date>
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) were separated from the main slab for X-ray computed microtomography (
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<emphasis bold="true" box="[274,287,1649,1666]" pageId="2" pageNumber="2">M</emphasis>
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CT) (
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<figureCitation box="[334,415,1645,1666]" captionStart="Figure 1" captionStartId="2.[1056,1116,271,290]" captionTargetBox="[160,1018,273,1413]" captionTargetId="figure-152@2.[159,1020,271,1414]" captionTargetPageId="2" captionText="Figure 1. The new Middle Jurassic pterosaur Dearc sgiathanach (A–C) Photographs of main slab (NMS G.2021.6.1– 2), bones in dorsal view (A); wing phalanges 2–3 (NMS G.2021.6.3–4), in dorsal view (B); and main counterslab (NMS G.2021.6.3), bones in ventral view (C). (D–F) Schematic drawings of (A)–(C). (G–H) Reconstruction of skull in dorsal (G) and ventral (I) views and skeleton in lateral view (H). ar, articular region; cd, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, digit; den, dentary;dors,dorsal vertebrae;dpc,deltopectoral crest; ep, extensor process; fm, femur; gas, gastralia; hdc, humeral distal condyle; hmt, humeral tubercle; isc, ischium; ju, jugal; max, maxilla; mc, metacarpal; mt,metatarsal;po,postorbital;r, ribs; sac, sacral plate; sca, scapula; scv, sacral (?) vertebrae; sm, sesamoid; sq, squamosal; st, sternum; symp, symphysis; uc, ulnar crest; ul, ulna; wp, wing phalanx; r/l, right/left. Blue on reconstructions are missing regions; red line in (E) is location of histological sectioning. Scale bars, 30 mm. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251706" httpUri="https://zenodo.org/record/6251706/files/figure.png" pageId="2" pageNumber="2">Figure 2</figureCitation>
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). Measurements in Data S1A.
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</paragraph>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.6251706" ID-Zenodo-Dep="6251706" httpUri="https://zenodo.org/record/6251706/files/figure.png" pageId="2" pageNumber="2" startId="2.[1056,1116,271,290]" subCaptionStartIDs="2.[1093,1154,878,895]" subCaptionStarts="Figures S" targetBox="[160,1018,273,1413]" targetPageId="2">
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<paragraph blockId="2.[1056,1468,271,895]" pageId="2" pageNumber="2">
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Figure 1. The new Middle Jurassic pterosaur
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<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[1103,1282,297,315]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="2" pageNumber="2" phylum="Chordata" rank="species" species="sgiathanach">Dearc sgiathanach</taxonomicName>
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</paragraph>
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<paragraph blockId="2.[1056,1468,271,895]" pageId="2" pageNumber="2">(A–C) Photographs of main slab (NMS G.2021.6.1– 2), bones in dorsal view (A); wing phalanges 2–3 (NMS G.2021.6.3–4), in dorsal view (B); and main counterslab (NMS G.2021.6.3), bones in ventral view (C).</paragraph>
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<paragraph blockId="2.[1056,1468,271,895]" box="[1056,1361,449,466]" pageId="2" pageNumber="2">(D–F) Schematic drawings of (A)–(C).</paragraph>
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<paragraph blockId="2.[1056,1468,271,895]" pageId="2" pageNumber="2">(G–H) Reconstruction of skull in dorsal (G) and ventral (I) views and skeleton in lateral view (H).</paragraph>
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<paragraph blockId="2.[1056,1468,271,895]" pageId="2" pageNumber="2">ar, articular region; cd, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, digit; den, dentary;dors,dorsal vertebrae;dpc,deltopectoral crest; ep, extensor process; fm, femur; gas, gastralia; hdc, humeral distal condyle; hmt, humeral tubercle; isc, ischium; ju, jugal; max, maxilla; mc, metacarpal; mt,metatarsal;po,postorbital;r, ribs; sac, sacral plate; sca, scapula; scv, sacral (?) vertebrae; sm, sesamoid; sq, squamosal; st, sternum; symp, symphysis; uc, ulnar crest; ul, ulna; wp, wing phalanx; r/l, right/left. Blue on reconstructions are missing regions; red line in (E) is location of histological sectioning. Scale bars, 30 mm.</paragraph>
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<paragraph blockId="2.[1056,1468,271,895]" box="[1056,1251,878,895]" pageId="2" pageNumber="2">
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See
|
||
<figureCitation box="[1093,1182,878,895]" captionStart="Figure 2" captionStartId="3.[142,202,940,959]" captionTargetBox="[143,1448,272,899]" captionTargetId="figure-424@3.[142,1449,271,900]" captionTargetPageId="3" captionText="Figure 2. Skull of the new Middle Jurassic pterosaur Dearc sgiathanach and comparisons of pterosaur brain and ear endocasts (A–L) Photographs (A–D), segmented MCT scan renderings (E–H), and schematic drawings (I–L) of the skull (NMS G.2021.6.2) in, from top to bottom, left lateral (reversed), dorsal, right lateral, and ventral views. (M–U) Brain and inner ear endocasts of Rhamphorhynchus (based on Witmer et al.7) (M, P, and Q), Dearc (N, R, and S), and Tapejara (based on Eck et al.8) (O, T, and U), shown in the skull in left lateral view and isolated in left lateral and dorsal views, respectively. asc, anterior semi-circular canal; bp, basisphenoid; cbl, cerebellum; cer, cerebrum; ch, choana; chl, channels; cv, third cervical; d, dentary; den, dentition; e, endocast; ec, ectopterygoid; et, edentulous anterior tip; f, frontal (?); hy, hyoid; ipv, interpterygoid vacuity; itf, inferior temporal fenestra; j, jugal; la, lacrimal; md, matrix infilled dent; mx, maxilla; n, nasal; olf, olfactory bulb; opl, optic lobe; p, parietal; pm, premaxilla; po, postorbital; ppd, post-palatal depression; psc, posterior semi-circular canal; ptf, pneumatic foramen; q, quadrate; qj, quadrojugal; rapr, retroarticular process; rh, probable placement of rhamphotheca; sof, suborbital fenestra; sq,squamosal;sbtf,subtemporal fenestra;stf, superior temporal fenestra.In (E)–(L), bones that cannot be distinguished in the MCT scan due to fusion or insufficient resolution are conservatively rendered together in one color. Scale bars, 30 mm. See Figure S2." figureDoi="http://doi.org/10.5281/zenodo.6251712" httpUri="https://zenodo.org/record/6251712/files/figure.png" pageId="2" pageNumber="2">Figures S2</figureCitation>
|
||
and S
|
||
<figureCitation box="[1236,1246,878,895]" captionStart="Figure 3" captionStartId="4.[159,219,1471,1490]" captionTargetBox="[161,1465,271,1431]" captionTargetId="figure-185@4.[161,1465,271,1431]" captionTargetPageId="4" captionText="Figure 3. Postcranial skeleton and dentition of the new Middle Jurassic pterosaur Dearc sgiathanach Photographs of the right manus (A), cervical series (B), pubic region (C), right humerus (D), left humerus (E), left metacarpal-phalanx articular region (F), right maxilla (G), and left pes (H) of NMS G.2021.6.1–4. ac, anterior caudal vertebrae; ar, articular region; c, condyle; cdv, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, dentary; dc, distal condyle; dpc, deltopectoral crest;dvrt, dorsal vertebrae; en,enamel;exp,extensor process; fem,femur; ft, flexor tubercle; gas,gastralia; h, humerus;ic, intercondylar groove; isc,ischium; max,maxilla; mcp,metacarpal; md, maxillary dentition; mg, medial groove; mt, metatarsal; nss,neural spine scar; pa, preacetabular process; poz, postzygapophysis ppb,prepubis; prez,prezygapophysis; rad,radius;rb,rib; sac,sacrum;sm,sesamoid;uc,ulnar crest; ul,ulna;un, ungual;vert, vertebra;wp1, wing phalanx one. Scale, 10 mm per bar. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251720" httpUri="https://zenodo.org/record/6251720/files/figure.png" pageId="2" pageNumber="2">3</figureCitation>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="2.[159,798,1706,1987]" box="[159,386,1706,1724]" pageId="2" pageNumber="2">
|
||
<heading bold="true" box="[159,386,1706,1724]" fontSize="8" level="6" pageId="2" pageNumber="2" reason="6">
|
||
<emphasis bold="true" box="[159,386,1706,1724]" pageId="2" pageNumber="2">Locality and horizon</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="2.[159,798,1706,1987]" pageId="2" pageNumber="2">
|
||
The specimen was discovered by A.P. in 2017 at
|
||
<location LSID="urn:lsid:plazi:treatment:03E587D1F10D5A30104A8B0B9DA6E108:8E93601CF10E5A3411AE8BD99F32E2DC" country="United Kingdom" municipality="Trotternish" name="Rubha nam Brathairean" pageId="2" pageNumber="2" stateProvince="Isle of Skye">Rubha nam Brathairean</location>
|
||
(
|
||
<location LSID="urn:lsid:plazi:treatment:03E587D1F10D5A30104A8B0B9DA6E108:8E93601CF10E5A3412178BFE9F98E2DC" box="[287,441,1762,1783]" country="United Kingdom" municipality="Trotternish" name="Brothers' Point" pageId="2" pageNumber="2" stateProvince="Isle of Skye">Brothers’ Point</location>
|
||
),
|
||
<collectingMunicipality box="[455,567,1762,1783]" pageId="2" pageNumber="2">Trotternish</collectingMunicipality>
|
||
,
|
||
<collectingRegion box="[575,693,1762,1783]" pageId="2" pageNumber="2">Isle of Skye</collectingRegion>
|
||
,
|
||
<collectingCountry box="[701,794,1762,1783]" name="United Kingdom" pageId="2" pageNumber="2">Scotland</collectingCountry>
|
||
, in the Lonfearn Member of the Lealt Shale Formation (Bathonian, Middle Jurassic)
|
||
<superScript attach="left" box="[324,357,1816,1830]" fontSize="6" pageId="2" pageNumber="2">
|
||
<bibRefCitation author="Hudson, J. D. & Wakefield, M. I." box="[324,333,1816,1830]" journalOrPublisher="Scott. J. Geol." pageId="2" pageNumber="2" pagination="87 - 97" part="54" refId="ref5817" refString="9. Hudson, J. D., and Wakefield, M. I. (2018). The Lonfearn Member, Lealt Shale Formation (Middle Jurassic) of the Inner Hebrides, Scotland. Scott. J. Geol. 54, 87 - 97." title="The Lonfearn Member, Lealt Shale Formation (Middle Jurassic) of the Inner Hebrides, Scotland" type="journal article" year="2018">9</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Harris, J. P. & Hudson, J. D." box="[338,357,1816,1830]" journalOrPublisher="Scott. J. Geol." pageId="2" pageNumber="2" pagination="231 - 250" part="16" refId="ref5867" refString="10. Harris, J. P., and Hudson, J. D. (1980). Lithostratigraphy of the Great Estuarine Group (Middle Jurassic), Inner Hebrides. Scott. J. Geol. 16, 231 - 250." title="Lithostratigraphy of the Great Estuarine Group (Middle Jurassic), Inner Hebrides" type="journal article" year="1980">10</bibRefCitation>
|
||
</superScript>
|
||
(
|
||
<figureCitation box="[368,464,1820,1841]" captionStart="Figure 1" captionStartId="2.[1056,1116,271,290]" captionTargetBox="[160,1018,273,1413]" captionTargetId="figure-152@2.[159,1020,271,1414]" captionTargetPageId="2" captionText="Figure 1. The new Middle Jurassic pterosaur Dearc sgiathanach (A–C) Photographs of main slab (NMS G.2021.6.1– 2), bones in dorsal view (A); wing phalanges 2–3 (NMS G.2021.6.3–4), in dorsal view (B); and main counterslab (NMS G.2021.6.3), bones in ventral view (C). (D–F) Schematic drawings of (A)–(C). (G–H) Reconstruction of skull in dorsal (G) and ventral (I) views and skeleton in lateral view (H). ar, articular region; cd, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, digit; den, dentary;dors,dorsal vertebrae;dpc,deltopectoral crest; ep, extensor process; fm, femur; gas, gastralia; hdc, humeral distal condyle; hmt, humeral tubercle; isc, ischium; ju, jugal; max, maxilla; mc, metacarpal; mt,metatarsal;po,postorbital;r, ribs; sac, sacral plate; sca, scapula; scv, sacral (?) vertebrae; sm, sesamoid; sq, squamosal; st, sternum; symp, symphysis; uc, ulnar crest; ul, ulna; wp, wing phalanx; r/l, right/left. Blue on reconstructions are missing regions; red line in (E) is location of histological sectioning. Scale bars, 30 mm. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251706" httpUri="https://zenodo.org/record/6251706/files/figure.png" pageId="2" pageNumber="2">Figure S1</figureCitation>
|
||
). The skeleton was
|
||
<collectedFrom pageId="2" pageNumber="2">embedded in a well-sorted lagoonal bioclastic limestone</collectedFrom>
|
||
(rich in
|
||
<emphasis box="[673,790,1849,1870]" italics="true" pageId="2" pageNumber="2">Neomiodon</emphasis>
|
||
, ostracods, and conchostracans), which overlies and infills dinosaur trackways impressed in subaerially exposed mudstones.
