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<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:ali="http://www.niso.org/schemas/ali/1.0/" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" article-type="research-article" dtd-version="1.2" xml:lang="en"><front><journal-meta><journal-id journal-id-type="publisher-id">RSOS</journal-id><journal-id journal-id-type="hwp">royopensci</journal-id><journal-title-group><journal-title>Royal Society Open Science</journal-title><abbrev-journal-title>R. Soc. Open Sci.</abbrev-journal-title></journal-title-group><issn publication-format="electronic">2054-5703</issn><publisher><publisher-name>The Royal Society</publisher-name></publisher></journal-meta><article-meta><article-id pub-id-type="publisher-id">rsos241090</article-id><article-id pub-id-type="doi">10.1098/rsos.241090</article-id><article-categories><subj-group subj-group-type="discipline-codes"><compound-subject><compound-subject-part content-type="code">1001</compound-subject-part></compound-subject><compound-subject><compound-subject-part content-type="code">1001</compound-subject-part></compound-subject><compound-subject><compound-subject-part content-type="code">1001</compound-subject-part></compound-subject></subj-group><subj-group subj-group-type="subject-codes"><compound-subject><compound-subject-part content-type="code">197</compound-subject-part></compound-subject><compound-subject><compound-subject-part content-type="code">60</compound-subject-part></compound-subject><compound-subject><compound-subject-part content-type="code">15</compound-subject-part></compound-subject></subj-group><subj-group subj-group-type="heading"><subject>Science, society and policy</subject></subj-group><subj-group subj-group-type="type-of-publication"><subject>Research articles</subject></subj-group></article-categories><title-group><article-title>Hiding in plain sight: the biomolecular identification of pinniped use in medieval manuscripts</article-title><alt-title alt-title-type="short">Hiding in plain sight: the biomolecular identification of pinniped use in medieval manuscripts</alt-title></title-group><contrib-group><contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0001-8420-792X</contrib-id><string-name name-style="western"><given-names>Élodie</given-names> <surname>Lévêque</surname></string-name><xref ref-type="aff" rid="aff1"><sup>1</sup></xref><x xml:space="preserve"><sup>, </sup></x><xref ref-type="aff" rid="aff2"><sup>2</sup></xref><x xml:space="preserve"><sup>, </sup></x><xref ref-type="aff" rid="aff3"><sup>3</sup></xref><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing – original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing – original draft</role><email>elodie.leveque@palaeome.org</email><x xml:space="preserve">, </x></contrib><contrib contrib-type="author"><string-name name-style="western"><given-names>Matthew D.</given-names> <surname>Teasdale</surname></string-name><xref ref-type="aff" rid="aff4"><sup>4</sup></xref><x xml:space="preserve"><sup>, </sup></x><xref ref-type="aff" rid="aff5"><sup>5</sup></xref><x xml:space="preserve"><sup>, </sup></x><xref ref-type="aff" rid="aff6"><sup>6</sup></xref><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing – original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing – original draft</role><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Data curation" vocab-term-identifier="https://credit.niso.org/contributor-roles/data-curation/">Data curation</role><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Methodology" vocab-term-identifier="https://credit.niso.org/contributor-roles/methodology/">Methodology</role><email>matthew.teasdale@palaeome.org</email><x xml:space="preserve">, </x></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0002-8991-2318</contrib-id><string-name name-style="western"><given-names>Sarah</given-names> <surname>Fiddyment</surname></string-name><xref ref-type="aff" rid="aff4"><sup>4</sup></xref><x xml:space="preserve"><sup>, </sup></x><xref ref-type="aff" rid="aff6"><sup>6</sup></xref><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role><email>sarah.fiddyment@palaeome.org</email><x xml:space="preserve">, </x></contrib><contrib contrib-type="author"><string-name name-style="western"><given-names>Maiken Hemme</given-names> <surname>Bro-Jørgensen</surname></string-name><xref ref-type="aff" rid="aff3"><sup>3</sup></xref><x xml:space="preserve"><sup>, </sup></x><xref ref-type="aff" rid="aff7"><sup>7</sup></xref><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing – review and editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing – review and editing</role><email>MBro-Jorgensen@ucc.ie</email><x xml:space="preserve">, </x></contrib><contrib contrib-type="author"><string-name name-style="western"><given-names>Luke</given-names> <surname>Spindler</surname></string-name><xref ref-type="aff" rid="aff6"><sup>6</sup></xref><x xml:space="preserve"><sup>, </sup></x><xref