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<title xml:lang="en">Chromosomal rearrangements maintain a polymorphic supergene controlling butterfly mimicry</title>
<author>
<name sortKey="Joron, Mathieu" sort="Joron, Mathieu" uniqKey="Joron M" first="Mathieu" last="Joron">Mathieu Joron</name>
<affiliation>
<nlm:aff id="A1">CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP50, 45 Rue Buffon, 75005 Paris, France.</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="A2">Institute of Evolutionary Biology, University of Edinburgh, Ashworth Laboratories, King’s Buildings, West Mains Road, Edinburgh EH9 3JT, UK.</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="A3">Institute of Biology, Leiden University, Postbus 9505, 2300 RA Leiden, The Netherlands.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Frezal, Lise" sort="Frezal, Lise" uniqKey="Frezal L" first="Lise" last="Frezal">Lise Frezal</name>
<affiliation>
<nlm:aff id="A1">CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP50, 45 Rue Buffon, 75005 Paris, France.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Jones, Robert T" sort="Jones, Robert T" uniqKey="Jones R" first="Robert T." last="Jones">Robert T. Jones</name>
<affiliation>
<nlm:aff id="A4">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn, Cornwall TR10 9EZ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Chamberlain, Nicola L" sort="Chamberlain, Nicola L" uniqKey="Chamberlain N" first="Nicola L." last="Chamberlain">Nicola L. Chamberlain</name>
<affiliation>
<nlm:aff id="A4">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn, Cornwall TR10 9EZ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Lee, Siu F" sort="Lee, Siu F" uniqKey="Lee S" first="Siu F." last="Lee">Siu F. Lee</name>
<affiliation>
<nlm:aff id="A5">Department of Genetics, Bio21 Institute, University of Melbourne, 30 Flemington Road, Parkville, 3010 Victoria, Australia.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Haag, Christoph R" sort="Haag, Christoph R" uniqKey="Haag C" first="Christoph R." last="Haag">Christoph R. Haag</name>
<affiliation>
<nlm:aff id="A6">Department of Biology, Ecology and Evolution, University of Fribourg, Chemin du Musée 10, CH-1700 Fribourg, Switzerland.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Whibley, Annabel" sort="Whibley, Annabel" uniqKey="Whibley A" first="Annabel" last="Whibley">Annabel Whibley</name>
<affiliation>
<nlm:aff id="A1">CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP50, 45 Rue Buffon, 75005 Paris, France.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Becuwe, Michel" sort="Becuwe, Michel" uniqKey="Becuwe M" first="Michel" last="Becuwe">Michel Becuwe</name>
<affiliation>
<nlm:aff id="A2">Institute of Evolutionary Biology, University of Edinburgh, Ashworth Laboratories, King’s Buildings, West Mains Road, Edinburgh EH9 3JT, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Baxter, Simon W" sort="Baxter, Simon W" uniqKey="Baxter S" first="Simon W." last="Baxter">Simon W. Baxter</name>
<affiliation>
<nlm:aff id="A7">Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Ferguson, Laura" sort="Ferguson, Laura" uniqKey="Ferguson L" first="Laura" last="Ferguson">Laura Ferguson</name>
<affiliation>
<nlm:aff id="A7">Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Wilkinson, Paul A" sort="Wilkinson, Paul A" uniqKey="Wilkinson P" first="Paul A." last="Wilkinson">Paul A. Wilkinson</name>
<affiliation>
<nlm:aff id="A4">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn, Cornwall TR10 9EZ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Salazar, Camilo" sort="Salazar, Camilo" uniqKey="Salazar C" first="Camilo" last="Salazar">Camilo Salazar</name>
<affiliation>