|
||
<bibRefCitation author="dePolo, P. E. & Brusatte, S. L. & Challands, T. J. & Foffa, D. & Wilkinson, M. & Clark, N. D. L. & Hoad, J. & Pereira, P. V. L. & Ross, D. A. & Wade, T. J." box="[778,797,1903,1917]" journalOrPublisher="PLoS ONE" pageId="2" pageNumber="2" pagination="0229640" part="15" refId="ref5913" refString="11. dePolo, P. E., Brusatte, S. L., Challands, T. J., Foffa, D., Wilkinson, M., Clark, N. D. L., Hoad, J., Pereira, P. V. L., Ross, D. A., and Wade, T. J. (2020). Novel track morphotypes from new tracksites indicate increased Middle Jurassic dinosaur diversity on the Isle of Skye, Scotland. PLoS ONE 15, e 0229640." title="Novel track morphotypes from new tracksites indicate increased Middle Jurassic dinosaur diversity on the Isle of Skye, Scotland" type="journal article" year="2020">
|
||
<superScript attach="none" box="[778,797,1903,1917]" fontSize="6" pageId="2" pageNumber="2">11</superScript>
|
||
</bibRefCitation>
|
||
These units formed in a marginal marine/nearshore environment that fluctuated between submerged and exposed.
|
||
</paragraph>
|
||
</materialsCitation>
|
||
</subSubSection>
|
||
<subSubSection pageId="2" pageNumber="2" type="diagnosis">
|
||
<paragraph blockId="2.[1056,1468,1005,1315]" box="[1056,1166,1005,1023]" pageId="2" pageNumber="2">
|
||
<heading bold="true" box="[1056,1166,1005,1023]" fontSize="8" level="6" pageId="2" pageNumber="2" reason="6">
|
||
<emphasis bold="true" box="[1056,1166,1005,1023]" pageId="2" pageNumber="2">Diagnosis</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="2.[1056,1468,1005,1315]" pageId="2" pageNumber="2">
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[1056,1251,1031,1052]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="2" pageNumber="2" phylum="Chordata" rank="species" species="sgiathanach">
|
||
<emphasis box="[1056,1251,1031,1052]" italics="true" pageId="2" pageNumber="2">Dearc sgiathanach</emphasis>
|
||
</taxonomicName>
|
||
is a rhamphorhynchine pterosaur with the following autapomorphies: tri-tubular vomers with ‘‘trident-shaped’’ precapillary contact, pre-choana depression on the palatal surface of the maxilla, enlarged optic lobes expanded anteroposteriorly, and fourth metatarsal more robust (diameter 2.5
|
||
<emphasis bold="true" box="[1087,1106,1267,1286]" pageId="2" pageNumber="2">×</emphasis>
|
||
) than mt1-3. For additional information, see STAR Methods.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="5" lastPageNumber="5" pageId="2" pageNumber="2" type="description">
|
||
<paragraph blockId="2.[829,1467,1355,1987]" box="[1056,1371,1355,1373]" pageId="2" pageNumber="2">
|
||
<heading bold="true" box="[1056,1371,1355,1373]" fontSize="8" level="6" pageId="2" pageNumber="2" reason="6">
|
||
<emphasis bold="true" box="[1056,1371,1355,1373]" pageId="2" pageNumber="2">Bone histology and maturity</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="2.[829,1467,1355,1987]" pageId="2" pageNumber="2">
|
||
Using Bennett’s
|
||
<bibRefCitation author="Bennett, S. C." box="[1223,1242,1377,1391]" journalOrPublisher="J. Paleontol." pageId="2" pageNumber="2" pagination="569 - 580" part="69" refId="ref6014" refString="12. Bennett, S. C. (1995). A statistical study of Rhamphorhynchus from the Solnhofen Limestone of Germany: year-classes of a single large species. J. Paleontol. 69, 569 - 580." title="A statistical study of Rhamphorhynchus from the Solnhofen Limestone of Germany: year-classes of a single large species" type="journal article" year="1995">
|
||
<superScript attach="left" box="[1223,1242,1377,1391]" fontSize="6" pageId="2" pageNumber="2">12</superScript>
|
||
</bibRefCitation>
|
||
osteologically based ontogenetic stages for the closely related
|
||
<taxonomicName authority=", NMS G." authorityName="NMS G." box="[1056,1353,1440,1461]" class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="2" pageNumber="2" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1056,1245,1440,1461]" italics="true" pageId="2" pageNumber="2">Rhamphorhynchus</emphasis>
|
||
, NMS G.
|
||
</taxonomicName>
|
||
2021.6.1–4 has features of terminal-stage adults, such as large and recurved premaxillary teeth, fused scapula-coracoid, well-developed humeral crests, smooth bone texture, and fused long bone epiphyses. However, some osteological features are indicative of immaturity according to Bennett:
|
||
<bibRefCitation author="Bennett, S. C." box="[1162,1181,1582,1596]" journalOrPublisher="J. Paleontol." pageId="2" pageNumber="2" pagination="569 - 580" part="69" refId="ref6014" refString="12. Bennett, S. C. (1995). A statistical study of Rhamphorhynchus from the Solnhofen Limestone of Germany: year-classes of a single large species. J. Paleontol. 69, 569 - 580." title="A statistical study of Rhamphorhynchus from the Solnhofen Limestone of Germany: year-classes of a single large species" type="journal article" year="1995">
|
||
<superScript attach="left" box="[1162,1181,1582,1596]" fontSize="6" pageId="2" pageNumber="2">12</superScript>
|
||
</bibRefCitation>
|
||
portions of the skull are unfused, such as the jugal with the lacrimal, and there appears to be limited fusion in the sacral vertebrae.
|
||
</paragraph>
|
||
<paragraph blockId="2.[829,1467,1355,1987]" lastBlockId="3.[142,779,1294,1491]" lastPageId="3" lastPageNumber="3" pageId="2" pageNumber="2">
|
||
Immaturity is corroborated by bone histology (STAR Methods; Data S2). The cortex of a sampled wing phalanx is composed entirely of primary fibrolamellar bone
|
||
<bibRefCitation author="Padian, K. & Lamm, E. L." box="[1192,1211,1728,1742]" journalOrPublisher="University of California Press" pageId="2" pageNumber="2" refId="ref6055" refString="13. Padian, K., and Lamm, E. L. (2013). Bone Histology of Fossil Tetrapods (University of California Press)." title="Bone Histology of Fossil Tetrapods" type="book" year="2013">
|
||
<superScript attach="left" box="[1192,1211,1728,1742]" fontSize="6" pageId="2" pageNumber="2">13</superScript>
|
||
</bibRefCitation>
|
||
and preserves two prominent lines of arrested growth (LAGs), which indicate that the individual was at least 2 years old at death.
|
||
<bibRefCitation author="Castanet, J. & Croci, S. & Aujard, F. & Perret, M. & Cubo, J. & de Margerie, E." box="[1251,1270,1786,1800]" journalOrPublisher="J. Zool." pageId="2" pageNumber="2" pagination="31 - 39" part="263" refId="ref6085" refString="14. Castanet, J., Croci, S., Aujard, F., Perret, M., Cubo, J., and de Margerie, E. (2004). Lines of arrested growth in bone and age estimation in a small primate: Microcebus murinus. J. Zool. 263, 31 - 39." title="Lines of arrested growth in bone and age estimation in a small primate: Microcebus murinus" type="journal article" year="2004">
|
||
<superScript attach="left" box="[1251,1270,1786,1800]" fontSize="6" pageId="2" pageNumber="2">14</superScript>
|
||
</bibRefCitation>
|
||
The position of the second LAG close to the external bone surface suggests that the individual died shortly after emerging from an annual growth hiatus. The cortex is densely vascularized and has a high proportion of woven bone, indicating a rapid rate of growth throughout life.
|
||
<bibRefCitation author="de Margerie, E. & Cubo, J. & Castanet, J." box="[864,883,1932,1946]" journalOrPublisher="C. R. Biol." pageId="2" pageNumber="2" pagination="221 - 230" part="325" refId="ref6149" refString="15. de Margerie, E., Cubo, J., and Castanet, J. (2002). Bone typology and growth rate: testing and quantifying ' Amprino's rule' in the mallard (Anas platyrhynchos). C. R. Biol. 325, 221 - 230." title="Bone typology and growth rate: testing and quantifying ' Amprino's rule' in the mallard (Anas platyrhynchos)" type="journal article" year="2002">
|
||
<superScript attach="left" box="[864,883,1932,1946]" fontSize="6" pageId="2" pageNumber="2">15</superScript>
|
||
</bibRefCitation>
|
||
The presence of vasculature extending to the external bone surface and the absence of an external fundamental system indicate that the individual was actively growing when it died. In many respects, the bone microstructure is similar to small, young individuals (<30% adult wingspan; size class I
|
||
<superScript attach="left" box="[755,774,1348,1362]" fontSize="6" pageId="3" pageNumber="3">12</superScript>
|
||
) of
|
||
<taxonomicName box="[176,365,1382,1403]" class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[176,365,1382,1403]" italics="true" pageId="3" pageNumber="3">Rhamphorhynchus</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Prondvai, E. & Stein, K. & Osi ", A. & Sander, M. P." box="[365,384,1377,1391]" journalOrPublisher="PLoS ONE" pageId="3" pageNumber="3" pagination="31392" part="7" refId="ref6203" refString="16. Prondvai, E., Stein, K., Osi ", A., and Sander, M. P. (2012). Life history of Rhamphorhynchus inferred from bone histology and the diversity of pterosaurian growth strategies. PLoS ONE 7, e 31392." title="Life history of Rhamphorhynchus inferred from bone histology and the diversity of pterosaurian growth strategies" type="journal article" year="2012">
|
||
<superScript attach="left" box="[365,384,1377,1391]" fontSize="6" pageId="3" pageNumber="3">16</superScript>
|
||
</bibRefCitation>
|
||
and other actively growing juvenile pterosaurs,
|
||
<superScript attach="left" box="[256,298,1406,1420]" fontSize="6" pageId="3" pageNumber="3">
|
||
<bibRefCitation author="Wang, X. & Kellner, A. W. A. & Jiang, S. & Cheng, X. & Wang, Q. & Ma, Y. & Paidoula, Y. & Rodrigues, T. & Chen, H. & Sayao, J. M. & al." box="[256,275,1406,1420]" journalOrPublisher="Science" pageId="3" pageNumber="3" pagination="1197 - 1201" part="358" refId="ref6254" refString="17. Wang, X., Kellner, A. W. A., Jiang, S., Cheng, X., Wang, Q., Ma, Y., Paidoula, Y., Rodrigues, T., Chen, H., Sayao, J. M., et al. (2017). Egg accumulation with 3 D embryos provides insight into the life history of a pterosaur. Science 358, 1197 - 1201." title="Egg accumulation with 3 D embryos provides insight into the life history of a pterosaur" type="journal article" year="2017">17</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Padian, K. & Horner, J. R. & De Ricqles, A." box="[279,298,1406,1420]" journalOrPublisher="J. Vertebr. Paleontol." pageId="3" pageNumber="3" pagination="555 - 571" part="24" refId="ref6342" refString="18. Padian, K., Horner, J. R., and De Ricqles, A. (2004). Growth in small dinosaurs and pterosaurs: the evolution of archosaurian growth strategies. J. Vertebr. Paleontol. 24, 555 - 571." title="Growth in small dinosaurs and pterosaurs: the evolution of archosaurian growth strategies" type="journal article" year="2004">18</bibRefCitation>
|
||
</superScript>
|
||
indicating that it is best interpreted as a juvenile or subadult
|
||
<bibRefCitation author="Woodward, H. N. & Freedman, E. A. & Farlow, J. O. & Horner, J. R." box="[261,280,1436,1450]" journalOrPublisher="Paleobiology" pageId="3" pageNumber="3" pagination="503 - 527" part="41" refId="ref6392" refString="19. Woodward, H. N., Freedman, E. A., Farlow, J. O., and Horner, J. R. (2015). Maiasaura, a model organism for extinct vertebrate population biology: a large sample statistical assessment of growth dynamics and survivorship. Paleobiology 41, 503 - 527." title="Maiasaura, a model organism for extinct vertebrate population biology: a large sample statistical assessment of growth dynamics and survivorship" type="journal article" year="2015">
|
||
<superScript attach="left" box="[261,280,1436,1450]" fontSize="6" pageId="3" pageNumber="3">19</superScript>
|
||
</bibRefCitation>
|
||
that had not reached adult body size when it perished.