ref-type="aff" rid="aff8"><sup>8</sup></xref><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role><email>l.spindler@bradford.ac.uk</email><x xml:space="preserve">, </x></contrib><contrib contrib-type="author"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0001-8086-4420</contrib-id><string-name name-style="western"><given-names>Ruairidh</given-names> <surname>Macleod</surname></string-name><xref ref-type="aff" rid="aff3"><sup>3</sup></xref><x xml:space="preserve"><sup>, </sup></x><xref ref-type="aff" rid="aff4"><sup>4</sup></xref><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role><email>ram88@cam.ac.uk; r@palaeome.org</email><x xml:space="preserve">, </x></contrib><contrib contrib-type="author"><string-name name-style="western"><given-names>François</given-names> <surname>Bougard</surname></string-name><xref ref-type="aff" rid="aff2"><sup>2</sup></xref><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Validation" vocab-term-identifier="https://credit.niso.org/contributor-roles/validation/">Validation</role><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Supervision" vocab-term-identifier="https://credit.niso.org/contributor-roles/supervision/">Supervision</role><email>f.bougard@irht.cnrs.fr</email><x xml:space="preserve">, </x></contrib><contrib contrib-type="author" equal-contrib="yes"><string-name name-style="western"><given-names>Morten</given-names> <surname>Tange Olsen</surname></string-name><xref ref-type="aff" rid="aff3"><sup>3</sup></xref><xref ref-type="author-notes" rid="equal-contrib1"><sup>†</sup></xref><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Supervision" vocab-term-identifier="https://credit.niso.org/contributor-roles/supervision/">Supervision</role><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing – review and editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing – review and editing</role><email>morten.olsen@sund.ku.dk</email><x xml:space="preserve"> and </x></contrib><contrib corresp="yes" contrib-type="author" equal-contrib="yes"><contrib-id contrib-id-type="orcid" authenticated="true">https://orcid.org/0000-0003-4226-5501</contrib-id><string-name name-style="western"><given-names>Matthew</given-names> <surname>Collins</surname></string-name><xref ref-type="aff" rid="aff3"><sup>3</sup></xref><x xml:space="preserve"><sup>, </sup></x><xref ref-type="aff" rid="aff4"><sup>4</sup></xref><xref ref-type="author-notes" rid="equal-contrib1"><sup>†</sup></xref><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Supervision" vocab-term-identifier="https://credit.niso.org/contributor-roles/supervision/">Supervision</role><role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Funding acquisition" vocab-term-identifier="https://credit.niso.org/contributor-roles/funding-acquisition/">Funding acquisition</role><email>matthew@palaeome.org</email></contrib><aff id="aff1"><label><sup>1</sup></label><institution-wrap><institution>HiCSA, Université Paris 1 Panthéon-Sorbonne</institution></institution-wrap>, <city>Paris</city>, <country>France</country></aff><aff id="aff2"><label><sup>2</sup></label><institution-wrap><institution>Institut de Recherche et d’Histoire des Textes</institution></institution-wrap>, <city>Paris, Île-de-France</city>, <country>France</country></aff><aff id="aff3"><label><sup>3</sup></label><institution-wrap><institution>Evolutionary Genomics, Globe Institute, University of Copenhagen</institution></institution-wrap>, <city>Copenhagen</city>, <country>Denmark</country></aff><aff id="aff4"><label><sup>4</sup></label><institution-wrap><institution>McDonald Institute for Archaeological Research, University of Cambridge</institution></institution-wrap>, <city>Cambridge</city>, <country>UK</country></aff><aff id="aff5"><label><sup>5</sup></label><institution-wrap><institution>Bioinformatics Support Unit, Faculty of Medical Sciences, Newcastle University</institution></institution-wrap>, <city>Newcastle Upon Tyne</city>, <country>UK</country></aff><aff id="aff6"><label><sup>6</sup></label><institution-wrap><institution>BioArCh, Department of Archaeology, University of York</institution></institution-wrap>, <city>York</city>, <country>UK</country></aff><aff id="aff7"><label><sup>7</sup></label><institution-wrap><institution>School of Biochemistry and Cell Biology, University College Cork</institution></institution-wrap>, <city>Cork</city>, <country>Ireland</country></aff><aff id="aff8"><label><sup>8</sup></label><institution-wrap><institution>School of Archaeological and Forensic Sciences, University of Bradford</institution></institution-wrap>, <city>Bradford</city>, <country>UK</country></aff></contrib-group><author-notes><fn fn-type="other"><p>Electronic supplementary material is available online at <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.6084/m9.figshare.c.7757095">https://doi.org/10.6084/m9.figshare.c.7757095</ext-link>.</p></fn><fn fn-type="other" id="equal-contrib1"><label><sup>†</sup></label><p>These authors contributed equally to the study.