<nlm:aff id="A8">Smithsonian Tropical Research Institute, NAOS island, Causeway Amador, Panamá, República de Panamá.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Davidson, Claire" sort="Davidson, Claire" uniqKey="Davidson C" first="Claire" last="Davidson">Claire Davidson</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Clark, Richard" sort="Clark, Richard" uniqKey="Clark R" first="Richard" last="Clark">Richard Clark</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Quail, Michael A" sort="Quail, Michael A" uniqKey="Quail M" first="Michael A." last="Quail">Michael A. Quail</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Beasley, Helen" sort="Beasley, Helen" uniqKey="Beasley H" first="Helen" last="Beasley">Helen Beasley</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Glithero, Rebecca" sort="Glithero, Rebecca" uniqKey="Glithero R" first="Rebecca" last="Glithero">Rebecca Glithero</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Lloyd, Christine" sort="Lloyd, Christine" uniqKey="Lloyd C" first="Christine" last="Lloyd">Christine Lloyd</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Sims, Sarah" sort="Sims, Sarah" uniqKey="Sims S" first="Sarah" last="Sims">Sarah Sims</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Jones, Matthew C" sort="Jones, Matthew C" uniqKey="Jones M" first="Matthew C." last="Jones">Matthew C. Jones</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Rogers, Jane" sort="Rogers, Jane" uniqKey="Rogers J" first="Jane" last="Rogers">Jane Rogers</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Jiggins, Chris D" sort="Jiggins, Chris D" uniqKey="Jiggins C" first="Chris D." last="Jiggins">Chris D. Jiggins</name>
<affiliation>
<nlm:aff id="A7">Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Ffrench Constant, Richard H" sort="Ffrench Constant, Richard H" uniqKey="Ffrench Constant R" first="Richard H." last="Ffrench-Constant">Richard H. Ffrench-Constant</name>
<affiliation>
<nlm:aff id="A4">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn, Cornwall TR10 9EZ, UK.</nlm:aff>
</affiliation>
</author>
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<date when="2011">2011</date>
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<title xml:lang="en" level="a" type="main">Chromosomal rearrangements maintain a polymorphic supergene controlling butterfly mimicry</title>
<author>
<name sortKey="Joron, Mathieu" sort="Joron, Mathieu" uniqKey="Joron M" first="Mathieu" last="Joron">Mathieu Joron</name>
<affiliation>
<nlm:aff id="A1">CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP50, 45 Rue Buffon, 75005 Paris, France.</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="A2">Institute of Evolutionary Biology, University of Edinburgh, Ashworth Laboratories, King’s Buildings, West Mains Road, Edinburgh EH9 3JT, UK.</nlm:aff>
</affiliation>
<affiliation>
<nlm:aff id="A3">Institute of Biology, Leiden University, Postbus 9505, 2300 RA Leiden, The Netherlands.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Frezal, Lise" sort="Frezal, Lise" uniqKey="Frezal L" first="Lise" last="Frezal">Lise Frezal</name>
<affiliation>
<nlm:aff id="A1">CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP50, 45 Rue Buffon, 75005 Paris, France.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Jones, Robert T" sort="Jones, Robert T" uniqKey="Jones R" first="Robert T." last="Jones">Robert T. Jones</name>
<affiliation>
<nlm:aff id="A4">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn, Cornwall TR10 9EZ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Chamberlain, Nicola L" sort="Chamberlain, Nicola L" uniqKey="Chamberlain N" first="Nicola L." last="Chamberlain">Nicola L. Chamberlain</name>
<affiliation>
<nlm:aff id="A4">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn, Cornwall TR10 9EZ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Lee, Siu F" sort="Lee, Siu F" uniqKey="Lee S" first="Siu F." last="Lee">Siu F. Lee</name>
<affiliation>
<nlm:aff id="A5">Department of Genetics, Bio21 Institute, University of Melbourne, 30 Flemington Road, Parkville, 3010 Victoria, Australia.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Haag, Christoph R" sort="Haag, Christoph R" uniqKey="Haag C" first="Christoph R." last="Haag">Christoph R. Haag</name>