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.6251712" ID-Zenodo-Dep="6251712" httpUri="https://zenodo.org/record/6251712/files/figure.png" pageId="3" pageNumber="3" startId="3.[142,202,940,959]" subCaptionStartIDs="3.[179,231,1219,1236]" subCaptionStarts="Figure S" targetBox="[143,1448,272,899]" targetPageId="3">
|
||
<paragraph blockId="3.[142,1450,940,1236]" box="[142,1354,940,959]" pageId="3" pageNumber="3">
|
||
Figure 2. Skull of the new Middle Jurassic pterosaur
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[645,824,940,958]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="species" species="sgiathanach">
|
||
<emphasis bold="true" box="[645,824,940,958]" italics="true" pageId="3" pageNumber="3">Dearc sgiathanach</emphasis>
|
||
</taxonomicName>
|
||
and comparisons of pterosaur brain and ear endocasts
|
||
</paragraph>
|
||
<paragraph blockId="3.[142,1450,940,1236]" pageId="3" pageNumber="3">
|
||
(A–L) Photographs (A–D), segmented
|
||
<emphasis bold="true" box="[452,462,967,984]" pageId="3" pageNumber="3">M</emphasis>
|
||
CT scan renderings (E–H), and schematic drawings (I–L) of the skull (NMS G.2021.6.2) in, from top to bottom, left lateral (reversed), dorsal, right lateral, and ventral views.
|
||
</paragraph>
|
||
<paragraph blockId="3.[142,1450,940,1236]" pageId="3" pageNumber="3">
|
||
(M–U) Brain and inner ear endocasts of
|
||
<taxonomicName box="[466,622,1017,1034]" class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[466,622,1017,1034]" italics="true" pageId="3" pageNumber="3">Rhamphorhynchus</emphasis>
|
||
</taxonomicName>
|
||
(based on Witmer et al.
|
||
<bibRefCitation author="Witmer, L. M. & Chatterjee, S. & Franzosa, J. & Rowe, T." box="[815,823,1013,1025]" journalOrPublisher="Nature" pageId="3" pageNumber="3" pagination="950 - 953" part="425" refId="ref5708" refString="7. Witmer, L. M., Chatterjee, S., Franzosa, J., and Rowe, T. (2003). Neuroanatomy of flying reptiles and implications for flight, posture and behaviour. Nature 425, 950 - 953." title="Neuroanatomy of flying reptiles and implications for flight, posture and behaviour" type="journal article" year="2003">
|
||
<superScript attach="none" box="[815,823,1013,1025]" fontSize="5" pageId="3" pageNumber="3">7</superScript>
|
||
</bibRefCitation>
|
||
) (M, P, and Q),
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[949,998,1017,1034]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[949,998,1017,1034]" italics="true" pageId="3" pageNumber="3">Dearc</emphasis>
|
||
</taxonomicName>
|
||
(N, R, and S), and
|
||
<taxonomicName authorityName="Kellner" authorityYear="1989" box="[1151,1222,1017,1034]" class="Reptilia" family="Tapejaridae" genus="Tapejara" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1151,1222,1017,1034]" italics="true" pageId="3" pageNumber="3">Tapejara</emphasis>
|
||
</taxonomicName>
|
||
(based on Eck et al.
|
||
<bibRefCitation author="Eck, K. E. & Elgin, R. A. & Frey, E." box="[1387,1395,1013,1025]" journalOrPublisher="Swiss J. Palaeontol." pageId="3" pageNumber="3" pagination="277" part="130" refId="ref5756" refString="8. Eck, K. E., Elgin, R. A., and Frey, E. (2011). On the osteology of Tapejara wellnhoferi Kellner 1989 and the first occurrence of a multiple specimen assemblage from the Santana Formation, Araripe Basin, NE-Brazil. Swiss J. Palaeontol. 130, 277." title="On the osteology of Tapejara wellnhoferi Kellner 1989 and the first occurrence of a multiple specimen assemblage from the Santana Formation, Araripe Basin, NE-Brazil" type="journal article" year="2011">
|
||
<superScript attach="none" box="[1387,1395,1013,1025]" fontSize="5" pageId="3" pageNumber="3">8</superScript>
|
||
</bibRefCitation>
|
||
) (O, T, and U), shown in the skull in left lateral view and isolated in left lateral and dorsal views, respectively.
|
||
</paragraph>
|
||
<paragraph blockId="3.[142,1450,940,1236]" pageId="3" pageNumber="3">
|
||
asc, anterior semi-circular canal; bp, basisphenoid; cbl, cerebellum; cer, cerebrum; ch, choana; chl, channels; cv, third cervical; d, dentary; den, dentition; e, endocast; ec, ectopterygoid; et, edentulous anterior tip; f, frontal (?); hy, hyoid; ipv, interpterygoid vacuity; itf, inferior temporal fenestra; j, jugal; la, lacrimal; md, matrix infilled dent; mx, maxilla; n, nasal; olf, olfactory bulb; opl, optic lobe; p, parietal; pm, premaxilla; po, postorbital; ppd, post-palatal depression; psc, posterior semi-circular canal; ptf, pneumatic foramen; q, quadrate; qj, quadrojugal; rapr, retroarticular process; rh, probable placement of rhamphotheca; sof, suborbital fenestra; sq,squamosal;sbtf,subtemporal fenestra;stf, superior temporal fenestra.In (E)–(L), bones that cannot be distinguished in the
|
||
<emphasis bold="true" box="[1335,1345,1169,1186]" pageId="3" pageNumber="3">M</emphasis>
|
||
CT scan due to fusion or insufficient resolution are conservatively rendered together in one color. Scale bars, 30 mm.
|
||
</paragraph>
|
||
<paragraph blockId="3.[142,1450,940,1236]" box="[142,264,1219,1236]" pageId="3" pageNumber="3">
|
||
See
|
||
<figureCitation box="[179,258,1219,1236]" captionStart="Figure 2" captionStartId="3.[142,202,940,959]" captionTargetBox="[143,1448,272,899]" captionTargetId="figure-424@3.[142,1449,271,900]" captionTargetPageId="3" captionText="Figure 2. Skull of the new Middle Jurassic pterosaur Dearc sgiathanach and comparisons of pterosaur brain and ear endocasts" figureDoi="http://doi.org/10.5281/zenodo.6251712" httpUri="https://zenodo.org/record/6251712/files/figure.png" pageId="3" pageNumber="3">Figure S2</figureCitation>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="3.[142,780,1531,1987]" box="[142,412,1531,1549]" pageId="3" pageNumber="3">
|
||
<heading bold="true" box="[142,412,1531,1549]" fontSize="8" level="6" pageId="3" pageNumber="3" reason="6">
|
||
<emphasis bold="true" box="[142,412,1531,1549]" pageId="3" pageNumber="3">Wingspan and body size</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="3.[142,780,1531,1987]" lastBlockId="3.[811,1450,1294,1870]" pageId="3" pageNumber="3">
|
||
Wingspan—defined as double the summed lengths of the bones of a single wing
|
||
<bibRefCitation author="Benson, R. B. J. & Frigot, R. A. & Goswami, A. & Andres, B. & Butler, R. J." box="[303,312,1582,1596]" journalOrPublisher="Nat. Commun." pageId="3" pageNumber="3" pagination="3567" part="5" refId="ref5589" refString="5. Benson, R. B. J., Frigot, R. A., Goswami, A., Andres, B., and Butler, R. J. (2014). Competition and constraint drove Cope's rule in the evolution of giant flying reptiles. Nat. Commun. 5, 3567." title="Competition and constraint drove Cope's rule in the evolution of giant flying reptiles" type="journal article" year="2014">
|
||
<superScript attach="right" box="[303,312,1582,1596]" fontSize="6" pageId="3" pageNumber="3">5</superScript>
|
||
</bibRefCitation>
|
||
—is tightly correlated to body mass and wing area in pterosaurs and thus a robust body size proxy.
|
||
<superScript attach="left" box="[680,699,1611,1625]" fontSize="6" pageId="3" pageNumber="3">20</superScript>
|
||
A complete wingspan cannot be measured directly from NMS G.2021.6.1–4 because some wing phalanges are missing. To estimate wingspan, we compiled measurements of complete wingspans of two non-monofenestratans represented by large sample sizes—
|
||
<taxonomicName box="[250,439,1762,1783]" class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[250,439,1762,1783]" italics="true" pageId="3" pageNumber="3">Rhamphorhynchus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName authorityName="Wagner" authorityYear="1860" box="[486,613,1762,1783]" class="Reptilia" family="Rhamphorhynchidae" genus="Dorygnathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[486,613,1762,1783]" italics="true" pageId="3" pageNumber="3">Dorygnathus</emphasis>
|
||
</taxonomicName>
|
||
—and regressed these against the lengths of individual bones to create predictor formulas (STAR Methods; Data S1C–S1Q). Using
|
||
<taxonomicName class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="3" pageNumber="3">Rhamphorhynchus</emphasis>
|
||
</taxonomicName>
|
||
scaling, the humerus length (112 mm) and skull length (222 mm) of NMS G.2021.6.1–4 indicate wingspans of 3.8 and 2.2 m, respectively. The largest known
|
||
<taxonomicName box="[534,723,1908,1929]" class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[534,723,1908,1929]" italics="true" pageId="3" pageNumber="3">Rhamphorhynchus</emphasis>
|
||
</taxonomicName>
|
||
(Natural History Museum UK 37002) is considerably smaller, with a wingspan of 1.8 m, humeral length of 79 mm, and skull length of 202 mm. Using
|
||
<taxonomicName authorityName="Wagner" authorityYear="1860" box="[1006,1133,1294,1315]" class="Reptilia" family="Rhamphorhynchidae" genus="Dorygnathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1006,1133,1294,1315]" italics="true" pageId="3" pageNumber="3">Dorygnathus</emphasis>
|
||
</taxonomicName>
|
||
scaling, the humeral length of NMS G.2021.6.1–4 indicates a wingspan of 1.9 m, approximately 10% larger than the largest
|
||
<taxonomicName authorityName="Wagner" authorityYear="1860" box="[1173,1300,1353,1374]" class="Reptilia" family="Rhamphorhynchidae" genus="Dorygnathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1173,1300,1353,1374]" italics="true" pageId="3" pageNumber="3">Dorygnathus</emphasis>
|
||
</taxonomicName>
|
||
(1.69 m wingspan, 84 mm humerus).
|
||
</paragraph>
|
||
<paragraph blockId="3.[811,1450,1294,1870]" pageId="3" pageNumber="3">
|
||
These results demonstrate that
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[1159,1219,1411,1432]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1159,1219,1411,1432]" italics="true" pageId="3" pageNumber="3">Dearc</emphasis>
|
||
</taxonomicName>
|
||
is the largest Jurassic pterosaur yet known, consistent with the fact that its humerus and skull are the longest of any Jurassic specimens. Furthermore, we interpret these results as evidence that
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[1330,1390,1499,1520]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1330,1390,1499,1520]" italics="true" pageId="3" pageNumber="3">Dearc</emphasis>
|
||
</taxonomicName>
|
||
likely achieved wingspans over 2.5 m, and perhaps larger (>3 m). This is based on two lines of reasoning. First, we consider the
|
||
<taxonomicName box="[811,1000,1586,1607]" class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[811,1000,1586,1607]" italics="true" pageId="3" pageNumber="3">Rhamphorhynchus</emphasis>
|
||
</taxonomicName>
|
||
equations, which give larger wingspan estimates, as the most valid predictors:
|
||
<taxonomicName box="[1209,1398,1616,1637]" class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1209,1398,1616,1637]" italics="true" pageId="3" pageNumber="3">Rhamphorhynchus</emphasis>
|
||
</taxonomicName>
|
||
is a closer relative of
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[991,1051,1645,1666]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[991,1051,1645,1666]" italics="true" pageId="3" pageNumber="3">Dearc</emphasis>
|
||
</taxonomicName>
|
||
than is
|
||
<taxonomicName authorityName="Wagner" authorityYear="1860" box="[1138,1265,1645,1666]" class="Reptilia" family="Rhamphorhynchidae" genus="Dorygnathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1138,1265,1645,1666]" italics="true" pageId="3" pageNumber="3">Dorygnathus</emphasis>
|
||
</taxonomicName>
|
||
, is known from a larger sample size (and thus generates a regression with tighter error bars and a higher r
|
||
<superScript attach="left" box="[1052,1061,1699,1713]" fontSize="6" pageId="3" pageNumber="3">2</superScript>
|
||
value), and has a well-established and nearly isometric growth trajectory that makes predicting wingspan from isolated skeletal elements more justifiable.
|
||
<bibRefCitation author="Hone, D. & Ratcliffe, J. M. & Riski, D. K. & Hermanson, J. W. & Reisz, R. R." box="[1340,1359,1757,1771]" journalOrPublisher="Lethaia" pageId="3" pageNumber="3" pagination="106 - 112" part="54" refId="ref6506" refString="21. Hone, D., Ratcliffe, J. M., Riski, D. K., Hermanson, J. W., and Reisz, R. R. (2021). Unique near isometric ontogeny in the pterosaur Rhamphorhynchus suggests hatchlings could fly. Lethaia 54, 106 - 112." title="Unique near isometric ontogeny in the pterosaur Rhamphorhynchus suggests hatchlings could fly" type="journal article" year="2021">
|
||
<superScript attach="left" box="[1340,1359,1757,1771]" fontSize="6" pageId="3" pageNumber="3">21</superScript>
|
||
</bibRefCitation>
|
||
Second, the holotype of
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[987,1047,1791,1812]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[987,1047,1791,1812]" italics="true" pageId="3" pageNumber="3">Dearc</emphasis>
|
||
</taxonomicName>
|
||
(NMS G.2021.6.1–4) was an actively growing juvenile-subadult at death and would have been larger as an adult (STAR Methods).
|
||
</paragraph>
|
||
<paragraph blockId="3.[811,1449,1911,1987]" box="[811,941,1911,1929]" pageId="3" pageNumber="3">
|
||
<heading bold="true" box="[811,941,1911,1929]" fontSize="8" level="6" pageId="3" pageNumber="3" reason="6">
|
||
<emphasis bold="true" box="[811,941,1911,1929]" pageId="3" pageNumber="3">Description</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="3.[811,1449,1911,1987]" lastBlockId="4.[829,1466,1733,1987]" lastPageId="4" lastPageNumber="4" pageId="3" pageNumber="3">
|
||
A detailed description is provided in Data S2, with salient features summarized here.
