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<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="doi">10.1098/rsos.221023</article-id>
<article-id pub-id-type="publisher-id">rsos221023</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>
</subj-group>
<subj-group subj-group-type="subject-codes"><compound-subject><compound-subject-part content-type="code">60</compound-subject-part></compound-subject><compound-subject><compound-subject-part content-type="code">70</compound-subject-part></compound-subject><compound-subject><compound-subject-part content-type="code">202</compound-subject-part></compound-subject>
</subj-group>
<subj-group subj-group-type="heading">
<subject>Organismal and evolutionary biology</subject>
</subj-group>
<subj-group subj-group-type="type-of-publication">
<subject>Research articles</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Eggshell composition and surface properties of avian brood-parasitic species compared with non-parasitic species</article-title>
<alt-title alt-title-type="short">Eggshell composition and surface properties of avian brood-parasitic species compared to non-parasitic species</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes"><contrib-id contrib-id-type="orcid" authenticated="true">http://orcid.org/0000-0002-8763-2291</contrib-id><string-name name-style="western"><given-names>Stephanie C.</given-names> <surname>McClelland</surname></string-name><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Conceptualization" vocab-term-identifier="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Data curation" vocab-term-identifier="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="http://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Investigation" vocab-term-identifier="http://credit.niso.org/contributor-roles/investigation/">Investigation</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Methodology" vocab-term-identifier="http://credit.niso.org/contributor-roles/methodology/">Methodology</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Project administration" vocab-term-identifier="http://credit.niso.org/contributor-roles/project-administration/">Project administration</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Validation" vocab-term-identifier="http://credit.niso.org/contributor-roles/validation/">Validation</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Visualization" vocab-term-identifier="http://credit.niso.org/contributor-roles/visualization/">Visualization</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Writing – original draft" vocab-term-identifier="http://credit.niso.org/contributor-roles/writing-–-original-draft/">Writing – original draft</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Writing – review &amp; editing" vocab-term-identifier="http://credit.niso.org/contributor-roles/writing-–-review-editing/">Writing – review &amp; editing</role><email>stephanie.mcclelland.2018@live.rhul.ac.uk</email>
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<contrib contrib-type="author"><string-name name-style="western"><given-names>Claire N.</given-names> <surname>Spottiswoode</surname></string-name><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Conceptualization" vocab-term-identifier="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Investigation" vocab-term-identifier="http://credit.niso.org/contributor-roles/investigation/">Investigation</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Methodology" vocab-term-identifier="http://credit.niso.org/contributor-roles/methodology/">Methodology</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Resources" vocab-term-identifier="http://credit.niso.org/contributor-roles/resources/">Resources</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Supervision" vocab-term-identifier="http://credit.niso.org/contributor-roles/supervision/">Supervision</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Validation" vocab-term-identifier="http://credit.niso.org/contributor-roles/validation/">Validation</role><role vocab="credit" vocab-identifier="http://credit.niso.org/" vocab-term="Writing – review &amp; editing" vocab-term-identifier="http://credit.niso.org/contributor-roles/writing-–-review-editing/">Writing – review &amp; editing</role>
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<aff id="af1"><label><sup>1</sup></label><addr-line>Department of Biological Sciences, School of Life and Environmental Sciences, <institution>Royal Holloway University of London</institution>, Egham, Surrey TW20 0EX, <country>UK</country></addr-line></aff>
<aff id="af2"><label><sup>2</sup></label><addr-line>School of Engineering and Innovation, <institution>Open University</institution>, Milton Keynes MK7 6AA, <country>UK</country></addr-line></aff>
<aff id="af3"><label><sup>3</sup></label><addr-line>Department of Zoology, <institution>University of Cambridge</institution>, Downing Street, Cambridge CB2 3EJ, <country>UK</country></addr-line></aff>