<affiliation>
<nlm:aff id="A6">Department of Biology, Ecology and Evolution, University of Fribourg, Chemin du Musée 10, CH-1700 Fribourg, Switzerland.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Whibley, Annabel" sort="Whibley, Annabel" uniqKey="Whibley A" first="Annabel" last="Whibley">Annabel Whibley</name>
<affiliation>
<nlm:aff id="A1">CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP50, 45 Rue Buffon, 75005 Paris, France.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Becuwe, Michel" sort="Becuwe, Michel" uniqKey="Becuwe M" first="Michel" last="Becuwe">Michel Becuwe</name>
<affiliation>
<nlm:aff id="A2">Institute of Evolutionary Biology, University of Edinburgh, Ashworth Laboratories, King’s Buildings, West Mains Road, Edinburgh EH9 3JT, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Baxter, Simon W" sort="Baxter, Simon W" uniqKey="Baxter S" first="Simon W." last="Baxter">Simon W. Baxter</name>
<affiliation>
<nlm:aff id="A7">Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Ferguson, Laura" sort="Ferguson, Laura" uniqKey="Ferguson L" first="Laura" last="Ferguson">Laura Ferguson</name>
<affiliation>
<nlm:aff id="A7">Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Wilkinson, Paul A" sort="Wilkinson, Paul A" uniqKey="Wilkinson P" first="Paul A." last="Wilkinson">Paul A. Wilkinson</name>
<affiliation>
<nlm:aff id="A4">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn, Cornwall TR10 9EZ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Salazar, Camilo" sort="Salazar, Camilo" uniqKey="Salazar C" first="Camilo" last="Salazar">Camilo Salazar</name>
<affiliation>
<nlm:aff id="A8">Smithsonian Tropical Research Institute, NAOS island, Causeway Amador, Panamá, República de Panamá.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Davidson, Claire" sort="Davidson, Claire" uniqKey="Davidson C" first="Claire" last="Davidson">Claire Davidson</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Clark, Richard" sort="Clark, Richard" uniqKey="Clark R" first="Richard" last="Clark">Richard Clark</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Quail, Michael A" sort="Quail, Michael A" uniqKey="Quail M" first="Michael A." last="Quail">Michael A. Quail</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Beasley, Helen" sort="Beasley, Helen" uniqKey="Beasley H" first="Helen" last="Beasley">Helen Beasley</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Glithero, Rebecca" sort="Glithero, Rebecca" uniqKey="Glithero R" first="Rebecca" last="Glithero">Rebecca Glithero</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Lloyd, Christine" sort="Lloyd, Christine" uniqKey="Lloyd C" first="Christine" last="Lloyd">Christine Lloyd</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Sims, Sarah" sort="Sims, Sarah" uniqKey="Sims S" first="Sarah" last="Sims">Sarah Sims</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Jones, Matthew C" sort="Jones, Matthew C" uniqKey="Jones M" first="Matthew C." last="Jones">Matthew C. Jones</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Rogers, Jane" sort="Rogers, Jane" uniqKey="Rogers J" first="Jane" last="Rogers">Jane Rogers</name>
<affiliation>
<nlm:aff id="A9">The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Jiggins, Chris D" sort="Jiggins, Chris D" uniqKey="Jiggins C" first="Chris D." last="Jiggins">Chris D. Jiggins</name>
<affiliation>
<nlm:aff id="A7">Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK.</nlm:aff>
</affiliation>
</author>
<author>
<name sortKey="Ffrench Constant, Richard H" sort="Ffrench Constant, Richard H" uniqKey="Ffrench Constant R" first="Richard H." last="Ffrench-Constant">Richard H. Ffrench-Constant</name>
<affiliation>
<nlm:aff id="A4">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn, Cornwall TR10 9EZ, UK.</nlm:aff>
</affiliation>
</author>
</analytic>
<series>
<title level="j">Nature</title>
<idno type="ISSN">0028-0836</idno>