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[1053,1113,1966,1987]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="3" pageNumber="3" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1053,1113,1966,1987]" italics="true" pageId="3" pageNumber="3">Dearc</emphasis>
|
||
</taxonomicName>
|
||
generally conforms to the classic non-monofenestratan body plan, as it has an elongate mandibular symphysis (>20% mandible length), cervical ribs (visible in
|
||
<emphasis bold="true" box="[159,172,1795,1812]" pageId="4" pageNumber="4">M</emphasis>
|
||
CT data of anterior cervicals), a neck shorter than the combined dorsal and sacral series, a short metacarpus (<80% humerus length), and an elongate tail comprised of elongate caudal vertebrae supported by interlocking zygapophyses (
|
||
<figureCitation box="[630,746,1879,1900]" captionStart="Figure 1" captionStartId="2.[1056,1116,271,290]" captionTargetBox="[160,1018,273,1413]" captionTargetId="figure-152@2.[159,1020,271,1414]" captionTargetPageId="2" captionText="Figure 1. The new Middle Jurassic pterosaur Dearc sgiathanach (A–C) Photographs of main slab (NMS G.2021.6.1– 2), bones in dorsal view (A); wing phalanges 2–3 (NMS G.2021.6.3–4), in dorsal view (B); and main counterslab (NMS G.2021.6.3), bones in ventral view (C). (D–F) Schematic drawings of (A)–(C). (G–H) Reconstruction of skull in dorsal (G) and ventral (I) views and skeleton in lateral view (H). ar, articular region; cd, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, digit; den, dentary;dors,dorsal vertebrae;dpc,deltopectoral crest; ep, extensor process; fm, femur; gas, gastralia; hdc, humeral distal condyle; hmt, humeral tubercle; isc, ischium; ju, jugal; max, maxilla; mc, metacarpal; mt,metatarsal;po,postorbital;r, ribs; sac, sacral plate; sca, scapula; scv, sacral (?) vertebrae; sm, sesamoid; sq, squamosal; st, sternum; symp, symphysis; uc, ulnar crest; ul, ulna; wp, wing phalanx; r/l, right/left. Blue on reconstructions are missing regions; red line in (E) is location of histological sectioning. Scale bars, 30 mm. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251706" httpUri="https://zenodo.org/record/6251706/files/figure.png" pageId="4" pageNumber="4">Figures 1, 2</figureCitation>
|
||
, and 3). It does, however, possess some features typical of pterodactyloids and often considered part of a ‘‘module’’ unique to their bauplan,
|
||
<bibRefCitation author="Lu, J. & Unwin, D. M. & Jin, X. & Liu, Y. & Ji, Q." box="[247,256,1962,1976]" journalOrPublisher="Proc. Biol. Sci." pageId="4" pageNumber="4" pagination="383 - 389" part="277" refId="ref5650" refString="6. Lu, J., Unwin, D. M., Jin, X., Liu, Y., and Ji, Q. (2010). Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proc. Biol. Sci. 277, 383 - 389." title="Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull" type="journal article" year="2010">
|
||
<superScript attach="left" box="[247,256,1962,1976]" fontSize="6" pageId="4" pageNumber="4">6</superScript>
|
||
</bibRefCitation>
|
||
including a skull that is longer than the combined dorsal and sacral series and a highly inclined quadrate (
|
||
<figureCitation box="[1374,1454,1733,1754]" captionStart="Figure 1" captionStartId="2.[1056,1116,271,290]" captionTargetBox="[160,1018,273,1413]" captionTargetId="figure-152@2.[159,1020,271,1414]" captionTargetPageId="2" captionText="Figure 1. The new Middle Jurassic pterosaur Dearc sgiathanach (A–C) Photographs of main slab (NMS G.2021.6.1– 2), bones in dorsal view (A); wing phalanges 2–3 (NMS G.2021.6.3–4), in dorsal view (B); and main counterslab (NMS G.2021.6.3), bones in ventral view (C). (D–F) Schematic drawings of (A)–(C). (G–H) Reconstruction of skull in dorsal (G) and ventral (I) views and skeleton in lateral view (H). ar, articular region; cd, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, digit; den, dentary;dors,dorsal vertebrae;dpc,deltopectoral crest; ep, extensor process; fm, femur; gas, gastralia; hdc, humeral distal condyle; hmt, humeral tubercle; isc, ischium; ju, jugal; max, maxilla; mc, metacarpal; mt,metatarsal;po,postorbital;r, ribs; sac, sacral plate; sca, scapula; scv, sacral (?) vertebrae; sm, sesamoid; sq, squamosal; st, sternum; symp, symphysis; uc, ulnar crest; ul, ulna; wp, wing phalanx; r/l, right/left. Blue on reconstructions are missing regions; red line in (E) is location of histological sectioning. Scale bars, 30 mm. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251706" httpUri="https://zenodo.org/record/6251706/files/figure.png" pageId="4" pageNumber="4">Figure 2</figureCitation>
|
||
). Furthermore, although the cervical vertebrae are short and squat as in non-monofenestratans, they are proportionally more elongate than most members of that grade, beginning to approach the proportions of more derived pterosaurs like
|
||
<emphasis italics="true" pageId="4" pageNumber="4">Wukongopterus lii</emphasis>
|
||
and
|
||
<emphasis box="[897,1135,1879,1900]" italics="true" pageId="4" pageNumber="4">Douzhanopterus zhengi</emphasis>
|
||
<superScript attach="left" box="[1135,1177,1874,1888]" fontSize="6" pageId="4" pageNumber="4">
|
||
<bibRefCitation author="Wang, X. & Kellner, A. W. & Jiang, S. & Meng, X." box="[1135,1154,1874,1888]" journalOrPublisher="An. Acad. Bras. Cienc." pageId="4" pageNumber="4" pagination="793 - 812" part="81" refId="ref6565" refString="22. Wang, X., Kellner, A. W., Jiang, S., and Meng, X. (2009). An unusual longtailed pterosaur with elongated neck from western Liaoning of China. An. Acad. Bras. Cienc. 81, 793 - 812." title="An unusual longtailed pterosaur with elongated neck from western Liaoning of China" type="journal article" year="2009">22</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Wang, X. & Jiang, S. & Zhang, J. & Cheng, X. & Yu, X. & Li, Y. & Wei, G. & Wang, X." box="[1158,1177,1874,1888]" journalOrPublisher="Sci. Rep." pageId="4" pageNumber="4" pagination="42763" part="7" refId="ref6620" refString="23. Wang, X., Jiang, S., Zhang, J., Cheng, X., Yu, X., Li, Y., Wei, G., and Wang, X. (2017). New evidence from China for the nature of the pterosaur evolutionary transition. Sci. Rep. 7, 42763." title="New evidence from China for the nature of the pterosaur evolutionary transition" type="journal article" year="2017">23</bibRefCitation>
|
||
</superScript>
|
||
(STAR Methods). There is a continuum between two distinct types of dentition: elongate fangs at the snout tip and conical pegs along much of the jaw length (
|
||
<figureCitation box="[903,996,1966,1987]" captionStart="Figure 1" captionStartId="2.[1056,1116,271,290]" captionTargetBox="[160,1018,273,1413]" captionTargetId="figure-152@2.[159,1020,271,1414]" captionTargetPageId="2" captionText="Figure 1. The new Middle Jurassic pterosaur Dearc sgiathanach (A–C) Photographs of main slab (NMS G.2021.6.1– 2), bones in dorsal view (A); wing phalanges 2–3 (NMS G.2021.6.3–4), in dorsal view (B); and main counterslab (NMS G.2021.6.3), bones in ventral view (C). (D–F) Schematic drawings of (A)–(C). (G–H) Reconstruction of skull in dorsal (G) and ventral (I) views and skeleton in lateral view (H). ar, articular region; cd, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, digit; den, dentary;dors,dorsal vertebrae;dpc,deltopectoral crest; ep, extensor process; fm, femur; gas, gastralia; hdc, humeral distal condyle; hmt, humeral tubercle; isc, ischium; ju, jugal; max, maxilla; mc, metacarpal; mt,metatarsal;po,postorbital;r, ribs; sac, sacral plate; sca, scapula; scv, sacral (?) vertebrae; sm, sesamoid; sq, squamosal; st, sternum; symp, symphysis; uc, ulnar crest; ul, ulna; wp, wing phalanx; r/l, right/left. Blue on reconstructions are missing regions; red line in (E) is location of histological sectioning. Scale bars, 30 mm. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251706" httpUri="https://zenodo.org/record/6251706/files/figure.png" pageId="4" pageNumber="4">Figures 2</figureCitation>
|
||
and
|
||
<figureCitation box="[1047,1059,1966,1987]" captionStart="Figure 3" captionStartId="4.[159,219,1471,1490]" captionTargetBox="[161,1465,271,1431]" captionTargetId="figure-185@4.[161,1465,271,1431]" captionTargetPageId="4" captionText="Figure 3. Postcranial skeleton and dentition of the new Middle Jurassic pterosaur Dearc sgiathanach Photographs of the right manus (A), cervical series (B), pubic region (C), right humerus (D), left humerus (E), left metacarpal-phalanx articular region (F), right maxilla (G), and left pes (H) of NMS G.2021.6.1–4. ac, anterior caudal vertebrae; ar, articular region; c, condyle; cdv, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, dentary; dc, distal condyle; dpc, deltopectoral crest;dvrt, dorsal vertebrae; en,enamel;exp,extensor process; fem,femur; ft, flexor tubercle; gas,gastralia; h, humerus;ic, intercondylar groove; isc,ischium; max,maxilla; mcp,metacarpal; md, maxillary dentition; mg, medial groove; mt, metatarsal; nss,neural spine scar; pa, preacetabular process; poz, postzygapophysis ppb,prepubis; prez,prezygapophysis; rad,radius;rb,rib; sac,sacrum;sm,sesamoid;uc,ulnar crest; ul,ulna;un, ungual;vert, vertebra;wp1, wing phalanx one. Scale, 10 mm per bar. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251720" httpUri="https://zenodo.org/record/6251720/files/figure.png" pageId="4" pageNumber="4">3</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.6251716" ID-Zenodo-Dep="6251716" httpUri="https://zenodo.org/record/6251716/files/figure.png" pageId="4" pageNumber="4" startId="4.[159,219,1471,1490]" subCaptionStartIDs="4.[197,258,1673,1690]" subCaptionStarts="Figures S" targetBox="[161,1465,271,1431]" targetPageId="4">
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.6251720" ID-Zenodo-Dep="6251720" httpUri="https://zenodo.org/record/6251720/files/figure.png" pageId="4" pageNumber="4" startId="4.[159,219,1471,1490]" targetBox="[161,1465,271,1431]" targetPageId="4">
|
||
<paragraph blockId="4.[159,1468,1471,1690]" box="[159,1121,1471,1490]" pageId="4" pageNumber="4">
|
||
<emphasis bold="true" box="[159,1121,1471,1490]" italics="true" pageId="4" pageNumber="4">
|
||
Figure 3. Postcranial skeleton and dentition of the new Middle Jurassic pterosaur
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[942,1121,1471,1489]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="4" pageNumber="4" phylum="Chordata" rank="species" species="sgiathanach">Dearc sgiathanach</taxonomicName>
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="4.[159,1468,1471,1690]" pageId="4" pageNumber="4">Photographs of the right manus (A), cervical series (B), pubic region (C), right humerus (D), left humerus (E), left metacarpal-phalanx articular region (F), right maxilla (G), and left pes (H) of NMS G.2021.6.1–4.</paragraph>
|
||
</caption>
|
||
<paragraph blockId="4.[159,1468,1471,1690]" pageId="4" pageNumber="4">ac, anterior caudal vertebrae; ar, articular region; c, condyle; cdv, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, dentary; dc, distal condyle; dpc, deltopectoral crest;dvrt, dorsal vertebrae; en,enamel;exp,extensor process; fem,femur; ft, flexor tubercle; gas,gastralia; h, humerus;ic, intercondylar groove; isc,ischium; max,maxilla; mcp,metacarpal; md, maxillary dentition; mg, medial groove; mt, metatarsal; nss,neural spine scar; pa, preacetabular process; poz, postzygapophysis ppb,prepubis; prez,prezygapophysis; rad,radius;rb,rib; sac,sacrum;sm,sesamoid;uc,ulnar crest; ul,ulna;un, ungual;vert, vertebra;wp1, wing phalanx one. Scale, 10 mm per bar.</paragraph>
|
||
<paragraph blockId="4.[159,1468,1471,1690]" box="[159,355,1673,1690]" pageId="4" pageNumber="4">
|
||
See
|
||