<aff id="af4"><label><sup>4</sup></label><addr-line>Cambridge Institute of Therapeutic Immunology and Infectious Disease (CITIID), Jeffrey Cheah Biomedical Centre, Cambridge Biomedical Campus, <institution>Cambridge University</institution>, Cambridge CB2 0AW, <country>UK</country></addr-line></aff>
<aff id="af5"><label><sup>5</sup></label><addr-line>FitzPatrick Institute of African Ornithology, DST-NRF Centre of Excellence, <institution>University of Cape Town</institution>, Rondebosch 7701, Cape Town, <country>South Africa</country></addr-line></aff>
<aff id="af6"><label><sup>6</sup></label><addr-line><institution>The Natural History Museum</institution>, Tring, Herts HP23 6AP, <country>UK</country></addr-line></aff>
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<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.6653130">https://doi.org/10.6084/m9.figshare.c.6653130</ext-link>.</p></fn>
</author-notes>
<pub-date publication-format="electronic" date-type="pub"><day>24</day><month>5</month><year>2023</year><string-date>May 24, 2023</string-date></pub-date>
<pub-date publication-format="electronic" date-type="collection"><month>5</month><year>2023</year></pub-date>
<volume>10</volume>
<issue>5</issue><elocation-id>221023</elocation-id>
<history>
<date date-type="received">
<day>16</day>
<month>8</month>
<year>2022</year><string-date>August 16, 2022</string-date></date>
<date date-type="accepted">
<day>5</day>
<month>5</month>
<year>2023</year><string-date>May 5, 2023</string-date></date>
</history><permissions>
<copyright-statement>© 2023 The Authors.</copyright-statement>
<copyright-year>2023</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>
<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/">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>
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<abstract>
<p>The eggs of avian obligate brood-parasitic species have multiple adaptations to deceive hosts and optimize development in host nests. While the structure and composition of the eggshell in all birds is essential for embryo growth and protection from external threats, parasitic eggs may face specific challenges such as high microbial loads, rapid laying and ejection by the host parents. We set out to assess whether eggshells of avian brood-parasitic species have either (i) specialized structural properties, to meet the demands of a brood-parasitic strategy or (ii) similar structural properties to eggs of their hosts, due to the similar nest environment. We measured the surface topography (roughness), wettability (how well surfaces repel water) and calcium content of eggshells of a phylogenetically and geographically diverse range of brood-parasitic species (representing four of the seven independent lineages of avian brood-parasitic species), their hosts and close relatives of the parasites. These components of the eggshell structure have been demonstrated previously to influence such factors as the risk of microbial infection and overall shell strength. Within a phylogenetically controlled framework, we found no overall significant differences in eggshell roughness, wettability and calcium content between (i) parasitic and non-parasitic species, or (ii) parasitic species and their hosts. Both the wettability and calcium content of the eggs from brood-parasitic species were not more similar to those of their hosts' eggs than expected by chance. By contrast, the mean surface roughness of the eggs of brood-parasitic species was more similar to that of their hosts’ eggs than expected by chance, suggesting brood-parasitic species may have evolved to lay eggs that match the host nest environment for this trait. The lack of significant overall differences between parasitic and non-parasitic species, including hosts, in the traits we measured, suggests that phylogenetic signal, as well as general adaptations to the nest environment and for embryo development, outweigh any influence of a parasitic lifestyle on these eggshell properties.</p>
</abstract>
<kwd-group>
<kwd>brood parasitism</kwd><x xml:space="preserve">, </x>
<kwd>calcium carbonate</kwd><x xml:space="preserve">, </x>
<kwd>coevolutionary arms race</kwd><x xml:space="preserve">, </x>
<kwd>cuckoos</kwd><x xml:space="preserve">, </x>
<kwd>profilometry</kwd><x xml:space="preserve">, </x>
<kwd>surface topography</kwd>
</kwd-group><funding-group specific-use="FundRef"><award-group><funding-source><institution-wrap><institution>The Leverhulme Trust</institution></institution-wrap></funding-source><award-id>RPG-2018-332</award-id></award-group></funding-group><funding-group specific-use="FundRef"><award-group><funding-source><institution-wrap><institution>Natural Environment Research Council</institution><institution-id>http://dx.doi.org/10.13039/501100000270</institution-id></institution-wrap></funding-source><award-id>PhD Studentship</award-id></award-group></funding-group>
</article-meta>
</front>
</article>