<idno type="eISSN">1476-4687</idno>
<imprint>
<date when="2011">2011</date>
</imprint>
</series>
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<front>
<div type="abstract" xml:lang="en">
<p id="P2">Supergenes are tight clusters of loci that facilitate the co-segregation of adaptive variation, providing integrated control of complex adaptive phenotypes
<sup>
<xref ref-type="bibr" rid="R1">1</xref>
</sup>
. Polymorphic supergenes, in which specific combinations of traits are maintained within a single population, were first described for ‘pin’ and ‘thrum’ floral types in
<italic>Primula</italic>
<sup>
<xref ref-type="bibr" rid="R1">1</xref>
</sup>
and
<italic>Fagopyrum</italic>
<sup>
<xref ref-type="bibr" rid="R2">2</xref>
</sup>
, but classic examples are also found in insect mimicry
<sup>
<xref ref-type="bibr" rid="R3">3</xref>
<xref ref-type="bibr" rid="R5">5</xref>
</sup>
and snail morphology
<sup>
<xref ref-type="bibr" rid="R6">6</xref>
</sup>
. Understanding the evolutionary mechanisms that generate these co-adapted gene sets, as well as the mode of limiting the production of unfit recombinant forms, remains a substantial challenge
<sup>
<xref ref-type="bibr" rid="R7">7</xref>
<xref ref-type="bibr" rid="R10">10</xref>
</sup>
. Here we show that individual wing-pattern morphs in the polymorphic mimetic butterfly
<italic>Heliconius numata</italic>
are associated with different genomic rearrangements at the supergene locus
<italic>P</italic>
. These rearrangements tighten the genetic linkage between at least two colour-pattern loci that are known to recombine in closely related species
<sup>
<xref ref-type="bibr" rid="R9">9</xref>
<xref ref-type="bibr" rid="R11">11</xref>
</sup>
, with complete suppression of recombination being observed in experimental crosses across a 400-kilobase interval containing at least 18 genes. In natural populations, notable patterns of linkage disequilibrium (LD) are observed across the entire
<italic>P</italic>
region. The resulting divergent haplotype clades and inversion breakpoints are found in complete association with wing-pattern morphs. Our results indicate that allelic combinations at known wing-patterning loci have become locked together in a polymorphic rearrangement at the
<italic>P</italic>
locus, forming a supergene that acts as a simple switch between complex adaptive phenotypes found in sympatry. These findings highlight how genomic rearrangements can have a central role in the coexistence of adaptive phenotypes involving several genes acting in concert, by locally limiting recombination and gene flow.</p>
</div>
</front>
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<pmc-comment>The publisher of this article does not allow downloading of the full text in XML form.</pmc-comment>
<pmc-dir>properties manuscript</pmc-dir>
<front>
<journal-meta>
<journal-id journal-id-type="nlm-journal-id">0410462</journal-id>
<journal-id journal-id-type="pubmed-jr-id">6011</journal-id>
<journal-id journal-id-type="nlm-ta">Nature</journal-id>
<journal-id journal-id-type="iso-abbrev">Nature</journal-id>
<journal-title-group>
<journal-title>Nature</journal-title>
</journal-title-group>
<issn pub-type="ppub">0028-0836</issn>
<issn pub-type="epub">1476-4687</issn>
</journal-meta>
<article-meta>
<article-id pub-id-type="pmid">21841803</article-id>
<article-id pub-id-type="pmc">3717454</article-id>
<article-id pub-id-type="doi">10.1038/nature10341</article-id>
<article-id pub-id-type="manuscript">EMS52362</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Article</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Chromosomal rearrangements maintain a polymorphic supergene controlling butterfly mimicry</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Joron</surname>
<given-names>Mathieu</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
<xref ref-type="aff" rid="A2">2</xref>
<xref ref-type="aff" rid="A3">3</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Frezal</surname>