<figureCitation box="[197,285,1673,1690]" captionStart="Figure 2" captionStartId="3.[142,202,940,959]" captionTargetBox="[143,1448,272,899]" captionTargetId="figure-424@3.[142,1449,271,900]" captionTargetPageId="3" captionText="Figure 2. Skull of the new Middle Jurassic pterosaur Dearc sgiathanach and comparisons of pterosaur brain and ear endocasts (A–L) Photographs (A–D), segmented MCT scan renderings (E–H), and schematic drawings (I–L) of the skull (NMS G.2021.6.2) in, from top to bottom, left lateral (reversed), dorsal, right lateral, and ventral views. (M–U) Brain and inner ear endocasts of Rhamphorhynchus (based on Witmer et al.7) (M, P, and Q), Dearc (N, R, and S), and Tapejara (based on Eck et al.8) (O, T, and U), shown in the skull in left lateral view and isolated in left lateral and dorsal views, respectively. asc, anterior semi-circular canal; bp, basisphenoid; cbl, cerebellum; cer, cerebrum; ch, choana; chl, channels; cv, third cervical; d, dentary; den, dentition; e, endocast; ec, ectopterygoid; et, edentulous anterior tip; f, frontal (?); hy, hyoid; ipv, interpterygoid vacuity; itf, inferior temporal fenestra; j, jugal; la, lacrimal; md, matrix infilled dent; mx, maxilla; n, nasal; olf, olfactory bulb; opl, optic lobe; p, parietal; pm, premaxilla; po, postorbital; ppd, post-palatal depression; psc, posterior semi-circular canal; ptf, pneumatic foramen; q, quadrate; qj, quadrojugal; rapr, retroarticular process; rh, probable placement of rhamphotheca; sof, suborbital fenestra; sq,squamosal;sbtf,subtemporal fenestra;stf, superior temporal fenestra.In (E)–(L), bones that cannot be distinguished in the MCT scan due to fusion or insufficient resolution are conservatively rendered together in one color. Scale bars, 30 mm. See Figure S2." figureDoi="http://doi.org/10.5281/zenodo.6251712" httpUri="https://zenodo.org/record/6251712/files/figure.png" pageId="4" pageNumber="4">Figures S2</figureCitation>
|
||
and S
|
||
<figureCitation box="[340,350,1673,1690]" captionStart="Figure 3" captionStartId="4.[159,219,1471,1490]" captionTargetBox="[161,1465,271,1431]" captionTargetId="figure-185@4.[161,1465,271,1431]" captionTargetPageId="4" captionText="Figure 3. Postcranial skeleton and dentition of the new Middle Jurassic pterosaur Dearc sgiathanach Photographs of the right manus (A), cervical series (B), pubic region (C), right humerus (D), left humerus (E), left metacarpal-phalanx articular region (F), right maxilla (G), and left pes (H) of NMS G.2021.6.1–4." figureDoi="http://doi.org/10.5281/zenodo.6251720" httpUri="https://zenodo.org/record/6251720/files/figure.png" pageId="4" pageNumber="4">3</figureCitation>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.6251726" ID-Zenodo-Dep="6251726" httpUri="https://zenodo.org/record/6251726/files/figure.png" pageId="5" pageNumber="5" startId="5.[142,202,1156,1175]" targetBox="[143,1448,272,1113]" targetPageId="5">
|
||
<paragraph blockId="5.[142,1450,1156,1351]" pageId="5" pageNumber="5">
|
||
<emphasis bold="true" box="[142,1449,1156,1175]" pageId="5" pageNumber="5">
|
||
Figure 4. Phylogenetic relationships of the Middle Jurassic pterosaur
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[806,984,1157,1175]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="sgiathanach">
|
||
<emphasis bold="true" box="[806,984,1157,1175]" italics="true" pageId="5" pageNumber="5">Dearc sgiathanach</emphasis>
|
||
</taxonomicName>
|
||
and wingspan estimates for Jurassic pterosaurs
|
||
</emphasis>
|
||
Strict consensus of most parsimonious trees from phylogenetic analysis,with silhouettes scaled to wingspan (
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[1040,1089,1182,1199]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1040,1089,1182,1199]" italics="true" pageId="5" pageNumber="5">Dearc</emphasis>
|
||
</taxonomicName>
|
||
= ca.2.0 m) (1), and skull reconstructions of key taxa: (A)
|
||
<emphasis box="[247,481,1208,1225]" italics="true" pageId="5" pageNumber="5">Scaphognathus crassirostris</emphasis>
|
||
(based on GPIB 1304), (B)
|
||
<emphasis box="[703,902,1208,1225]" italics="true" pageId="5" pageNumber="5">
|
||
<taxonomicName authorityName="Wagner" authorityYear="1860" box="[703,808,1208,1225]" class="Reptilia" family="Rhamphorhynchidae" genus="Dorygnathus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">Dorygnathus</taxonomicName>
|
||
banthensis
|
||
</emphasis>
|
||
(based on SMNS 55886), (C)
|
||
<emphasis box="[1145,1388,1208,1225]" italics="true" pageId="5" pageNumber="5">
|
||
<taxonomicName box="[1145,1301,1208,1225]" class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">Rhamphorhynchus</taxonomicName>
|
||
muensteri
|
||
</emphasis>
|
||
(based on NHMUK R 37002), (D)
|
||
<emphasis box="[353,617,1233,1250]" italics="true" pageId="5" pageNumber="5">
|
||
<taxonomicName box="[353,498,1233,1250]" class="Reptilia" family="Rhamphorhynchidae" genus="Angustinaripterus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">Angustinaripterus</taxonomicName>
|
||
longicephalus
|
||
</emphasis>
|
||
(based on ZDM T8001), and (E)
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[884,1037,1233,1250]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="sgiathanach">
|
||
<emphasis box="[884,1037,1233,1250]" italics="true" pageId="5" pageNumber="5">Dearc sgiathanach</emphasis>
|
||
</taxonomicName>
|
||
. Wingspan plot (2): estimated wingspan range for isolated pterosaur remains from the Taynton Limestone collection. See Data S1D for complete element list. (A) NHMUK PV R 36634, (B) GSM 113726, (C) OUM J28352,(D) NHMUK PV R 38016, (E) NHMUK PV R1362, (F) LL12158, (G) NHMUK PV R 40126,(H) NMS G.2021.6.1-4 (
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[1110,1159,1283,1300]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1110,1159,1283,1300]" italics="true" pageId="5" pageNumber="5">Dearc</emphasis>
|
||
</taxonomicName>
|
||
), (I) OUM J23046, (J) OUM J28273, (K) NHM UK PV R 40126 B, (L) OUM J28319, (M) OUM J28307, (N) OUM J28271, (O) OUM J28354, (P) LL12160, (Q) OUM J23047, and (R) MJM L K1995.
|
||
</paragraph>
|
||
<paragraph blockId="5.[142,1450,1156,1351]" box="[142,436,1334,1351]" pageId="5" pageNumber="5">See Figure S4 and Data S1 and S2.</paragraph>
|
||
</caption>
|
||
<paragraph blockId="5.[142,781,1411,1987]" pageId="5" pageNumber="5">
|
||
<emphasis bold="true" box="[164,177,1415,1432]" pageId="5" pageNumber="5">M</emphasis>
|
||
CT data provide a stellar view of a complete, articulated palate and hyoid of a non-monofenestratan pterosaur in dorsal and ventral view (
|
||
<figureCitation box="[331,417,1470,1491]" captionStart="Figure 1" captionStartId="2.[1056,1116,271,290]" captionTargetBox="[160,1018,273,1413]" captionTargetId="figure-152@2.[159,1020,271,1414]" captionTargetPageId="2" captionText="Figure 1. The new Middle Jurassic pterosaur Dearc sgiathanach (A–C) Photographs of main slab (NMS G.2021.6.1– 2), bones in dorsal view (A); wing phalanges 2–3 (NMS G.2021.6.3–4), in dorsal view (B); and main counterslab (NMS G.2021.6.3), bones in ventral view (C). (D–F) Schematic drawings of (A)–(C). (G–H) Reconstruction of skull in dorsal (G) and ventral (I) views and skeleton in lateral view (H). ar, articular region; cd, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, digit; den, dentary;dors,dorsal vertebrae;dpc,deltopectoral crest; ep, extensor process; fm, femur; gas, gastralia; hdc, humeral distal condyle; hmt, humeral tubercle; isc, ischium; ju, jugal; max, maxilla; mc, metacarpal; mt,metatarsal;po,postorbital;r, ribs; sac, sacral plate; sca, scapula; scv, sacral (?) vertebrae; sm, sesamoid; sq, squamosal; st, sternum; symp, symphysis; uc, ulnar crest; ul, ulna; wp, wing phalanx; r/l, right/left. Blue on reconstructions are missing regions; red line in (E) is location of histological sectioning. Scale bars, 30 mm. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251706" httpUri="https://zenodo.org/record/6251706/files/figure.png" pageId="5" pageNumber="5">Figure 2</figureCitation>
|
||
). The heart-shaped choana is cut medially by forking vomers, comprised of three cylindrical rods that converge anteriorly in a trident-shaped contact. There is a thin extension of the ectopterygoid, which rotates around its own axis, forming an elevated ventral border of the postpalatine fenestra, joining the vomers at a perpendicular angle. This ‘‘contorted’’ morphology has not been described in other pterosaurs.
|
||
</paragraph>
|
||
<paragraph blockId="5.[142,781,1411,1987]" lastBlockId="5.[811,1449,1411,1491]" pageId="5" pageNumber="5">
|
||
<emphasis bold="true" box="[164,177,1707,1724]" pageId="5" pageNumber="5">M</emphasis>
|
||
CT data also provide one of the few brain and inner ear endocasts of a basal pterosaur (
|
||
<figureCitation box="[467,553,1733,1754]" captionStart="Figure 1" captionStartId="2.[1056,1116,271,290]" captionTargetBox="[160,1018,273,1413]" captionTargetId="figure-152@2.[159,1020,271,1414]" captionTargetPageId="2" captionText="Figure 1. The new Middle Jurassic pterosaur Dearc sgiathanach (A–C) Photographs of main slab (NMS G.2021.6.1– 2), bones in dorsal view (A); wing phalanges 2–3 (NMS G.2021.6.3–4), in dorsal view (B); and main counterslab (NMS G.2021.6.3), bones in ventral view (C). (D–F) Schematic drawings of (A)–(C). (G–H) Reconstruction of skull in dorsal (G) and ventral (I) views and skeleton in lateral view (H). ar, articular region; cd, caudal vertebrae; cor, coracoid; cv, cervical vertebrae; d, digit; den, dentary;dors,dorsal vertebrae;dpc,deltopectoral crest; ep, extensor process; fm, femur; gas, gastralia; hdc, humeral distal condyle; hmt, humeral tubercle; isc, ischium; ju, jugal; max, maxilla; mc, metacarpal; mt,metatarsal;po,postorbital;r, ribs; sac, sacral plate; sca, scapula; scv, sacral (?) vertebrae; sm, sesamoid; sq, squamosal; st, sternum; symp, symphysis; uc, ulnar crest; ul, ulna; wp, wing phalanx; r/l, right/left. Blue on reconstructions are missing regions; red line in (E) is location of histological sectioning. Scale bars, 30 mm. See Figures S2 and S3." figureDoi="http://doi.org/10.5281/zenodo.6251706" httpUri="https://zenodo.org/record/6251706/files/figure.png" pageId="5" pageNumber="5">Figure 2</figureCitation>
|
||
). Like
|
||
<taxonomicName class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="5" pageNumber="5">Rhamphorhynchus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<bibRefCitation author="Witmer, L. M. & Chatterjee, S. & Franzosa, J. & Rowe, T." box="[196,205,1757,1771]" journalOrPublisher="Nature" pageId="5" pageNumber="5" pagination="950 - 953" part="425" refId="ref5708" refString="7. Witmer, L. M., Chatterjee, S., Franzosa, J., and Rowe, T. (2003). Neuroanatomy of flying reptiles and implications for flight, posture and behaviour. Nature 425, 950 - 953." title="Neuroanatomy of flying reptiles and implications for flight, posture and behaviour" type="journal article" year="2003">
|
||
<superScript attach="left" box="[196,205,1757,1771]" fontSize="6" pageId="5" pageNumber="5">7</superScript>
|
||
</bibRefCitation>
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[212,272,1762,1783]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[212,272,1762,1783]" italics="true" pageId="5" pageNumber="5">Dearc</emphasis>
|
||
</taxonomicName>
|
||
had a large cerebrum with optic lobes positioned at the same level as the forebrain and a large flocculus, around which thin and arched semicircular canals looped, that nonetheless did not project to the same lateral level as the cerebrum. In pterodactyloids, however, the brain is highly flexed so that the cerebrum is elevated relative to the optic lobes, and the flocculus is expanded beyond the cerebrum laterally.
