<?xml version="1.0" encoding="utf-8"?>
<TEI xmlns="http://www.tei-c.org/ns/1.0" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" xmlns:hal="http://hal.archives-ouvertes.fr/" xmlns:gml="http://www.opengis.net/gml/3.3/" xmlns:gmlce="http://www.opengis.net/gml/3.3/ce" version="1.1" xsi:schemaLocation="http://www.tei-c.org/ns/1.0 http://api.archives-ouvertes.fr/documents/aofr-sword.xsd">
  <teiHeader>
    <fileDesc>
      <titleStmt>
        <title>HAL TEI export of hal-02907425</title>
      </titleStmt>
      <publicationStmt>
        <distributor>CCSD</distributor>
        <availability status="restricted">
          <licence target="https://creativecommons.org/publicdomain/zero/1.0/">CC0 1.0 - Universal</licence>
        </availability>
        <date when="2026-05-18T13:21:39+02:00"/>
      </publicationStmt>
      <sourceDesc>
        <p part="N">HAL API Platform</p>
      </sourceDesc>
    </fileDesc>
  </teiHeader>
  <text>
    <body>
      <listBibl>
        <biblFull>
          <titleStmt>
            <title xml:lang="en">Wonky whales: the evolution of cranial asymmetry in cetaceans</title>
            <author role="aut">
              <persName>
                <forename type="first">Ellen</forename>
                <surname>Coombs</surname>
              </persName>
              <idno type="halauthorid">2005967-0</idno>
              <affiliation ref="#struct-431958"/>
            </author>
            <author role="aut">
              <persName>
                <forename type="first">Julien</forename>
                <surname>Clavel</surname>
              </persName>
              <email type="md5">265bb9c62b7ed67c9303694ee49c37df</email>
              <email type="domain">univ-lyon1.fr</email>
              <idno type="idhal" notation="string">julien-clavel</idno>
              <idno type="idhal" notation="numeric">750439</idno>
              <idno type="halauthorid" notation="string">21863-750439</idno>
              <idno type="RESEARCHERID">http://www.researcherid.com/rid/O-3215-2017</idno>
              <idno type="ORCID">https://orcid.org/0000-0002-2715-4988</idno>
              <idno type="IDREF">https://www.idref.fr/225622467</idno>
              <idno type="RESEARCHERID">http://www.researcherid.com/rid/http://www.researcherid.com/rid/O-3215-2017</idno>
              <affiliation ref="#struct-215697"/>
            </author>
            <author role="aut">
              <persName>
                <forename type="first">Travis</forename>
                <surname>Park</surname>
              </persName>
              <idno type="halauthorid">1691109-0</idno>
              <affiliation ref="#struct-310641"/>
            </author>
            <author role="aut">
              <persName>
                <forename type="first">Morgan</forename>
                <surname>Churchill</surname>
              </persName>
              <idno type="halauthorid">1858221-0</idno>
              <affiliation ref="#struct-328948"/>
            </author>
            <author role="aut">
              <persName>
                <forename type="first">Anjali</forename>
                <surname>Goswami</surname>
              </persName>
              <idno type="halauthorid">1422832-0</idno>
              <affiliation ref="#struct-431958"/>
            </author>
            <editor role="depositor">
              <persName>
                <forename>Nathalie</forename>
                <surname>Lyvet</surname>
              </persName>
              <email type="md5">d610acca244c9a817cb89db4405e6403</email>
              <email type="domain">univ-lyon1.fr</email>
            </editor>
          </titleStmt>
          <editionStmt>
            <edition n="v1" type="current">
              <date type="whenSubmitted">2021-11-29 15:36:50</date>
              <date type="whenModified">2026-02-07 05:22:14</date>
              <date type="whenReleased">2021-12-14 17:42:27</date>
              <date type="whenProduced">2020</date>
              <date type="whenEndEmbargoed">2021-11-29</date>
              <ref type="file" target="https://univ-lyon1.hal.science/hal-02907425v1/document">
                <date notBefore="2021-11-29"/>
              </ref>
              <ref type="file" subtype="greenPublisher" n="1" target="https://univ-lyon1.hal.science/hal-02907425v1/file/8-Coombs%20et%20al%202020.pdf" id="file-3455253-3052360">
                <date notBefore="2021-11-29"/>
              </ref>
              <ref type="externalLink" target="https://bmcbiol.biomedcentral.com/track/pdf/10.1186/s12915-020-00805-4"/>
            </edition>
            <respStmt>
              <resp>contributor</resp>
              <name key="121211">
                <persName>
                  <forename>Nathalie</forename>
                  <surname>Lyvet</surname>
                </persName>
                <email type="md5">d610acca244c9a817cb89db4405e6403</email>