<given-names>Lise</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
<xref ref-type="author-notes" rid="FN1">*</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Jones</surname>
<given-names>Robert T.</given-names>
</name>
<xref ref-type="aff" rid="A4">4</xref>
<xref ref-type="author-notes" rid="FN1">*</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Chamberlain</surname>
<given-names>Nicola L.</given-names>
</name>
<xref ref-type="aff" rid="A4">4</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Lee</surname>
<given-names>Siu F.</given-names>
</name>
<xref ref-type="aff" rid="A5">5</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Haag</surname>
<given-names>Christoph R.</given-names>
</name>
<xref ref-type="aff" rid="A6">6</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Whibley</surname>
<given-names>Annabel</given-names>
</name>
<xref ref-type="aff" rid="A1">1</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Becuwe</surname>
<given-names>Michel</given-names>
</name>
<xref ref-type="aff" rid="A2">2</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Baxter</surname>
<given-names>Simon W.</given-names>
</name>
<xref ref-type="aff" rid="A7">7</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ferguson</surname>
<given-names>Laura</given-names>
</name>
<xref ref-type="aff" rid="A7">7</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Wilkinson</surname>
<given-names>Paul A.</given-names>
</name>
<xref ref-type="aff" rid="A4">4</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Salazar</surname>
<given-names>Camilo</given-names>
</name>
<xref ref-type="aff" rid="A8">8</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Davidson</surname>
<given-names>Claire</given-names>
</name>
<xref ref-type="aff" rid="A9">9</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Clark</surname>
<given-names>Richard</given-names>
</name>
<xref ref-type="aff" rid="A9">9</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Quail</surname>
<given-names>Michael A.</given-names>
</name>
<xref ref-type="aff" rid="A9">9</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Beasley</surname>
<given-names>Helen</given-names>
</name>
<xref ref-type="aff" rid="A9">9</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Glithero</surname>
<given-names>Rebecca</given-names>
</name>
<xref ref-type="aff" rid="A9">9</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Lloyd</surname>
<given-names>Christine</given-names>
</name>
<xref ref-type="aff" rid="A9">9</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Sims</surname>
<given-names>Sarah</given-names>
</name>
<xref ref-type="aff" rid="A9">9</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Jones</surname>
<given-names>Matthew C.</given-names>
</name>
<xref ref-type="aff" rid="A9">9</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Rogers</surname>
<given-names>Jane</given-names>
</name>
<xref ref-type="aff" rid="A9">9</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Jiggins</surname>
<given-names>Chris D.</given-names>
</name>
<xref ref-type="aff" rid="A7">7</xref>
</contrib>
<contrib contrib-type="author">
<name>
<surname>ffrench-Constant</surname>
<given-names>Richard H.</given-names>
</name>
<xref ref-type="aff" rid="A4">4</xref>
</contrib>
</contrib-group>
<aff id="A1">
<label>1</label>
CNRS UMR 7205, Muséum National d’Histoire Naturelle, CP50, 45 Rue Buffon, 75005 Paris, France.</aff>
<aff id="A2">
<label>2</label>
Institute of Evolutionary Biology, University of Edinburgh, Ashworth Laboratories, King’s Buildings, West Mains Road, Edinburgh EH9 3JT, UK.</aff>
<aff id="A3">
<label>3</label>
Institute of Biology, Leiden University, Postbus 9505, 2300 RA Leiden, The Netherlands.</aff>
<aff id="A4">
<label>4</label>
Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Cornwall Campus, Penryn, Cornwall TR10 9EZ, UK.</aff>
<aff id="A5">
<label>5</label>