</p></fn></author-notes><pub-date date-type="collection" publication-format="electronic"><month>04</month><year>2025</year></pub-date><pub-date date-type="pub" publication-format="electronic"><day>09</day><month>04</month><year>2025</year><string-date>April 9, 2025</string-date></pub-date><volume>12</volume><issue>4</issue><elocation-id>241090</elocation-id><history><date date-type="received"><day>08</day><month>07</month><year>2024</year><string-date>July 8, 2024</string-date></date><date date-type="rev-recd"><day>10</day><month>02</month><year>2025</year><string-date>February 10, 2025</string-date></date><date date-type="accepted"><day>10</day><month>03</month><year>2025</year><string-date>March 10, 2025</string-date></date></history><permissions><copyright-statement>© 2025 The Author(s).</copyright-statement><copyright-year>2025</copyright-year><license license-type="open-access" xlink:href="http://creativecommons.org/licenses/by/4.0/"><ali:license_ref specific-use="vor">http://creativecommons.org/licenses/by/4.0/</ali:license_ref><ali:license_ref specific-use="tdm">http://creativecommons.org/licenses/by/4.0/</ali:license_ref><ali:license_ref specific-use="version-of-record">http://creativecommons.org/licenses/by/4.0/</ali:license_ref><ali:license_ref specific-use="text-data-mining">http://creativecommons.org/licenses/by/4.0/</ali:license_ref><license-p>Published by the Royal Society under the terms of the Creative Commons Attribution License <ext-link ext-link-type="uri" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">http://creativecommons.org/licenses/by/4.0/</ext-link>, which permits unrestricted use, provided the original author and source are credited.</license-p></license></permissions><self-uri content-type="pdf" xlink:href="rsos.241090.pdf" /><related-object id="ss1" source-id="https://www.webofscience.com/api/gateway/wos/peer-review/10.1098/rsos.241090" source-id-type="url" source-type="peer-review-report" /><abstract><p>The survival of medieval manuscripts in their original bindings remains a rare occurrence. Taking advantage of the diversity of bindings in Cistercian libraries such as Clairvaux and its daughter abbeys during the twelfth and thirteenth centuries, this study focuses on the biocodicological analysis of medieval manuscript bindings, with particular emphasis on the use of sealskins. Using innovative methods such as electrostatic zooarchaeology by mass spectrometry (eZooMS) and ancient DNA (aDNA) analysis, this research identifies the animal species and origin of the leather used in these bindings as predominantly pinniped (seal) species. In particular, the collagen-based eZooMS technique facilitated the classification of seven chemises into the pinniped clade, although species identification remained elusive, except in one additional case where a bearded seal (<italic toggle="yes">Erignathus barbatus</italic>) was definitively identified. aDNA analysis was instrumental in verifying the origin of the sealskins, with four samples identified as harbour seals and a single sample as a harp seal and sourced to (contemporary) populations in Scandinavia, Scotland and Iceland or Greenland. This geographical inference supports the notion of a robust medieval trade network that went well beyond local sourcing, linking the Cistercians to wider economic circuits that included fur trade with the Norse. The study, therefore, highlights the use of an unexpected skin (seal) from an unexpected source (the northwestern Atlantic). The widespread use of sealskins in Cistercian libraries such as Clairvaux and its daughter abbeys during the twelfth and thirteenth centuries hints at broader trade networks that brought, for example, walrus ivory from the far north into continental Europe. This integration of the biological sciences into the study of historical manuscripts not only provides a clearer picture of the material culture of medieval Europe, but also illustrates the extensive trade networks that Cistercian monasteries were part of, challenging previous assumptions about local resource use in manuscript production.</p></abstract><kwd-group><kwd>medieval manuscripts</kwd><x xml:space="preserve">; </x><kwd>sealskin</kwd><x xml:space="preserve">; </x><kwd>pinniped</kwd><x xml:space="preserve">; </x><kwd>Romanesque bindings</kwd><x xml:space="preserve">; </x><kwd>Norse trade networks</kwd><x xml:space="preserve">; </x><kwd>biocodicology</kwd></kwd-group><funding-group specific-use="FundRef"><award-group award-type="grant" id="award-1"><funding-source><institution-wrap><institution>Richard Lounsbury Foundation</institution></institution-wrap></funding-source></award-group><award-group award-type="grant" id="award-2"><funding-source><institution-wrap><institution>EU Framework Programme</institution></institution-wrap></funding-source></award-group><award-group award-type="grant" id="award-3"><funding-source><institution-wrap><institution>ERC</institution></institution-wrap></funding-source></award-group></funding-group></article-meta></front></article>