                <email type="domain">univ-lyon1.fr</email>
              </name>
            </respStmt>
          </editionStmt>
          <publicationStmt>
            <distributor>CCSD</distributor>
            <idno type="halId">hal-02907425</idno>
            <idno type="halUri">https://univ-lyon1.hal.science/hal-02907425</idno>
            <idno type="halBibtex">coombs:hal-02907425</idno>
            <idno type="halRefHtml">&lt;i&gt;BMC Biology&lt;/i&gt;, 2020, 18 (1), pp.1-24. &lt;a target="_blank" href="https://dx.doi.org/10.1186/s12915-020-00805-4"&gt;&amp;#x27E8;10.1186/s12915-020-00805-4&amp;#x27E9;&lt;/a&gt;</idno>
            <idno type="halRef">BMC Biology, 2020, 18 (1), pp.1-24. &amp;#x27E8;10.1186/s12915-020-00805-4&amp;#x27E9;</idno>
            <availability status="restricted">
              <licence target="https://about.hal.science/hal-authorisation-v1/">HAL Authorization<ref corresp="#file-3455253-3052360"/></licence>
            </availability>
          </publicationStmt>
          <seriesStmt>
            <idno type="stamp" n="CNRS">CNRS - Centre national de la recherche scientifique</idno>
            <idno type="stamp" n="UNIV-LYON1">Université Claude Bernard - Lyon I</idno>
            <idno type="stamp" n="ENTPE">Ecole Nationale des Travaux Publics de l'Etat</idno>
            <idno type="stamp" n="GIP-BE">GIP Bretagne Environnement</idno>
            <idno type="stamp" n="UDL">UDL</idno>
            <idno type="stamp" n="UNIV-LYON">Université de Lyon</idno>
            <idno type="stamp" n="TEST-HALCNRS">Collection test HAL CNRS</idno>
            <idno type="stamp" n="LEHNA" corresp="UNIV-LYON1">Laboratoire d'Ecologie des Hydrosystèmes Naturels et Anthropisés</idno>
            <idno type="stamp" n="TEST3-HALCNRS">TEST3-HALCNRS</idno>
            <idno type="stamp" n="TEST4-HALCNRS">collection test</idno>
            <idno type="stamp" n="INEE-CNRS">Institut écologie et environnement du CNRS</idno>
          </seriesStmt>
          <notesStmt>
            <note type="audience" n="2">International</note>
            <note type="popular" n="0">No</note>
            <note type="peer" n="1">Yes</note>
          </notesStmt>
          <sourceDesc>
            <biblStruct>
              <analytic>
                <title xml:lang="en">Wonky whales: the evolution of cranial asymmetry in cetaceans</title>
                <author role="aut">
                  <persName>
                    <forename type="first">Ellen</forename>
                    <surname>Coombs</surname>
                  </persName>
                  <idno type="halauthorid">2005967-0</idno>
                  <affiliation ref="#struct-431958"/>
                </author>
                <author role="aut">
                  <persName>
                    <forename type="first">Julien</forename>
                    <surname>Clavel</surname>
                  </persName>
                  <email type="md5">265bb9c62b7ed67c9303694ee49c37df</email>
                  <email type="domain">univ-lyon1.fr</email>
                  <idno type="idhal" notation="string">julien-clavel</idno>
                  <idno type="idhal" notation="numeric">750439</idno>
                  <idno type="halauthorid" notation="string">21863-750439</idno>
                  <idno type="RESEARCHERID">http://www.researcherid.com/rid/O-3215-2017</idno>
                  <idno type="ORCID">https://orcid.org/0000-0002-2715-4988</idno>
                  <idno type="IDREF">https://www.idref.fr/225622467</idno>
                  <idno type="RESEARCHERID">http://www.researcherid.com/rid/http://www.researcherid.com/rid/O-3215-2017</idno>
                  <affiliation ref="#struct-215697"/>
                </author>
                <author role="aut">
                  <persName>
                    <forename type="first">Travis</forename>
                    <surname>Park</surname>
                  </persName>
                  <idno type="halauthorid">1691109-0</idno>
                  <affiliation ref="#struct-310641"/>
                </author>
                <author role="aut">
                  <persName>
                    <forename type="first">Morgan</forename>
                    <surname>Churchill</surname>
                  </persName>
                  <idno type="halauthorid">1858221-0</idno>
                  <affiliation ref="#struct-328948"/>
                </author>
                <author role="aut">
                  <persName>
                    <forename type="first">Anjali</forename>
                    <surname>Goswami</surname>
                  </persName>
                  <idno type="halauthorid">1422832-0</idno>
                  <affiliation ref="#struct-431958"/>
                </author>
              </analytic>
              <monogr>
                <idno type="halJournalId" status="VALID">313</idno>