Department of Genetics, Bio21 Institute, University of Melbourne, 30 Flemington Road, Parkville, 3010 Victoria, Australia.</aff>
<aff id="A6">
<label>6</label>
Department of Biology, Ecology and Evolution, University of Fribourg, Chemin du Musée 10, CH-1700 Fribourg, Switzerland.</aff>
<aff id="A7">
<label>7</label>
Department of Zoology, University of Cambridge, Downing Street, Cambridge CB2 3EJ, UK.</aff>
<aff id="A8">
<label>8</label>
Smithsonian Tropical Research Institute, NAOS island, Causeway Amador, Panamá, República de Panamá.</aff>
<aff id="A9">
<label>9</label>
The Wellcome Trust Sanger Institute, Hinxton, Cambridge CB10 1HH, UK.</aff>
<author-notes>
<corresp id="CR1">Correspondence and requests for materials should be addressed to M.J. (
<email>joron@mnhn.fr</email>
).</corresp>
<fn fn-type="equal" id="FN1">
<label>*</label>
<p id="P1">These authors contributed equally to this work.</p>
</fn>
</author-notes>
<pub-date pub-type="nihms-submitted">
<day>5</day>
<month>7</month>
<year>2013</year>
</pub-date>
<pub-date pub-type="epub">
<day>14</day>
<month>8</month>
<year>2011</year>
</pub-date>
<pub-date pub-type="pmc-release">
<day>22</day>
<month>7</month>
<year>2013</year>
</pub-date>
<volume>477</volume>
<issue>7363</issue>
<fpage>203</fpage>
<lpage>206</lpage>
<abstract>
<p id="P2">Supergenes are tight clusters of loci that facilitate the co-segregation of adaptive variation, providing integrated control of complex adaptive phenotypes
<sup>
<xref ref-type="bibr" rid="R1">1</xref>
</sup>
. Polymorphic supergenes, in which specific combinations of traits are maintained within a single population, were first described for ‘pin’ and ‘thrum’ floral types in
<italic>Primula</italic>
<sup>
<xref ref-type="bibr" rid="R1">1</xref>
</sup>
and
<italic>Fagopyrum</italic>
<sup>
<xref ref-type="bibr" rid="R2">2</xref>
</sup>
, but classic examples are also found in insect mimicry
<sup>
<xref ref-type="bibr" rid="R3">3</xref>
<xref ref-type="bibr" rid="R5">5</xref>
</sup>
and snail morphology
<sup>
<xref ref-type="bibr" rid="R6">6</xref>
</sup>
. Understanding the evolutionary mechanisms that generate these co-adapted gene sets, as well as the mode of limiting the production of unfit recombinant forms, remains a substantial challenge
<sup>
<xref ref-type="bibr" rid="R7">7</xref>
<xref ref-type="bibr" rid="R10">10</xref>
</sup>
. Here we show that individual wing-pattern morphs in the polymorphic mimetic butterfly
<italic>Heliconius numata</italic>
are associated with different genomic rearrangements at the supergene locus
<italic>P</italic>
. These rearrangements tighten the genetic linkage between at least two colour-pattern loci that are known to recombine in closely related species
<sup>
<xref ref-type="bibr" rid="R9">9</xref>
<xref ref-type="bibr" rid="R11">11</xref>
</sup>
, with complete suppression of recombination being observed in experimental crosses across a 400-kilobase interval containing at least 18 genes. In natural populations, notable patterns of linkage disequilibrium (LD) are observed across the entire
<italic>P</italic>
region. The resulting divergent haplotype clades and inversion breakpoints are found in complete association with wing-pattern morphs. Our results indicate that allelic combinations at known wing-patterning loci have become locked together in a polymorphic rearrangement at the
<italic>P</italic>
locus, forming a supergene that acts as a simple switch between complex adaptive phenotypes found in sympatry. These findings highlight how genomic rearrangements can have a central role in the coexistence of adaptive phenotypes involving several genes acting in concert, by locally limiting recombination and gene flow.</p>
</abstract>
<funding-group>
<award-group>
<funding-source country="United Kingdom">Wellcome Trust : </funding-source>
<award-id>098051 || WT</award-id>
</award-group>
</funding-group>
</article-meta>
</front>
</pmc>
</record>

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