|
||
<superScript attach="left" box="[181,229,1962,1976]" fontSize="6" pageId="5" pageNumber="5">
|
||
<bibRefCitation author="Witmer, L. M. & Chatterjee, S. & Franzosa, J. & Rowe, T." box="[181,190,1962,1976]" journalOrPublisher="Nature" pageId="5" pageNumber="5" pagination="950 - 953" part="425" refId="ref5708" refString="7. Witmer, L. M., Chatterjee, S., Franzosa, J., and Rowe, T. (2003). Neuroanatomy of flying reptiles and implications for flight, posture and behaviour. Nature 425, 950 - 953." title="Neuroanatomy of flying reptiles and implications for flight, posture and behaviour" type="journal article" year="2003">7</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Eck, K. E. & Elgin, R. A. & Frey, E." box="[195,204,1962,1976]" journalOrPublisher="Swiss J. Palaeontol." pageId="5" pageNumber="5" pagination="277" part="130" refId="ref5756" refString="8. Eck, K. E., Elgin, R. A., and Frey, E. (2011). On the osteology of Tapejara wellnhoferi Kellner 1989 and the first occurrence of a multiple specimen assemblage from the Santana Formation, Araripe Basin, NE-Brazil. Swiss J. Palaeontol. 130, 277." title="On the osteology of Tapejara wellnhoferi Kellner 1989 and the first occurrence of a multiple specimen assemblage from the Santana Formation, Araripe Basin, NE-Brazil" type="journal article" year="2011">8</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Codorniu, L. & Paulina Carabajal, A. & Pol, D. & Unwin, D. & Rauhut, O. W." box="[210,229,1962,1976]" journalOrPublisher="PeerJ" pageId="5" pageNumber="5" pagination="2311" part="4" refId="ref6687" refString="24. Codorniu, L., Paulina Carabajal, A., Pol, D., Unwin, D., and Rauhut, O. W. (2016). A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium. PeerJ 4, e 2311." title="A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium" type="journal article" year="2016">24</bibRefCitation>
|
||
</superScript>
|
||
In
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[262,322,1966,1987]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[262,322,1966,1987]" italics="true" pageId="5" pageNumber="5">Dearc</emphasis>
|
||
</taxonomicName>
|
||
, the optic lobes are larger, anteroposteriorly longer, and more widely exposed dorsally than in any known basal pterosaur or pterodactyloid (
|
||
<taxonomicName box="[1167,1356,1440,1461]" class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1167,1356,1440,1461]" italics="true" pageId="5" pageNumber="5">Rhamphorhynchus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<bibRefCitation author="Witmer, L. M. & Chatterjee, S. & Franzosa, J. & Rowe, T." box="[1362,1371,1436,1450]" journalOrPublisher="Nature" pageId="5" pageNumber="5" pagination="950 - 953" part="425" refId="ref5708" refString="7. Witmer, L. M., Chatterjee, S., Franzosa, J., and Rowe, T. (2003). Neuroanatomy of flying reptiles and implications for flight, posture and behaviour. Nature 425, 950 - 953." title="Neuroanatomy of flying reptiles and implications for flight, posture and behaviour" type="journal article" year="2003">
|
||
<superScript attach="left" box="[1362,1371,1436,1450]" fontSize="6" pageId="5" pageNumber="5">7</superScript>
|
||
</bibRefCitation>
|
||
<emphasis italics="true" pageId="5" pageNumber="5">Allkauren</emphasis>
|
||
,
|
||
<bibRefCitation author="Codorniu, L. & Paulina Carabajal, A. & Pol, D. & Unwin, D. & Rauhut, O. W." box="[850,869,1465,1479]" journalOrPublisher="PeerJ" pageId="5" pageNumber="5" pagination="2311" part="4" refId="ref6687" refString="24. Codorniu, L., Paulina Carabajal, A., Pol, D., Unwin, D., and Rauhut, O. W. (2016). A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium. PeerJ 4, e 2311." title="A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium" type="journal article" year="2016">
|
||
<superScript attach="left" box="[850,869,1465,1479]" fontSize="6" pageId="5" pageNumber="5">24</superScript>
|
||
</bibRefCitation>
|
||
and
|
||
<taxonomicName authorityName="Kellner" authorityYear="1989" box="[922,1008,1470,1491]" class="Reptilia" family="Tapejaridae" genus="Tapejara" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[922,1008,1470,1491]" italics="true" pageId="5" pageNumber="5">Tapejara</emphasis>
|
||
</taxonomicName>
|
||
<bibRefCitation author="Eck, K. E. & Elgin, R. A. & Frey, E." box="[1009,1018,1465,1479]" journalOrPublisher="Swiss J. Palaeontol." pageId="5" pageNumber="5" pagination="277" part="130" refId="ref5756" refString="8. Eck, K. E., Elgin, R. A., and Frey, E. (2011). On the osteology of Tapejara wellnhoferi Kellner 1989 and the first occurrence of a multiple specimen assemblage from the Santana Formation, Araripe Basin, NE-Brazil. Swiss J. Palaeontol. 130, 277." title="On the osteology of Tapejara wellnhoferi Kellner 1989 and the first occurrence of a multiple specimen assemblage from the Santana Formation, Araripe Basin, NE-Brazil" type="journal article" year="2011">
|
||
<superScript attach="right" box="[1009,1018,1465,1479]" fontSize="6" pageId="5" pageNumber="5">8</superScript>
|
||
</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection lastPageId="6" lastPageNumber="6" pageId="5" pageNumber="5" type="discussion">
|
||
<paragraph blockId="5.[811,1449,1560,1987]" box="[811,1055,1560,1578]" pageId="5" pageNumber="5">
|
||
<heading bold="true" box="[811,1055,1560,1578]" fontSize="8" level="6" pageId="5" pageNumber="5" reason="6">
|
||
<emphasis bold="true" box="[811,1055,1560,1578]" pageId="5" pageNumber="5">Phylogenetic analysis</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="5.[811,1449,1560,1987]" pageId="5" pageNumber="5">
|
||
Our phylogenetic analysis focuses on non-monofenestratan pterosaurs and combines data from several independent published analyses
|
||
<superScript attach="left" box="[971,1040,1640,1654]" fontSize="6" pageId="5" pageNumber="5">
|
||
<bibRefCitation author="Wang, X. & Kellner, A. W. & Jiang, S. & Meng, X." box="[971,990,1640,1654]" journalOrPublisher="An. Acad. Bras. Cienc." pageId="5" pageNumber="5" pagination="793 - 812" part="81" refId="ref6565" refString="22. Wang, X., Kellner, A. W., Jiang, S., and Meng, X. (2009). An unusual longtailed pterosaur with elongated neck from western Liaoning of China. An. Acad. Bras. Cienc. 81, 793 - 812." title="An unusual longtailed pterosaur with elongated neck from western Liaoning of China" type="journal article" year="2009">22</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Andres, B. & Myers, T. S." box="[994,1013,1640,1654]" journalOrPublisher="Earth Environ. Sci. Trans. R. Soc. Edinb." pageId="5" pageNumber="5" pagination="383 - 398" part="103" refId="ref6740" refString="25. Andres, B., and Myers, T. S. (2012). Lone star pterosaurs. Earth Environ. Sci. Trans. R. Soc. Edinb. 103, 383 - 398." title="Lone star pterosaurs" type="journal article" year="2012">25</bibRefCitation>
|
||
–
|
||
<bibRefCitation author="Vidovic, S. U." box="[1021,1040,1640,1654]" pageId="5" pageNumber="5" refId="ref7244" refString="35. Vidovic, S. U. (2016). A discourse on pterosaur phylogeny (University of Portsmouth), PhD thesis." title="A discourse on pterosaur phylogeny (University of Portsmouth)" type="book" year="2016">35</bibRefCitation>
|
||
</superScript>
|
||
with new characters, while excluding taxa known only from highly immature specimens and characters that exhibit strong ontogenetic variation, resulting in a dataset of 58 taxa scored for 155 characters (Data S1B and S2).
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[811,1002,1762,1783]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="species" species="sgiathanach">
|
||
<emphasis box="[811,1002,1762,1783]" italics="true" pageId="5" pageNumber="5">Dearc sgiathanach</emphasis>
|
||
</taxonomicName>
|
||
is recovered within a large grade of nonmonofenestratans, including subclades centered on
|
||
<taxonomicName class="Reptilia" family="Rhamphorhynchidae" genus="Rhamphorhynchus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="5" pageNumber="5">Rhamphorhynchus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<emphasis box="[955,1112,1820,1841]" italics="true" pageId="5" pageNumber="5">Scaphognathus</emphasis>
|
||
(
|
||
<figureCitation box="[1126,1208,1820,1841]" captionStart="Figure 4" captionStartId="5.[142,202,1156,1175]" captionTargetBox="[143,1448,272,1113]" captionTargetId="figure-356@5.[142,1449,271,1116]" captionTargetPageId="5" captionText="Figure 4. Phylogenetic relationships of the Middle Jurassic pterosaur Dearc sgiathanach and wingspan estimates for Jurassic pterosaurs Strict consensus of most parsimonious trees from phylogenetic analysis,with silhouettes scaled to wingspan (Dearc = ca.2.0 m) (1), and skull reconstructions of key taxa: (A) Scaphognathus crassirostris (based on GPIB 1304), (B) Dorygnathus banthensis (based on SMNS 55886), (C) Rhamphorhynchus muensteri (based on NHMUK R 37002), (D) Angustinaripterus longicephalus (based on ZDM T8001), and (E) Dearc sgiathanach. Wingspan plot (2): estimated wingspan range for isolated pterosaur remains from the Taynton Limestone collection. See Data S1D for complete element list. (A) NHMUK PV R 36634, (B) GSM 113726, (C) OUM J28352,(D) NHMUK PV R 38016, (E) NHMUK PV R1362, (F) LL12158, (G) NHMUK PV R 40126,(H) NMS G.2021.6.1-4 (Dearc), (I) OUM J23046, (J) OUM J28273, (K) NHM UK PV R 40126 B, (L) OUM J28319, (M) OUM J28307, (N) OUM J28271, (O) OUM J28354, (P) LL12160, (Q) OUM J23047, and (R) MJM L K1995. See Figure S4 and Data S1 and S2." figureDoi="http://doi.org/10.5281/zenodo.6251726" httpUri="https://zenodo.org/record/6251726/files/figure.png" pageId="5" pageNumber="5">Figure 4</figureCitation>
|
||
).
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[1228,1288,1820,1841]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1228,1288,1820,1841]" italics="true" pageId="5" pageNumber="5">Dearc</emphasis>
|
||
</taxonomicName>
|
||
is in the former subclade, where it groups with the Chinese
|
||
<taxonomicName box="[1273,1449,1849,1870]" class="Reptilia" family="Rhamphorhynchidae" genus="Angustinaripterus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1273,1449,1849,1870]" italics="true" pageId="5" pageNumber="5">Angustinaripterus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName box="[863,983,1879,1900]" class="Reptilia" family="Rhamphorhynchidae" genus="Sericipterus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="5" pageNumber="5" phylum="Chordata" rank="genus">
|
||
<emphasis box="[863,983,1879,1900]" italics="true" pageId="5" pageNumber="5">Sericipterus</emphasis>
|
||
</taxonomicName>
|
||
, in the clade Angustinaripterini, diagnosed here for the first time by several features including a large antorbital fenestra, reclined quadrate, and proportionally elongate anterior cervicals (see above).
|
||
</paragraph>
|
||
<paragraph blockId="6.[159,798,274,1987]" box="[159,259,274,292]" pageId="6" pageNumber="6">
|
||
<heading bold="true" box="[159,259,274,292]" fontSize="8" level="6" pageId="6" pageNumber="6" reason="6">
|
||
<emphasis bold="true" box="[159,259,274,292]" pageId="6" pageNumber="6">Remarks</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph blockId="6.[159,798,274,1987]" pageId="6" pageNumber="6">
|
||
The holotype of
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[322,508,300,321]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="species" species="sgiathanach">
|
||
<emphasis box="[322,508,300,321]" italics="true" pageId="6" pageNumber="6">Dearc sgiathanach</emphasis>
|
||
</taxonomicName>
|
||
is a rare three-dimensionally preserved pterosaur from the Jurassic, which gives unique insight into the osteology, size, growth, and neuroanatomy of a basal non-monofenestratan. Its most remarkable attribute is its size: its wingspan was ca. 1.9–3.8 m, roughly the size of the largest flying birds today (e.g., wandering albatross), and it was not fully grown at death. Triassic and Jurassic pterosaurs have long been stereotyped as relatively small animals, constrained to wingspans of approximately 1.6–1.8 m or less over the first ca. 70 million years of their evolutionary history,
|
||
<bibRefCitation author="Benson, R. B. J. & Frigot, R. A. & Goswami, A. & Andres, B. & Butler, R. J." box="[707,716,559,573]" journalOrPublisher="Nat. Commun." pageId="6" pageNumber="6" pagination="3567" part="5" refId="ref5589" refString="5. Benson, R. B. J., Frigot, R. A., Goswami, A., Andres, B., and Butler, R. J. (2014). Competition and constraint drove Cope's rule in the evolution of giant flying reptiles. Nat. Commun. 5, 3567." title="Competition and constraint drove Cope's rule in the evolution of giant flying reptiles" type="journal article" year="2014">
|
||
<superScript attach="left" box="[707,716,559,573]" fontSize="6" pageId="6" pageNumber="6">5</superScript>
|
||
</bibRefCitation>
|
||
before becoming larger in the latest Jurassic or Early Cretaceous, culminating in airplane-sized giants like
|
||
<emphasis box="[594,742,622,643]" italics="true" pageId="6" pageNumber="6">Quetzalcoatlus</emphasis>
|
||
with 10-m wingspans.
|
||
<superScript attach="left" box="[335,344,646,660]" fontSize="6" pageId="6" pageNumber="6">5</superScript>
|
||
A few tantalizing specimens have hinted at larger Jurassic pterosaurs,
|
||
<superScript attach="left" box="[434,480,676,690]" fontSize="6" pageId="6" pageNumber="6">
|
||
<bibRefCitation author="Carpenter, K. & Unwin, D. & Cloward, K. & Miles, C." box="[434,453,676,690]" journalOrPublisher="Geol. Soc. Lond. Spec. Publ." pageId="6" pageNumber="6" pagination="45 - 54" part="217" refId="ref7270" refString="36. Carpenter, K., Unwin, D., Cloward, K., and Miles, C. (2003). A new scaphognathine pterosaur from the Upper Jurassic Morrison Formation of Wyoming, USA. Geol. Soc. Lond. Spec. Publ. 217, 45 - 54." title="A new scaphognathine pterosaur from the Upper Jurassic Morrison Formation of Wyoming, USA" type="journal article" year="2003">36</bibRefCitation>
|
||
–
|
||
<bibRefCitation author="Andres, B. & Clark, J. M. & Xing, X." box="[461,480,676,690]" journalOrPublisher="J. Vertebr. Paleontol." pageId="6" pageNumber="6" pagination="163 - 187" part="30" refId="ref7385" refString="38. Andres, B., Clark, J. M., and Xing, X. (2010). A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs. J. Vertebr. Paleontol. 30, 163 - 187." title="A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs" type="journal article" year="2010">38</bibRefCitation>
|
||
</superScript>
|
||
but these are often limited to one or a few bones, which make body size estimations difficult.