                <idno type="issn">1741-7007</idno>
                <idno type="eissn">1741-7007</idno>
                <title level="j">BMC Biology</title>
                <imprint>
                  <publisher>BioMed Central</publisher>
                  <biblScope unit="volume">18</biblScope>
                  <biblScope unit="issue">1</biblScope>
                  <biblScope unit="pp">1-24</biblScope>
                  <date type="datePub">2020</date>
                </imprint>
              </monogr>
              <idno type="doi">10.1186/s12915-020-00805-4</idno>
            </biblStruct>
          </sourceDesc>
          <profileDesc>
            <langUsage>
              <language ident="en">English</language>
            </langUsage>
            <textClass>
              <keywords scheme="author">
                <term xml:lang="en">Macroevolution</term>
                <term xml:lang="en">Morphometrics</term>
                <term xml:lang="en">Cetaceans</term>
                <term xml:lang="en">Asymmetry</term>
                <term xml:lang="en">Trait evolution</term>
              </keywords>
              <classCode scheme="halDomain" n="sdv.ba.zv">Life Sciences [q-bio]/Animal biology/Vertebrate Zoology</classCode>
              <classCode scheme="halDomain" n="sdv.bid.evo">Life Sciences [q-bio]/Biodiversity/Populations and Evolution [q-bio.PE]</classCode>
              <classCode scheme="halDomain" n="sdu.stu.pg">Sciences of the Universe [physics]/Earth Sciences/Paleontology</classCode>
              <classCode scheme="halTypology" n="ART">Journal articles</classCode>
              <classCode scheme="halOldTypology" n="ART">Journal articles</classCode>
              <classCode scheme="halTreeTypology" n="ART">Journal articles</classCode>
            </textClass>
            <abstract xml:lang="en">
              <p>Background: Unlike most mammals, toothed whale (Odontoceti) skulls lack symmetry in the nasal and facial (nasofacial) region. This asymmetry is hypothesised to relate to echolocation, which may have evolved in the earliest diverging odontocetes. Early cetaceans (whales, dolphins, and porpoises) such as archaeocetes, namely the protocetids and basilosaurids, have asymmetric rostra, but it is unclear when nasofacial asymmetry evolved during the transition from archaeocetes to modern whales. We used three-dimensional geometric morphometrics and phylogenetic comparative methods to reconstruct the evolution of asymmetry in the skulls of 162 living and extinct cetaceans over 50 million years.Results: In archaeocetes, we found asymmetry is prevalent in the rostrum and also in the squamosal, jugal, and orbit, possibly reflecting preservational deformation. Asymmetry in odontocetes is predominant in the nasofacial region. Mysticetes (baleen whales) show symmetry similar to terrestrial artiodactyls such as bovines. The first significant shift in asymmetry occurred in the stem odontocete family Xenorophidae during the Early Oligocene. Further increases in asymmetry occur in the physeteroids in the Late Oligocene, Squalodelphinidae and Platanistidae in the Late Oligocene/Early Miocene, and in the Monodontidae in the Late Miocene/Early Pliocene. Additional episodes of rapid change in odontocete skull asymmetry were found in the Mid-Late Oligocene, a period of rapid evolution and diversification. No high-probability increases or jumps in asymmetry were found in mysticetes or archaeocetes. Unexpectedly, no increases in asymmetry were recovered within the highly asymmetric ziphiids, which may result from the extreme, asymmetric shape of premaxillary crests in these taxa not being captured by landmarks alone.Conclusions: Early ancestors of living whales had little cranial asymmetry and likely were not able to echolocate. Archaeocetes display high levels of asymmetry in the rostrum, potentially related to directional hearing, which is lost in early neocetes—the taxon including the most recent common ancestor of living cetaceans. Nasofacial asymmetry becomes a significant feature of Odontoceti skulls in the Early Oligocene, reaching its highest levels in extant taxa. Separate evolutionary regimes are reconstructed for odontocetes living in acoustically complex environments, suggesting that these niches impose strong selective pressure on echolocation ability and thus increased cranial asymmetry.</p>
            </abstract>
          </profileDesc>
        </biblFull>
      </listBibl>
    </body>
    <back>
      <listOrg type="structures">
        <org type="laboratory" xml:id="struct-431958" status="INCOMING">
          <orgName>Department of Genetics, Evolution and Environment</orgName>
          <desc>
            <address>
              <country key="GB"/>
            </address>
          </desc>
          <listRelation>
            <relation active="#struct-300875" type="direct"/>
          </listRelation>
        </org>
        <org type="laboratory" xml:id="struct-215697" status="VALID">
          <idno type="IdRef">151765413</idno>
          <idno type="ISNI">0000000404461170</idno>
          <idno type="RNSR">199911718W</idno>
          <idno type="ROR">https://ror.org/05pny1q12</idno>
          <orgName>Laboratoire d'Ecologie des Hydrosystèmes Naturels et Anthropisés</orgName>
          <orgName type="acronym">LEHNA</orgName>
          <date type="start">2015-01-01</date>
          <desc>
            <address>
              <addrLine>Université Claude Bernard Lyon 13-6, rue Raphaël Dubois - Bâtiments Darwin C &amp; Forel, 69622 Villeurbanne Cedex43, Boulevard du 11 novembre 1918ENTPE3, rue Maurice Audin69518 Vaulx-en-Velin</addrLine>
              <country key="FR"/>
            </address>
            <ref type="url">http://umr5023.univ-lyon1.fr</ref>
          </desc>
          <listRelation>
            <relation active="#struct-194495" type="direct"/>
            <relation active="#struct-301088" type="indirect"/>
            <relation active="#struct-300365" type="direct"/>
            <relation name="UMR5023" active="#struct-441569" type="direct"/>
          </listRelation>
        </org>
        <org type="institution" xml:id="struct-310641" status="VALID">
          <idno type="ROR">https://ror.org/039zvsn29</idno>
          <orgName>The Natural History Museum [London]</orgName>
          <orgName type="acronym">NHM</orgName>
          <desc>
            <address>
              <country key="GB"/>
            </address>
          </desc>
        </org>
        <org type="institution" xml:id="struct-328948" status="VALID">
          <idno type="ROR">https://ror.org/05w22af52</idno>
          <orgName>University of Wisconsin–Oshkosh</orgName>
          <orgName type="acronym">UWO</orgName>
          <desc>
            <address>
              <country key="US"/>
            </address>
            <ref type="url">https://www.uwosh.edu/</ref>
          </desc>
        </org>
        <org type="institution" xml:id="struct-300875" status="VALID">
          <idno type="IdRef">029789753</idno>
          <idno type="ROR">https://ror.org/02jx3x895</idno>
          <orgName>University College London [UCL]</orgName>
          <orgName type="acronym">UCL</orgName>
          <desc>
            <address>
              <addrLine>Gower Street, London WC1E 6BT</addrLine>
              <country key="GB"/>
            </address>
            <ref type="url">http://www.ucl.ac.uk/</ref>
          </desc>
        </org>
        <org type="institution" xml:id="struct-194495" status="VALID">
          <idno type="IdRef">026402823</idno>
          <idno type="ISNI">0000000121686185</idno>
          <idno type="ROR">https://ror.org/029brtt94</idno>
          <orgName>Université Claude Bernard Lyon 1</orgName>
          <orgName type="acronym">UCBL</orgName>
          <desc>
            <address>
              <addrLine>43, boulevard du 11 novembre 1918, 69622 Villeurbanne cedex</addrLine>
              <country key="FR"/>
            </address>
            <ref type="url">http://www.univ-lyon1.fr/</ref>
          </desc>
          <listRelation>
            <relation active="#struct-301088" type="direct"/>
          </listRelation>
        </org>
        <org type="regroupinstitution" xml:id="struct-301088" status="VALID">
          <idno type="ROR">https://ror.org/01rk35k63</idno>
          <orgName>Université de Lyon</orgName>
          <desc>
            <address>
              <addrLine>92 rue Pasteur - CS 30122, 69361 Lyon Cedex 07</addrLine>
              <country key="FR"/>
            </address>
            <ref type="url">https://www.universite-lyon.fr/</ref>
          </desc>
        </org>
        <org type="institution" xml:id="struct-300365" status="VALID">
          <idno type="ROR">https://ror.org/05r0ccb59</idno>
          <orgName>École Nationale des Travaux Publics de l'État</orgName>
          <orgName type="acronym">ENTPE</orgName>
          <desc>
            <address>
              <addrLine>3, rue Maurice Audin - 69518 Vaulx-en-Velin Cedex</addrLine>
              <country key="FR"/>
            </address>
            <ref type="url">http://www.entpe.fr</ref>
          </desc>
        </org>
        <org type="regroupinstitution" xml:id="struct-441569" status="VALID">
          <idno type="IdRef">02636817X</idno>
          <idno type="ISNI">0000000122597504</idno>
          <idno type="ROR">https://ror.org/02feahw73</idno>
          <orgName>Centre National de la Recherche Scientifique</orgName>
          <orgName type="acronym">CNRS</orgName>
          <date type="start">1939-10-19</date>
          <desc>
            <address>
              <country key="FR"/>
            </address>
            <ref type="url">https://www.cnrs.fr/</ref>
          </desc>
        </org>
      </listOrg>
    </back>
  </text>
</TEI>