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[159,219,739,760]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[159,219,739,760]" italics="true" pageId="6" pageNumber="6">Dearc</emphasis>
|
||
</taxonomicName>
|
||
is the first Jurassic pterosaur whose wingspan can confidently be estimated at ca. 2.5 m or greater, based on a well-preserved, articulated skeleton.
|
||
<bibRefCitation author="Benson, R. B. J. & Frigot, R. A. & Goswami, A. & Andres, B. & Butler, R. J." box="[443,452,792,806]" journalOrPublisher="Nat. Commun." pageId="6" pageNumber="6" pagination="3567" part="5" refId="ref5589" refString="5. Benson, R. B. J., Frigot, R. A., Goswami, A., Andres, B., and Butler, R. J. (2014). Competition and constraint drove Cope's rule in the evolution of giant flying reptiles. Nat. Commun. 5, 3567." title="Competition and constraint drove Cope's rule in the evolution of giant flying reptiles" type="journal article" year="2014">
|
||
<superScript attach="left" box="[443,452,792,806]" fontSize="6" pageId="6" pageNumber="6">5</superScript>
|
||
</bibRefCitation>
|
||
Its closest relatives,
|
||
<taxonomicName class="Reptilia" family="Rhamphorhynchidae" genus="Angustinaripterus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis italics="true" pageId="6" pageNumber="6">Angustinaripterus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName box="[266,386,826,847]" class="Reptilia" family="Rhamphorhynchidae" genus="Sericipterus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[266,386,826,847]" italics="true" pageId="6" pageNumber="6">Sericipterus</emphasis>
|
||
</taxonomicName>
|
||
, are also sizeable for Jurassic species, with wingspans previously estimated at 1.6
|
||
<bibRefCitation author="Wellnhofer, P." box="[604,613,851,865]" journalOrPublisher="Salamander Books" pageId="6" pageNumber="6" refId="ref5475" refString="1. Wellnhofer, P. (1991). The Illustrated Encyclopedia of Pterosaurs (Salamander Books)." title="The Illustrated Encyclopedia of Pterosaurs" type="book" year="1991">
|
||
<superScript attach="none" box="[604,613,851,865]" fontSize="6" pageId="6" pageNumber="6">1</superScript>
|
||
</bibRefCitation>
|
||
–1.7
|
||
<bibRefCitation author="Andres, B. & Clark, J. M. & Xing, X." box="[656,675,851,865]" journalOrPublisher="J. Vertebr. Paleontol." pageId="6" pageNumber="6" pagination="163 - 187" part="30" refId="ref7385" refString="38. Andres, B., Clark, J. M., and Xing, X. (2010). A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs. J. Vertebr. Paleontol. 30, 163 - 187." title="A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs" type="journal article" year="2010">
|
||
<superScript attach="left" box="[656,675,851,865]" fontSize="6" pageId="6" pageNumber="6">38</superScript>
|
||
</bibRefCitation>
|
||
m extrapolated from patchy fossils. Our regression equations indicate larger wingspans for these taxa: ca. 2–3 m, which is still approximately 10% smaller than
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[435,495,943,964]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[435,495,943,964]" italics="true" pageId="6" pageNumber="6">Dearc</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[511,571,943,964]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[511,571,943,964]" italics="true" pageId="6" pageNumber="6">Dearc</emphasis>
|
||
</taxonomicName>
|
||
, therefore, anchors a clade of large, long-snouted Jurassic non-monofenestratans: Angustinaripterini.
|
||
<bibRefCitation author="Andres, B. & Clark, J. M. & Xing, X." box="[341,360,997,1011]" journalOrPublisher="J. Vertebr. Paleontol." pageId="6" pageNumber="6" pagination="163 - 187" part="30" refId="ref7385" refString="38. Andres, B., Clark, J. M., and Xing, X. (2010). A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs. J. Vertebr. Paleontol. 30, 163 - 187." title="A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs" type="journal article" year="2010">
|
||
<superScript attach="left" box="[341,360,997,1011]" fontSize="6" pageId="6" pageNumber="6">38</superScript>
|
||
</bibRefCitation>
|
||
</paragraph>
|
||
<paragraph blockId="6.[159,798,274,1987]" pageId="6" pageNumber="6">
|
||
The large size of
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[355,415,1031,1052]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[355,415,1031,1052]" italics="true" pageId="6" pageNumber="6">Dearc</emphasis>
|
||
</taxonomicName>
|
||
prompted us to re-examine fragmentary specimens from the Taynton Limestone,
|
||
<bibRefCitation author="O'Sullivan, M. & Martill, D. M." box="[619,638,1056,1070]" journalOrPublisher="Acta Palaeontol. Pol." pageId="6" pageNumber="6" pagination="617 - 644" part="63" refId="ref7441" refString="39. O'Sullivan, M., and Martill, D. M. (2018). Pterosauria of the Great Oolite Group (Bathonian, Middle Jurassic) of Oxfordshire and Gloucestershire, England. Acta Palaeontol. Pol. 63, 617 - 644." title="Pterosauria of the Great Oolite Group (Bathonian, Middle Jurassic) of Oxfordshire and Gloucestershire, England" type="journal article" year="2018">
|
||
<superScript attach="left" box="[619,638,1056,1070]" fontSize="6" pageId="6" pageNumber="6">39</superScript>
|
||
</bibRefCitation>
|
||
an English unit that formed at the same general time as the Scottish Middle Jurassic deposits, in or along the margins of the same seaway. We identified 17 specimens—all single bones—that yield wingspan estimates of over 1.7 m based on our predictor formulas (above), including several that may have had wingspans of over 3.0 m. The discovery of
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[457,517,1236,1257]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[457,517,1236,1257]" italics="true" pageId="6" pageNumber="6">Dearc</emphasis>
|
||
</taxonomicName>
|
||
, and our survey of Taynton specimens, reveals that Jurassic pterosaurs were capable of achieving considerably larger sizes than previously thought. Jurassic pterosaurs may still have been constrained in size— and certainly there is no evidence they approached the grandeur of giant Cretaceous pterodactyloids—but if so, that constraint was at a substantially greater wingspan than 1.6–1.8 m.
|
||
<bibRefCitation author="Benson, R. B. J. & Frigot, R. A. & Goswami, A. & Andres, B. & Butler, R. J." box="[720,729,1406,1420]" journalOrPublisher="Nat. Commun." pageId="6" pageNumber="6" pagination="3567" part="5" refId="ref5589" refString="5. Benson, R. B. J., Frigot, R. A., Goswami, A., Andres, B., and Butler, R. J. (2014). Competition and constraint drove Cope's rule in the evolution of giant flying reptiles. Nat. Commun. 5, 3567." title="Competition and constraint drove Cope's rule in the evolution of giant flying reptiles" type="journal article" year="2014">
|
||
<superScript attach="left" box="[720,729,1406,1420]" fontSize="6" pageId="6" pageNumber="6">5</superScript>
|
||
</bibRefCitation>
|
||
</paragraph>
|
||
<paragraph blockId="6.[159,798,274,1987]" pageId="6" pageNumber="6">
|
||
Trends in pterosaur size evolution, particularly the shift to increasingly larger species in the Cretaceous, have been interpreted in terms of two main hypotheses, which are not mutually exclusive: (1) advances in the pterodactyloid body plan allowed them to become larger and more efficient fliers than non-pterodactyloids, and (2) the diversification of birds (Avialae) may have driven latest Jurassic/Cretaceous pterosaurs into ever-larger size niches.
|
||
<bibRefCitation author="Benson, R. B. J. & Frigot, R. A. & Goswami, A. & Andres, B. & Butler, R. J." box="[284,293,1640,1654]" journalOrPublisher="Nat. Commun." pageId="6" pageNumber="6" pagination="3567" part="5" refId="ref5589" refString="5. Benson, R. B. J., Frigot, R. A., Goswami, A., Andres, B., and Butler, R. J. (2014). Competition and constraint drove Cope's rule in the evolution of giant flying reptiles. Nat. Commun. 5, 3567." title="Competition and constraint drove Cope's rule in the evolution of giant flying reptiles" type="journal article" year="2014">
|
||
<superScript attach="left" box="[284,293,1640,1654]" fontSize="6" pageId="6" pageNumber="6">5</superScript>
|
||
</bibRefCitation>
|
||
Our identification of
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[532,592,1645,1666]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[532,592,1645,1666]" italics="true" pageId="6" pageNumber="6">Dearc</emphasis>
|
||
</taxonomicName>
|
||
demonstrates that non-pterodactyloids were able to grow to larger sizes by the Middle Jurassic, with some evidence for large pterosaurs back to the Early Jurassic,
|
||
<bibRefCitation author="O'Sullivan, M. & Martill, D. M. & Groocock, D." box="[369,388,1728,1742]" journalOrPublisher="Proc. Geol. Assoc." pageId="6" pageNumber="6" pagination="973 - 981" part="124" refId="ref7327" refString="37. O'Sullivan, M., Martill, D. M., and Groocock, D. (2013). A pterosaur humerus and scapulocoracoid from the Jurassic Whitby Mudstone Formation, and the evolution of large body size in early pterosaurs. Proc. Geol. Assoc. 124, 973 - 981." title="A pterosaur humerus and scapulocoracoid from the Jurassic Whitby Mudstone Formation, and the evolution of large body size in early pterosaurs" type="journal article" year="2013">
|
||
<superScript attach="left" box="[369,388,1728,1742]" fontSize="6" pageId="6" pageNumber="6">37</superScript>
|
||
</bibRefCitation>
|
||
tens of millions of years before birds underwent their adaptive Cretaceous radiation.
|
||
<superScript attach="left" box="[609,651,1757,1771]" fontSize="6" pageId="6" pageNumber="6">
|
||
<bibRefCitation author="Brusatte, S. L. & O'Connor, J. K. & Jarvis, E. D." box="[609,628,1757,1771]" journalOrPublisher="Curr. Biol." pageId="6" pageNumber="6" pagination="888 - R 898" part="25" refId="ref7489" refString="40. Brusatte, S. L., O'Connor, J. K., and Jarvis, E. D. (2015). The origin and diversification of birds. Curr. Biol. 25, R 888 - R 898." title="The origin and diversification of birds" type="journal article" year="2015">40</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Benson, R. B. J. & Choiniere, J. N." box="[632,651,1757,1771]" journalOrPublisher="Proc. Biol. Sci." pageId="6" pageNumber="6" pagination="20131780" part="280" refId="ref7535" refString="41. Benson, R. B. J., and Choiniere, J. N. (2013). Rates of dinosaur limb evolution provide evidence for exceptional radiation in Mesozoic birds. Proc. Biol. Sci. 280, 20131780." title="Rates of dinosaur limb evolution provide evidence for exceptional radiation in Mesozoic birds" type="journal article" year="2013">41</bibRefCitation>
|
||
</superScript>
|
||
These size increases seemingly occurred too early for avialans, which are first definitively known from the Late Jurassic, to have been a direct cause. Alternatively, if there was pressure on pterosaurs to become larger, it may have started deep in the Jurassic and involved competition with unrecognized early avialans or other animals,
|
||
<bibRefCitation author="Brusatte, S. L. & O'Connor, J. K. & Jarvis, E. D." box="[243,262,1932,1946]" journalOrPublisher="Curr. Biol." pageId="6" pageNumber="6" pagination="888 - R 898" part="25" refId="ref7489" refString="40. Brusatte, S. L., O'Connor, J. K., and Jarvis, E. D. (2015). The origin and diversification of birds. Curr. Biol. 25, R 888 - R 898." title="The origin and diversification of birds" type="journal article" year="2015">
|
||
<superScript attach="left" box="[243,262,1932,1946]" fontSize="6" pageId="6" pageNumber="6">40</superScript>
|
||
</bibRefCitation>
|
||
like non-avialan feathered dinosaurs or other pterosaurs.
|
||
</paragraph>
|
||
<paragraph blockId="6.[829,1468,271,1315]" pageId="6" pageNumber="6">
|
||
Not only is
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[963,1023,271,292]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[963,1023,271,292]" italics="true" pageId="6" pageNumber="6">Dearc</emphasis>
|
||
</taxonomicName>
|
||
large, but it and its closest angustinaripterin relatives possess derived characters considered keystones of the pterodactyloid skull ‘‘module,’’ notably an elongate skull and inclined quadrate. Previous work has argued that the transition between non-pterodactyloids and pterodactyloids involved a nearly perfect modular shift, in which features of the skull changed together in a tightly integrated unit, followed by the body, limbs, and tail.
|
||
<bibRefCitation author="Lu, J. & Unwin, D. M. & Jin, X. & Liu, Y. & Ji, Q." box="[1041,1050,471,485]" journalOrPublisher="Proc. Biol. Sci." pageId="6" pageNumber="6" pagination="383 - 389" part="277" refId="ref5650" refString="6. Lu, J., Unwin, D. M., Jin, X., Liu, Y., and Ji, Q. (2010). Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proc. Biol. Sci. 277, 383 - 389." title="Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull" type="journal article" year="2010">
|
||
<superScript attach="left" box="[1041,1050,471,485]" fontSize="6" pageId="6" pageNumber="6">6</superScript>
|
||
</bibRefCitation>
|
||
While we do not dispute the overarching modularity of the pterosaur skeleton,
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[1213,1273,505,526]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1213,1273,505,526]" italics="true" pageId="6" pageNumber="6">Dearc</emphasis>
|
||
</taxonomicName>
|
||
hints that some of the trademark ‘‘pterodactyloid’’ features convergently evolved in other groups,
|
||
<bibRefCitation author="Sullivan, C. & Yuan, W. & Hone, D. W. E. & Wang, Y. & Xu, X. & Zhang, F." box="[988,1007,559,573]" journalOrPublisher="J. Vertebr. Paleontol." pageId="6" pageNumber="6" pagination="243 - 280" part="34" refId="ref7581" refString="42. Sullivan, C., Yuan, W., Hone, D. W. E., Wang, Y., Xu, X., and Zhang, F. (2014). The vertebrates of the Jurassic Daohugou Biota of northeastern China. J. Vertebr. Paleontol. 34, 243 - 280." title="The vertebrates of the Jurassic Daohugou Biota of northeastern China" type="journal article" year="2014">
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||
<superScript attach="left" box="[988,1007,559,573]" fontSize="6" pageId="6" pageNumber="6">42</superScript>
|
||
</bibRefCitation>
|
||
perhaps due to feeding ecology or other factors. Yet the endocranial anatomy of
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[1194,1254,593,614]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1194,1254,593,614]" italics="true" pageId="6" pageNumber="6">Dearc</emphasis>
|
||
</taxonomicName>
|
||
is distinctly primitive, as it has the unflexed brain and smaller flocculus of basal pterosaurs
|
||
<superScript attach="left" box="[884,917,646,660]" fontSize="6" pageId="6" pageNumber="6">
|
||
<bibRefCitation author="Witmer, L. M. & Chatterjee, S. & Franzosa, J. & Rowe, T." box="[884,893,646,660]" journalOrPublisher="Nature" pageId="6" pageNumber="6" pagination="950 - 953" part="425" refId="ref5708" refString="7. Witmer, L. M., Chatterjee, S., Franzosa, J., and Rowe, T. (2003). Neuroanatomy of flying reptiles and implications for flight, posture and behaviour. Nature 425, 950 - 953." title="Neuroanatomy of flying reptiles and implications for flight, posture and behaviour" type="journal article" year="2003">7</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Codorniu, L. & Paulina Carabajal, A. & Pol, D. & Unwin, D. & Rauhut, O. W." box="[898,917,646,660]" journalOrPublisher="PeerJ" pageId="6" pageNumber="6" pagination="2311" part="4" refId="ref6687" refString="24. Codorniu, L., Paulina Carabajal, A., Pol, D., Unwin, D., and Rauhut, O. W. (2016). A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium. PeerJ 4, e 2311." title="A Jurassic pterosaur from Patagonia and the origin of the pterodactyloid neurocranium" type="journal article" year="2016">24</bibRefCitation>
|
||
</superScript>
|
||
and not the transformed brain of pterodactyloids. Thus, it seems, non-pterodactyloids had a similar neuroanatomy regardless of body size.
|
||
</paragraph>
|
||
<paragraph blockId="6.[829,1468,271,1315]" pageId="6" pageNumber="6">
|
||
Our recognition of large Middle Jurassic pterosaurs exposes a taphonomic bias, to add to the already notoriously problematic record of pterosaurs.
|
||
<superScript attach="left" box="[1045,1088,792,806]" fontSize="6" pageId="6" pageNumber="6">
|
||
<bibRefCitation author="Butler, R. J. & Benson, R. B. & Barrett, P. M." box="[1045,1064,792,806]" journalOrPublisher="Palaeogeogr. Palaeoclimatol. Palaeoecol." pageId="6" pageNumber="6" pagination="78 - 87" part="372" refId="ref7644" refString="43. Butler, R. J., Benson, R. B., and Barrett, P. M. (2013). Pterosaur diversity: untangling the influence of sampling biases, Lagerstatten ¨, and genuine biodiversity signals. Palaeogeogr. Palaeoclimatol. Palaeoecol. 372, 78 - 87." title="Pterosaur diversity: untangling the influence of sampling biases, Lagerstatten ¨, and genuine biodiversity signals" type="journal article" year="2013">43</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Dean, C. D. & Mannion, P. D. & Butler, R. J." box="[1069,1088,792,806]" journalOrPublisher="Palaeontology" pageId="6" pageNumber="6" pagination="225 - 247" part="59" refId="ref7700" refString="44. Dean, C. D., Mannion, P. D., and Butler, R. J. (2016). Preservational bias controls the fossil record of pterosaurs. Palaeontology 59, 225 - 247." title="Preservational bias controls the fossil record of pterosaurs" type="journal article" year="2016">44</bibRefCitation>
|
||
</superScript>
|
||
There were fairly large pterosaurs in the Jurassic (wingspans> 1.8 m), but reasonably complete adult or near-mature skeletons are mostly lacking, for reasons unclear and worthy of further study. The Middle Jurassic age of
|
||
<taxonomicName authorityName="Jagielska & O’Sullivan & Funston & Butler & Challands & Clark & Fraser & Penny & Ross & Wilkinson & Brusatte" authorityYear="2022" box="[1407,1467,885,906]" class="Reptilia" family="Rhamphorhynchidae" genus="Dearc" higherTaxonomySource="GBIF" kingdom="Animalia" order="Pterosauria" pageId="6" pageNumber="6" phylum="Chordata" rank="genus">
|
||
<emphasis box="[1407,1467,885,906]" italics="true" pageId="6" pageNumber="6">Dearc</emphasis>
|
||
</taxonomicName>
|
||
adds to increasing evidence that this interval—once a frustrating gap in the pterosaur record—was in fact a dynamic time of diversification, in which a variety of basal taxa and early monofenestratan lineages
|
||
<superScript attach="left" box="[978,1012,997,1011]" fontSize="6" pageId="6" pageNumber="6">
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||
<bibRefCitation author="Lu, J. & Unwin, D. M. & Jin, X. & Liu, Y. & Ji, Q." box="[978,987,997,1011]" journalOrPublisher="Proc. Biol. Sci." pageId="6" pageNumber="6" pagination="383 - 389" part="277" refId="ref5650" refString="6. Lu, J., Unwin, D. M., Jin, X., Liu, Y., and Ji, Q. (2010). Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull. Proc. Biol. Sci. 277, 383 - 389." title="Evidence for modular evolution in a long-tailed pterosaur with a pterodactyloid skull" type="journal article" year="2010">6</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="O'Sullivan, M. & Martill, D. M." box="[993,1012,997,1011]" journalOrPublisher="Acta Palaeontol. Pol." pageId="6" pageNumber="6" pagination="617 - 644" part="63" refId="ref7441" refString="39. O'Sullivan, M., and Martill, D. M. (2018). Pterosauria of the Great Oolite Group (Bathonian, Middle Jurassic) of Oxfordshire and Gloucestershire, England. Acta Palaeontol. Pol. 63, 617 - 644." title="Pterosauria of the Great Oolite Group (Bathonian, Middle Jurassic) of Oxfordshire and Gloucestershire, England" type="journal article" year="2018">39</bibRefCitation>
|
||
</superScript>
|
||
coexisted and occupied a range of environments, from open marine to lagoonal, nearshore to desert, around the world. We can now add larger taxa, with derived pterodactyloid-type skull characters, to that roster. As with dinosaurs
|
||
<bibRefCitation author="Benson, R. B. J." box="[884,903,1114,1128]" journalOrPublisher="Annu. Rev. Earth Planet. Sci." pageId="6" pageNumber="6" pagination="379 - 408" part="49" refId="ref7743" refString="45. Benson, R. B. J. (2018). Dinosaur macroevolution and macroecology. Annu. Rev. Earth Planet. Sci. 49, 379 - 408." title="Dinosaur macroevolution and macroecology" type="journal article" year="2018">
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||
<superScript attach="left" box="[884,903,1114,1128]" fontSize="6" pageId="6" pageNumber="6">45</superScript>
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||
</bibRefCitation>
|
||
and mammals,
|
||
<bibRefCitation author="Close, R. A. & Friedman, M. & Lloyd, G. T. & Benson, R. B." box="[1056,1075,1114,1128]" journalOrPublisher="Curr. Biol." pageId="6" pageNumber="6" pagination="2137 - 2142" part="25" refId="ref7777" refString="46. Close, R. A., Friedman, M., Lloyd, G. T., and Benson, R. B. (2015). Evidence for a mid-Jurassic adaptive radiation in mammals. Curr. Biol. 25, 2137 - 2142." title="Evidence for a mid-Jurassic adaptive radiation in mammals" type="journal article" year="2015">
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<superScript attach="left" box="[1056,1075,1114,1128]" fontSize="6" pageId="6" pageNumber="6">46</superScript>
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||
</bibRefCitation>
|
||
the Middle Jurassic was likely a vibrant time in pterosaur history, not a static and archaic prelude to a Cretaceous explosion of larger,
|
||
<bibRefCitation author="Benson, R. B. J. & Frigot, R. A. & Goswami, A. & Andres, B. & Butler, R. J." box="[1138,1147,1172,1186]" journalOrPublisher="Nat. Commun." pageId="6" pageNumber="6" pagination="3567" part="5" refId="ref5589" refString="5. Benson, R. B. J., Frigot, R. A., Goswami, A., Andres, B., and Butler, R. J. (2014). Competition and constraint drove Cope's rule in the evolution of giant flying reptiles. Nat. Commun. 5, 3567." title="Competition and constraint drove Cope's rule in the evolution of giant flying reptiles" type="journal article" year="2014">
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||
<superScript attach="left" box="[1138,1147,1172,1186]" fontSize="6" pageId="6" pageNumber="6">5</superScript>
|
||
</bibRefCitation>
|
||
more disparate,
|
||
<bibRefCitation author="Prentice, K. C. & Ruta, M. & Benton, M. J." box="[1308,1327,1172,1186]" journalOrPublisher="J. Syst. Palaeontology" pageId="6" pageNumber="6" pagination="337 - 353" part="9" refId="ref7828" refString="47. Prentice, K. C., Ruta, M., and Benton, M. J. (2011). Evolution of morphological disparity in pterosaurs. J. Syst. Palaeontology 9, 337 - 353." title="Evolution of morphological disparity in pterosaurs" type="journal article" year="2011">
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<superScript attach="left" box="[1308,1327,1172,1186]" fontSize="6" pageId="6" pageNumber="6">47</superScript>
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||
</bibRefCitation>
|
||
more efficient fliers.
|
||
<bibRefCitation author="Venditti, C. & Baker, J. & Benton, M. J. & Meade, A. & Humphries, S." box="[883,902,1202,1216]" journalOrPublisher="Nature" pageId="6" pageNumber="6" pagination="83 - 86" part="587" refId="ref6455" refString="20. Venditti, C., Baker, J., Benton, M. J., Meade, A., and Humphries, S. (2020). 150 million years of sustained increase in pterosaur flight efficiency. Nature 587, 83 - 86." title="150 million years of sustained increase in pterosaur flight efficiency" type="journal article" year="2020">
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<superScript attach="left" box="[883,902,1202,1216]" fontSize="6" pageId="6" pageNumber="6">20</superScript>
|
||
</bibRefCitation>
|
||
With the dawn of the Cretaceous, however, the diverse non-monofenestratans disappeared, including larger ones with pterodactyloid convergences. This mystery remains to be solved.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |