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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Butterfly genome reveals promiscuous exchange of mimicry adaptations among species</title><author><name sortKey="Dasmahapatra, Kanchon K" sort="Dasmahapatra, Kanchon K" uniqKey="Dasmahapatra K" first="Kanchon K" last="Dasmahapatra">Kanchon K. Dasmahapatra</name><affiliation><nlm:aff id="A1">Department of Genetics, Evolution and Environment, University College London, Gower Street, London, WC1E 6BT, UK.</nlm:aff></affiliation></author><author><name sortKey="Walters, James R" sort="Walters, James R" uniqKey="Walters J" first="James R." last="Walters">James R. Walters</name><affiliation><nlm:aff id="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, UK.</nlm:aff></affiliation></author><author><name sortKey="Briscoe, Adriana D" sort="Briscoe, Adriana D" uniqKey="Briscoe A" first="Adriana D." last="Briscoe">Adriana D. Briscoe</name><affiliation><nlm:aff id="A3">Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA.</nlm:aff></affiliation></author><author><name sortKey="Davey, John W" sort="Davey, John W" uniqKey="Davey J" first="John W." last="Davey">John W. Davey</name><affiliation><nlm:aff id="A4">Institute of Evolutionary Biology, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</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="A5">CNRS UMR 7205, Muséum National d’Histoire Naturelle, 45 rue Buffon, Paris 75005, France.</nlm:aff></affiliation></author><author><name sortKey="Nadeau, Nicola J" sort="Nadeau, Nicola J" uniqKey="Nadeau N" first="Nicola J." last="Nadeau">Nicola J. Nadeau</name><affiliation><nlm:aff id="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, UK.</nlm:aff></affiliation></author><author><name sortKey="Zimin, Aleksey V" sort="Zimin, Aleksey V" uniqKey="Zimin A" first="Aleksey V." last="Zimin">Aleksey V. Zimin</name><affiliation><nlm:aff id="A6">Institute for Physical Science and Technology, University of Maryland, College Park, MD 20742, USA.</nlm:aff></affiliation></author><author><name sortKey="Hughes, Daniel S T" sort="Hughes, Daniel S T" uniqKey="Hughes D" first="Daniel S. T." last="Hughes">Daniel S. T. Hughes</name><affiliation><nlm:aff id="A7">European Bioinformatics Institute, Hinxton, CB10 1SD, UK.</nlm:aff></affiliation></author><author><name sortKey="Ferguson, Laura C" sort="Ferguson, Laura C" uniqKey="Ferguson L" first="Laura C." last="Ferguson">Laura C. Ferguson</name><affiliation><nlm:aff id="A8">Department of Zoology, University of Oxford, South Parks Road, Oxford, OX1 3PS, UK.</nlm:aff></affiliation></author><author><name sortKey="Martin, Simon H" sort="Martin, Simon H" uniqKey="Martin S" first="Simon H." last="Martin">Simon H. Martin</name><affiliation><nlm:aff id="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, 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="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, UK.</nlm:aff></affiliation><affiliation><nlm:aff id="A9">Smithsonian Tropical Research Institute, Naos Island, Causeway Amador, Panamá, Republic de Panamá.</nlm:aff></affiliation></author><author><name sortKey="Lewis, James J" sort="Lewis, James J" uniqKey="Lewis J" first="James J." last="Lewis">James J. Lewis</name><affiliation><nlm:aff id="A3">Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA.</nlm:aff></affiliation></author><author><name sortKey="Adler, Sebastian" sort="Adler, Sebastian" uniqKey="Adler S" first="Sebastian" last="Adler">Sebastian Adler</name><affiliation><nlm:aff id="A10">Institut für Mathematik und Informatik, Universität Greifswald, 17487 Greifswald, Germany.</nlm:aff></affiliation></author><author><name sortKey="Ahn, Seung Joon" sort="Ahn, Seung Joon" uniqKey="Ahn S" first="Seung-Joon" last="Ahn">Seung-Joon Ahn</name><affiliation><nlm:aff id="A11">Max Planck Institute for Chemical Ecology, D-07745 Jena, Germany.</nlm:aff></affiliation></author><author><name sortKey="Baker, Dean A" sort="Baker, Dean A" uniqKey="Baker D" first="Dean A." last="Baker">Dean A. Baker</name><affiliation><nlm:aff id="A12">Ecology and Evolution, Imperial College London, London, 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="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, 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="A13">FAS Center for Systems Biology, Harvard University, Cambridge, MA 02138, USA.</nlm:aff></affiliation></author><author><name sortKey="Chauhan, Ritika" sort="Chauhan, Ritika" uniqKey="Chauhan R" first="Ritika" last="Chauhan">Ritika Chauhan</name><affiliation><nlm:aff id="A14">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Penryn, TR10 9EZ, UK.</nlm:aff></affiliation></author><author><name sortKey="Counterman, Brian A" sort="Counterman, Brian A" uniqKey="Counterman B" first="Brian A." last="Counterman">Brian A. Counterman</name><affiliation><nlm:aff id="A15">Department of Biology, Mississippi State University, Mississippi State, Mississippi 39762, USA.</nlm:aff></affiliation></author><author><name sortKey="Dalmay, Tamas" sort="Dalmay, Tamas" uniqKey="Dalmay T" first="Tamas" last="Dalmay">Tamas Dalmay</name><affiliation><nlm:aff id="A16">School of Biological Sciences, University of East Anglia, Norwich Research Park, Norwich, NR4 7TJ, UK.</nlm:aff></affiliation></author><author><name sortKey="Gilbert, Lawrence E" sort="Gilbert, Lawrence E" uniqKey="Gilbert L" first="Lawrence E." last="Gilbert">Lawrence E. Gilbert</name><affiliation><nlm:aff id="A17">Section of Integrative Biology and Brackenridge Field Laboratory, University of Texas, Austin, TX 78712, USA.</nlm:aff></affiliation></author><author><name sortKey="Gordon, Karl" sort="Gordon, Karl" uniqKey="Gordon K" first="Karl" last="Gordon">Karl Gordon</name><affiliation><nlm:aff id="A18">Black Mountain Laboratories, CSIRO Ecosystem Sciences, Clunies Ross Street, Canberra, ACT 2601, Australia.</nlm:aff></affiliation></author><author><name sortKey="Heckel, David G" sort="Heckel, David G" uniqKey="Heckel D" first="David G." last="Heckel">David G. Heckel</name><affiliation><nlm:aff id="A11">Max Planck Institute for Chemical Ecology, D-07745 Jena, Germany.</nlm:aff></affiliation></author><author><name sortKey="Hines, Heather M" sort="Hines, Heather M" uniqKey="Hines H" first="Heather M." last="Hines">Heather M. Hines</name><affiliation><nlm:aff id="A19">Department of Genetics, North Carolina State University, Raleigh, NC 27695, USA.</nlm:aff></affiliation></author><author><name sortKey="Hoff, Katharina J" sort="Hoff, Katharina J" uniqKey="Hoff K" first="Katharina J." last="Hoff">Katharina J. Hoff</name><affiliation><nlm:aff id="A10">Institut für Mathematik und Informatik, Universität Greifswald, 17487 Greifswald, Germany.</nlm:aff></affiliation></author><author><name sortKey="Holland, Peter W H" sort="Holland, Peter W H" uniqKey="Holland P" first="Peter W. H." last="Holland">Peter W. H. Holland</name><affiliation><nlm:aff id="A8">Department of Zoology, University of Oxford, South Parks Road, Oxford, OX1 3PS, UK.</nlm:aff></affiliation></author><author><name sortKey="Jacquin Joly, Emmanuelle" sort="Jacquin Joly, Emmanuelle" uniqKey="Jacquin Joly E" first="Emmanuelle" last="Jacquin-Joly">Emmanuelle Jacquin-Joly</name><affiliation><nlm:aff id="A20">Inra, UMR-A 1272 Inra-Université Pierre et Marie Curie, Physiologie de l’Insecte: Signalisation et Communication, Route de Saint-Cyr, Versailles, France.</nlm:aff></affiliation></author><author><name sortKey="Jiggins, Francis M" sort="Jiggins, Francis M" uniqKey="Jiggins F" first="Francis M." last="Jiggins">Francis M. Jiggins</name><affiliation><nlm:aff id="A21">Department of Genetics, Downing Street, University of Cambridge, Cambridge, CB2 3EH, UK.</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="A5">CNRS UMR 7205, Muséum National d’Histoire Naturelle, 45 rue Buffon, Paris 75005, France.</nlm:aff></affiliation></author><author><name sortKey="Kapan, Durrell D" sort="Kapan, Durrell D" uniqKey="Kapan D" first="Durrell D." last="Kapan">Durrell D. Kapan</name><affiliation><nlm:aff id="A22">Department of Entomology, Center for Comparative Genomics, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, California 94118, USA.</nlm:aff></affiliation><affiliation><nlm:aff id="A23">Center for Conservation and Research Training, Pacific Biosciences Research Center, University of Hawaii at Manoa, 3050 Maile Way, Gilmore 406, Honolulu, HI 96822, USA.</nlm:aff></affiliation></author><author><name sortKey="Kersey, Paul" sort="Kersey, Paul" uniqKey="Kersey P" first="Paul" last="Kersey">Paul Kersey</name><affiliation><nlm:aff id="A7">European Bioinformatics Institute, Hinxton, CB10 1SD, UK.</nlm:aff></affiliation></author><author><name sortKey="Lamas, Gerardo" sort="Lamas, Gerardo" uniqKey="Lamas G" first="Gerardo" last="Lamas">Gerardo Lamas</name><affiliation><nlm:aff id="A24">Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Apartado 14-0434, Lima, Peru.</nlm:aff></affiliation></author><author><name sortKey="Lawson, Daniel" sort="Lawson, Daniel" uniqKey="Lawson D" first="Daniel" last="Lawson">Daniel Lawson</name><affiliation><nlm:aff id="A7">European Bioinformatics Institute, Hinxton, CB10 1SD, UK.</nlm:aff></affiliation></author><author><name sortKey="Mapleson, Daniel" sort="Mapleson, Daniel" uniqKey="Mapleson D" first="Daniel" last="Mapleson">Daniel Mapleson</name><affiliation><nlm:aff id="A25">School of Computing Sciences, University of East Anglia, Norwich Research Park, Norwich, NR4 7TJ, UK.</nlm:aff></affiliation></author><author><name sortKey="Maroja, Luana S" sort="Maroja, Luana S" uniqKey="Maroja L" first="Luana S." last="Maroja">Luana S. Maroja</name><affiliation><nlm:aff id="A26">Department of Biology, Williams College, Williamstown, MA 01267, USA.</nlm:aff></affiliation></author><author><name sortKey="Martin, Arnaud" sort="Martin, Arnaud" uniqKey="Martin A" first="Arnaud" last="Martin">Arnaud Martin</name><affiliation><nlm:aff id="A3">Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA.</nlm:aff></affiliation></author><author><name sortKey="Moxon, Simon" sort="Moxon, Simon" uniqKey="Moxon S" first="Simon" last="Moxon">Simon Moxon</name><affiliation><nlm:aff id="A27">Department of Genetics, Yale University School of Medicine, 333 Cedar Street, New Haven, CT 06520, USA.</nlm:aff></affiliation></author><author><name sortKey="Palmer, William J" sort="Palmer, William J" uniqKey="Palmer W" first="William J." last="Palmer">William J. Palmer</name><affiliation><nlm:aff id="A21">Department of Genetics, Downing Street, University of Cambridge, Cambridge, CB2 3EH, UK.</nlm:aff></affiliation></author><author><name sortKey="Papa, Riccardo" sort="Papa, Riccardo" uniqKey="Papa R" first="Riccardo" last="Papa">Riccardo Papa</name><affiliation><nlm:aff id="A28">Department of Biology and Center for Applied Tropical Ecology and Conservation, University of Puerto Rico, Rio Piedras, Puerto Rico.</nlm:aff></affiliation></author><author><name sortKey="Papanicolaou, Alexie" sort="Papanicolaou, Alexie" uniqKey="Papanicolaou A" first="Alexie" last="Papanicolaou">Alexie Papanicolaou</name><affiliation><nlm:aff id="A18">Black Mountain Laboratories, CSIRO Ecosystem Sciences, Clunies Ross Street, Canberra, ACT 2601, Australia.</nlm:aff></affiliation></author><author><name sortKey="Pauchet, Yannick" sort="Pauchet, Yannick" uniqKey="Pauchet Y" first="Yannick" last="Pauchet">Yannick Pauchet</name><affiliation><nlm:aff id="A11">Max Planck Institute for Chemical Ecology, D-07745 Jena, Germany.</nlm:aff></affiliation></author><author><name sortKey="Ray, David A" sort="Ray, David A" uniqKey="Ray D" first="David A." last="Ray">David A. Ray</name><affiliation><nlm:aff id="A29">Department of Biochemistry, Molecular Biology, Entomology and Plant Pathology, Mississippi State University, Mississippi State, Mississippi 39762, USA.</nlm:aff></affiliation><affiliation><nlm:aff id="A30">Institute for Genomics, Biocomputing and Biotechnology, Mississippi State University, Mississippi State, Mississippi 39759, USA.</nlm:aff></affiliation></author><author><name sortKey="Rosser, Neil" sort="Rosser, Neil" uniqKey="Rosser N" first="Neil" last="Rosser">Neil Rosser</name><affiliation><nlm:aff id="A1">Department of Genetics, Evolution and Environment, University College London, Gower Street, London, WC1E 6BT, UK.</nlm:aff></affiliation></author><author><name sortKey="Salzberg, Steven L" sort="Salzberg, Steven L" uniqKey="Salzberg S" first="Steven L." last="Salzberg">Steven L. Salzberg</name><affiliation><nlm:aff id="A31">McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University, Baltimore, MD 21205, USA.</nlm:aff></affiliation></author><author><name sortKey="Supple, Megan A" sort="Supple, Megan A" uniqKey="Supple M" first="Megan A." last="Supple">Megan A. Supple</name><affiliation><nlm:aff id="A32">Biomathematics Program, North Carolina State University, Raleigh, NC 27695, USA.</nlm:aff></affiliation></author><author><name sortKey="Surridge, Alison" sort="Surridge, Alison" uniqKey="Surridge A" first="Alison" last="Surridge">Alison Surridge</name><affiliation><nlm:aff id="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, UK.</nlm:aff></affiliation></author><author><name sortKey="Tenger Trolander, Ayse" sort="Tenger Trolander, Ayse" uniqKey="Tenger Trolander A" first="Ayse" last="Tenger-Trolander">Ayse Tenger-Trolander</name><affiliation><nlm:aff id="A13">FAS Center for Systems Biology, Harvard University, Cambridge, MA 02138, USA.</nlm:aff></affiliation></author><author><name sortKey="Vogel, Heiko" sort="Vogel, Heiko" uniqKey="Vogel H" first="Heiko" last="Vogel">Heiko Vogel</name><affiliation><nlm:aff id="A11">Max Planck Institute for Chemical Ecology, D-07745 Jena, Germany.</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="A33">School of Biological Sciences, University of Bristol, Bristol, BS8 1UG, UK.</nlm:aff></affiliation></author><author><name sortKey="Wilson, Derek" sort="Wilson, Derek" uniqKey="Wilson D" first="Derek" last="Wilson">Derek Wilson</name><affiliation><nlm:aff id="A7">European Bioinformatics Institute, Hinxton, CB10 1SD, UK.</nlm:aff></affiliation></author><author><name sortKey="Yorke, James A" sort="Yorke, James A" uniqKey="Yorke J" first="James A." last="Yorke">James A. Yorke</name><affiliation><nlm:aff id="A6">Institute for Physical Science and Technology, University of Maryland, College Park, MD 20742, USA.</nlm:aff></affiliation></author><author><name sortKey="Yuan, Furong" sort="Yuan, Furong" uniqKey="Yuan F" first="Furong" last="Yuan">Furong Yuan</name><affiliation><nlm:aff id="A3">Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA.</nlm:aff></affiliation></author><author><name sortKey="Balmuth, Alexi L" sort="Balmuth, Alexi L" uniqKey="Balmuth A" first="Alexi L." last="Balmuth">Alexi L. Balmuth</name><affiliation><nlm:aff id="A34">The GenePool, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation></author><author><name sortKey="Eland, Cathlene" sort="Eland, Cathlene" uniqKey="Eland C" first="Cathlene" last="Eland">Cathlene Eland</name><affiliation><nlm:aff id="A34">The GenePool, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation></author><author><name sortKey="Gharbi, Karim" sort="Gharbi, Karim" uniqKey="Gharbi K" first="Karim" last="Gharbi">Karim Gharbi</name><affiliation><nlm:aff id="A34">The GenePool, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation></author><author><name sortKey="Thomson, Marian" sort="Thomson, Marian" uniqKey="Thomson M" first="Marian" last="Thomson">Marian Thomson</name><affiliation><nlm:aff id="A34">The GenePool, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation></author><author><name sortKey="Gibbs, Richard A" sort="Gibbs, Richard A" uniqKey="Gibbs R" first="Richard A." last="Gibbs">Richard A. Gibbs</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Han, Yi" sort="Han, Yi" uniqKey="Han Y" first="Yi" last="Han">Yi Han</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Jayaseelan, Joy C" sort="Jayaseelan, Joy C" uniqKey="Jayaseelan J" first="Joy C." last="Jayaseelan">Joy C. Jayaseelan</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Kovar, Christie" sort="Kovar, Christie" uniqKey="Kovar C" first="Christie" last="Kovar">Christie Kovar</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Mathew, Tittu" sort="Mathew, Tittu" uniqKey="Mathew T" first="Tittu" last="Mathew">Tittu Mathew</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Muzny, Donna M" sort="Muzny, Donna M" uniqKey="Muzny D" first="Donna M." last="Muzny">Donna M. Muzny</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Ongeri, Fiona" sort="Ongeri, Fiona" uniqKey="Ongeri F" first="Fiona" last="Ongeri">Fiona Ongeri</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Pu, Ling Ling" sort="Pu, Ling Ling" uniqKey="Pu L" first="Ling-Ling" last="Pu">Ling-Ling Pu</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Qu, Jiaxin" sort="Qu, Jiaxin" uniqKey="Qu J" first="Jiaxin" last="Qu">Jiaxin Qu</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Thornton, Rebecca L" sort="Thornton, Rebecca L" uniqKey="Thornton R" first="Rebecca L." last="Thornton">Rebecca L. Thornton</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Worley, Kim C" sort="Worley, Kim C" uniqKey="Worley K" first="Kim C." last="Worley">Kim C. Worley</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Wu, Yuan Qing" sort="Wu, Yuan Qing" uniqKey="Wu Y" first="Yuan-Qing" last="Wu">Yuan-Qing Wu</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Linares, Mauricio" sort="Linares, Mauricio" uniqKey="Linares M" first="Mauricio" last="Linares">Mauricio Linares</name><affiliation><nlm:aff id="A36">Facultad de Ciencias Naturales y Matemáticas, Universidad del Rosario and Instituto de Genética, Universidad de los Andes, Bogotá, Colombia.</nlm:aff></affiliation></author><author><name sortKey="Blaxter, Mark L" sort="Blaxter, Mark L" uniqKey="Blaxter M" first="Mark L." last="Blaxter">Mark L. Blaxter</name><affiliation><nlm:aff id="A4">Institute of Evolutionary Biology, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation><affiliation><nlm:aff id="A34">The GenePool, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation></author><author><name sortKey="Constant, Richard H Ffrench" sort="Constant, Richard H Ffrench" uniqKey="Constant R" first="Richard H. Ffrench" last="Constant">Richard H. Ffrench Constant</name><affiliation><nlm:aff id="A14">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Penryn, TR10 9EZ, UK.</nlm:aff></affiliation></author><author><name sortKey="Joron, Mathieu" sort="Joron, Mathieu" uniqKey="Joron M" first="Mathieu" last="Joron">Mathieu Joron</name><affiliation><nlm:aff id="A5">CNRS UMR 7205, Muséum National d’Histoire Naturelle, 45 rue Buffon, Paris 75005, France.</nlm:aff></affiliation></author><author><name sortKey="Kronforst, Marcus R" sort="Kronforst, Marcus R" uniqKey="Kronforst M" first="Marcus R." last="Kronforst">Marcus R. Kronforst</name><affiliation><nlm:aff id="A13">FAS Center for Systems Biology, Harvard University, Cambridge, MA 02138, USA.</nlm:aff></affiliation></author><author><name sortKey="Mullen, Sean P" sort="Mullen, Sean P" uniqKey="Mullen S" first="Sean P." last="Mullen">Sean P. Mullen</name><affiliation><nlm:aff id="A37">Department of Biology, Boston University, 5 Cummington Street, Boston, MA 02215, USA.</nlm:aff></affiliation></author><author><name sortKey="Reed, Robert D" sort="Reed, Robert D" uniqKey="Reed R" first="Robert D." last="Reed">Robert D. Reed</name><affiliation><nlm:aff id="A3">Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA.</nlm:aff></affiliation></author><author><name sortKey="Scherer, Steven E" sort="Scherer, Steven E" uniqKey="Scherer S" first="Steven E." last="Scherer">Steven E. Scherer</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Richards, Stephen" sort="Richards, Stephen" uniqKey="Richards S" first="Stephen" last="Richards">Stephen Richards</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Mallet, James" sort="Mallet, James" uniqKey="Mallet J" first="James" last="Mallet">James Mallet</name><affiliation><nlm:aff id="A1">Department of Genetics, Evolution and Environment, University College London, Gower Street, London, WC1E 6BT, UK.</nlm:aff></affiliation><affiliation><nlm:aff id="A38">Department of Organismic and Evolutionary Biology, Harvard University, 16 Divinity Avenue, Cambridge, MA 02138, USA.</nlm:aff></affiliation></author><author><name sortKey="Mcmillan, W Owen" sort="Mcmillan, W Owen" uniqKey="Mcmillan W" first="W. Owen" last="Mcmillan">W. Owen Mcmillan</name><affiliation><nlm:aff id="A9">Smithsonian Tropical Research Institute, Naos Island, Causeway Amador, Panamá, Republic de Panamá.</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="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, UK.</nlm:aff></affiliation><affiliation><nlm:aff id="A9">Smithsonian Tropical Research Institute, Naos Island, Causeway Amador, Panamá, Republic de Panamá.</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">22722851</idno><idno type="pmc">3398145</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3398145</idno><idno type="RBID">PMC:3398145</idno><idno type="doi">10.1038/nature11041</idno><date when="2012">2012</date><idno type="wicri:Area/Pmc/Corpus">002D61</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">002D61</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Butterfly genome reveals promiscuous exchange of mimicry adaptations among species</title><author><name sortKey="Dasmahapatra, Kanchon K" sort="Dasmahapatra, Kanchon K" uniqKey="Dasmahapatra K" first="Kanchon K" last="Dasmahapatra">Kanchon K. Dasmahapatra</name><affiliation><nlm:aff id="A1">Department of Genetics, Evolution and Environment, University College London, Gower Street, London, WC1E 6BT, UK.</nlm:aff></affiliation></author><author><name sortKey="Walters, James R" sort="Walters, James R" uniqKey="Walters J" first="James R." last="Walters">James R. Walters</name><affiliation><nlm:aff id="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, UK.</nlm:aff></affiliation></author><author><name sortKey="Briscoe, Adriana D" sort="Briscoe, Adriana D" uniqKey="Briscoe A" first="Adriana D." last="Briscoe">Adriana D. Briscoe</name><affiliation><nlm:aff id="A3">Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA.</nlm:aff></affiliation></author><author><name sortKey="Davey, John W" sort="Davey, John W" uniqKey="Davey J" first="John W." last="Davey">John W. Davey</name><affiliation><nlm:aff id="A4">Institute of Evolutionary Biology, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</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="A5">CNRS UMR 7205, Muséum National d’Histoire Naturelle, 45 rue Buffon, Paris 75005, France.</nlm:aff></affiliation></author><author><name sortKey="Nadeau, Nicola J" sort="Nadeau, Nicola J" uniqKey="Nadeau N" first="Nicola J." last="Nadeau">Nicola J. Nadeau</name><affiliation><nlm:aff id="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, UK.</nlm:aff></affiliation></author><author><name sortKey="Zimin, Aleksey V" sort="Zimin, Aleksey V" uniqKey="Zimin A" first="Aleksey V." last="Zimin">Aleksey V. Zimin</name><affiliation><nlm:aff id="A6">Institute for Physical Science and Technology, University of Maryland, College Park, MD 20742, USA.</nlm:aff></affiliation></author><author><name sortKey="Hughes, Daniel S T" sort="Hughes, Daniel S T" uniqKey="Hughes D" first="Daniel S. T." last="Hughes">Daniel S. T. Hughes</name><affiliation><nlm:aff id="A7">European Bioinformatics Institute, Hinxton, CB10 1SD, UK.</nlm:aff></affiliation></author><author><name sortKey="Ferguson, Laura C" sort="Ferguson, Laura C" uniqKey="Ferguson L" first="Laura C." last="Ferguson">Laura C. Ferguson</name><affiliation><nlm:aff id="A8">Department of Zoology, University of Oxford, South Parks Road, Oxford, OX1 3PS, UK.</nlm:aff></affiliation></author><author><name sortKey="Martin, Simon H" sort="Martin, Simon H" uniqKey="Martin S" first="Simon H." last="Martin">Simon H. Martin</name><affiliation><nlm:aff id="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, 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="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, UK.</nlm:aff></affiliation><affiliation><nlm:aff id="A9">Smithsonian Tropical Research Institute, Naos Island, Causeway Amador, Panamá, Republic de Panamá.</nlm:aff></affiliation></author><author><name sortKey="Lewis, James J" sort="Lewis, James J" uniqKey="Lewis J" first="James J." last="Lewis">James J. Lewis</name><affiliation><nlm:aff id="A3">Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA.</nlm:aff></affiliation></author><author><name sortKey="Adler, Sebastian" sort="Adler, Sebastian" uniqKey="Adler S" first="Sebastian" last="Adler">Sebastian Adler</name><affiliation><nlm:aff id="A10">Institut für Mathematik und Informatik, Universität Greifswald, 17487 Greifswald, Germany.</nlm:aff></affiliation></author><author><name sortKey="Ahn, Seung Joon" sort="Ahn, Seung Joon" uniqKey="Ahn S" first="Seung-Joon" last="Ahn">Seung-Joon Ahn</name><affiliation><nlm:aff id="A11">Max Planck Institute for Chemical Ecology, D-07745 Jena, Germany.</nlm:aff></affiliation></author><author><name sortKey="Baker, Dean A" sort="Baker, Dean A" uniqKey="Baker D" first="Dean A." last="Baker">Dean A. Baker</name><affiliation><nlm:aff id="A12">Ecology and Evolution, Imperial College London, London, 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="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, 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="A13">FAS Center for Systems Biology, Harvard University, Cambridge, MA 02138, USA.</nlm:aff></affiliation></author><author><name sortKey="Chauhan, Ritika" sort="Chauhan, Ritika" uniqKey="Chauhan R" first="Ritika" last="Chauhan">Ritika Chauhan</name><affiliation><nlm:aff id="A14">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Penryn, TR10 9EZ, UK.</nlm:aff></affiliation></author><author><name sortKey="Counterman, Brian A" sort="Counterman, Brian A" uniqKey="Counterman B" first="Brian A." last="Counterman">Brian A. Counterman</name><affiliation><nlm:aff id="A15">Department of Biology, Mississippi State University, Mississippi State, Mississippi 39762, USA.</nlm:aff></affiliation></author><author><name sortKey="Dalmay, Tamas" sort="Dalmay, Tamas" uniqKey="Dalmay T" first="Tamas" last="Dalmay">Tamas Dalmay</name><affiliation><nlm:aff id="A16">School of Biological Sciences, University of East Anglia, Norwich Research Park, Norwich, NR4 7TJ, UK.</nlm:aff></affiliation></author><author><name sortKey="Gilbert, Lawrence E" sort="Gilbert, Lawrence E" uniqKey="Gilbert L" first="Lawrence E." last="Gilbert">Lawrence E. Gilbert</name><affiliation><nlm:aff id="A17">Section of Integrative Biology and Brackenridge Field Laboratory, University of Texas, Austin, TX 78712, USA.</nlm:aff></affiliation></author><author><name sortKey="Gordon, Karl" sort="Gordon, Karl" uniqKey="Gordon K" first="Karl" last="Gordon">Karl Gordon</name><affiliation><nlm:aff id="A18">Black Mountain Laboratories, CSIRO Ecosystem Sciences, Clunies Ross Street, Canberra, ACT 2601, Australia.</nlm:aff></affiliation></author><author><name sortKey="Heckel, David G" sort="Heckel, David G" uniqKey="Heckel D" first="David G." last="Heckel">David G. Heckel</name><affiliation><nlm:aff id="A11">Max Planck Institute for Chemical Ecology, D-07745 Jena, Germany.</nlm:aff></affiliation></author><author><name sortKey="Hines, Heather M" sort="Hines, Heather M" uniqKey="Hines H" first="Heather M." last="Hines">Heather M. Hines</name><affiliation><nlm:aff id="A19">Department of Genetics, North Carolina State University, Raleigh, NC 27695, USA.</nlm:aff></affiliation></author><author><name sortKey="Hoff, Katharina J" sort="Hoff, Katharina J" uniqKey="Hoff K" first="Katharina J." last="Hoff">Katharina J. Hoff</name><affiliation><nlm:aff id="A10">Institut für Mathematik und Informatik, Universität Greifswald, 17487 Greifswald, Germany.</nlm:aff></affiliation></author><author><name sortKey="Holland, Peter W H" sort="Holland, Peter W H" uniqKey="Holland P" first="Peter W. H." last="Holland">Peter W. H. Holland</name><affiliation><nlm:aff id="A8">Department of Zoology, University of Oxford, South Parks Road, Oxford, OX1 3PS, UK.</nlm:aff></affiliation></author><author><name sortKey="Jacquin Joly, Emmanuelle" sort="Jacquin Joly, Emmanuelle" uniqKey="Jacquin Joly E" first="Emmanuelle" last="Jacquin-Joly">Emmanuelle Jacquin-Joly</name><affiliation><nlm:aff id="A20">Inra, UMR-A 1272 Inra-Université Pierre et Marie Curie, Physiologie de l’Insecte: Signalisation et Communication, Route de Saint-Cyr, Versailles, France.</nlm:aff></affiliation></author><author><name sortKey="Jiggins, Francis M" sort="Jiggins, Francis M" uniqKey="Jiggins F" first="Francis M." last="Jiggins">Francis M. Jiggins</name><affiliation><nlm:aff id="A21">Department of Genetics, Downing Street, University of Cambridge, Cambridge, CB2 3EH, UK.</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="A5">CNRS UMR 7205, Muséum National d’Histoire Naturelle, 45 rue Buffon, Paris 75005, France.</nlm:aff></affiliation></author><author><name sortKey="Kapan, Durrell D" sort="Kapan, Durrell D" uniqKey="Kapan D" first="Durrell D." last="Kapan">Durrell D. Kapan</name><affiliation><nlm:aff id="A22">Department of Entomology, Center for Comparative Genomics, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, California 94118, USA.</nlm:aff></affiliation><affiliation><nlm:aff id="A23">Center for Conservation and Research Training, Pacific Biosciences Research Center, University of Hawaii at Manoa, 3050 Maile Way, Gilmore 406, Honolulu, HI 96822, USA.</nlm:aff></affiliation></author><author><name sortKey="Kersey, Paul" sort="Kersey, Paul" uniqKey="Kersey P" first="Paul" last="Kersey">Paul Kersey</name><affiliation><nlm:aff id="A7">European Bioinformatics Institute, Hinxton, CB10 1SD, UK.</nlm:aff></affiliation></author><author><name sortKey="Lamas, Gerardo" sort="Lamas, Gerardo" uniqKey="Lamas G" first="Gerardo" last="Lamas">Gerardo Lamas</name><affiliation><nlm:aff id="A24">Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Apartado 14-0434, Lima, Peru.</nlm:aff></affiliation></author><author><name sortKey="Lawson, Daniel" sort="Lawson, Daniel" uniqKey="Lawson D" first="Daniel" last="Lawson">Daniel Lawson</name><affiliation><nlm:aff id="A7">European Bioinformatics Institute, Hinxton, CB10 1SD, UK.</nlm:aff></affiliation></author><author><name sortKey="Mapleson, Daniel" sort="Mapleson, Daniel" uniqKey="Mapleson D" first="Daniel" last="Mapleson">Daniel Mapleson</name><affiliation><nlm:aff id="A25">School of Computing Sciences, University of East Anglia, Norwich Research Park, Norwich, NR4 7TJ, UK.</nlm:aff></affiliation></author><author><name sortKey="Maroja, Luana S" sort="Maroja, Luana S" uniqKey="Maroja L" first="Luana S." last="Maroja">Luana S. Maroja</name><affiliation><nlm:aff id="A26">Department of Biology, Williams College, Williamstown, MA 01267, USA.</nlm:aff></affiliation></author><author><name sortKey="Martin, Arnaud" sort="Martin, Arnaud" uniqKey="Martin A" first="Arnaud" last="Martin">Arnaud Martin</name><affiliation><nlm:aff id="A3">Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA.</nlm:aff></affiliation></author><author><name sortKey="Moxon, Simon" sort="Moxon, Simon" uniqKey="Moxon S" first="Simon" last="Moxon">Simon Moxon</name><affiliation><nlm:aff id="A27">Department of Genetics, Yale University School of Medicine, 333 Cedar Street, New Haven, CT 06520, USA.</nlm:aff></affiliation></author><author><name sortKey="Palmer, William J" sort="Palmer, William J" uniqKey="Palmer W" first="William J." last="Palmer">William J. Palmer</name><affiliation><nlm:aff id="A21">Department of Genetics, Downing Street, University of Cambridge, Cambridge, CB2 3EH, UK.</nlm:aff></affiliation></author><author><name sortKey="Papa, Riccardo" sort="Papa, Riccardo" uniqKey="Papa R" first="Riccardo" last="Papa">Riccardo Papa</name><affiliation><nlm:aff id="A28">Department of Biology and Center for Applied Tropical Ecology and Conservation, University of Puerto Rico, Rio Piedras, Puerto Rico.</nlm:aff></affiliation></author><author><name sortKey="Papanicolaou, Alexie" sort="Papanicolaou, Alexie" uniqKey="Papanicolaou A" first="Alexie" last="Papanicolaou">Alexie Papanicolaou</name><affiliation><nlm:aff id="A18">Black Mountain Laboratories, CSIRO Ecosystem Sciences, Clunies Ross Street, Canberra, ACT 2601, Australia.</nlm:aff></affiliation></author><author><name sortKey="Pauchet, Yannick" sort="Pauchet, Yannick" uniqKey="Pauchet Y" first="Yannick" last="Pauchet">Yannick Pauchet</name><affiliation><nlm:aff id="A11">Max Planck Institute for Chemical Ecology, D-07745 Jena, Germany.</nlm:aff></affiliation></author><author><name sortKey="Ray, David A" sort="Ray, David A" uniqKey="Ray D" first="David A." last="Ray">David A. Ray</name><affiliation><nlm:aff id="A29">Department of Biochemistry, Molecular Biology, Entomology and Plant Pathology, Mississippi State University, Mississippi State, Mississippi 39762, USA.</nlm:aff></affiliation><affiliation><nlm:aff id="A30">Institute for Genomics, Biocomputing and Biotechnology, Mississippi State University, Mississippi State, Mississippi 39759, USA.</nlm:aff></affiliation></author><author><name sortKey="Rosser, Neil" sort="Rosser, Neil" uniqKey="Rosser N" first="Neil" last="Rosser">Neil Rosser</name><affiliation><nlm:aff id="A1">Department of Genetics, Evolution and Environment, University College London, Gower Street, London, WC1E 6BT, UK.</nlm:aff></affiliation></author><author><name sortKey="Salzberg, Steven L" sort="Salzberg, Steven L" uniqKey="Salzberg S" first="Steven L." last="Salzberg">Steven L. Salzberg</name><affiliation><nlm:aff id="A31">McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University, Baltimore, MD 21205, USA.</nlm:aff></affiliation></author><author><name sortKey="Supple, Megan A" sort="Supple, Megan A" uniqKey="Supple M" first="Megan A." last="Supple">Megan A. Supple</name><affiliation><nlm:aff id="A32">Biomathematics Program, North Carolina State University, Raleigh, NC 27695, USA.</nlm:aff></affiliation></author><author><name sortKey="Surridge, Alison" sort="Surridge, Alison" uniqKey="Surridge A" first="Alison" last="Surridge">Alison Surridge</name><affiliation><nlm:aff id="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, UK.</nlm:aff></affiliation></author><author><name sortKey="Tenger Trolander, Ayse" sort="Tenger Trolander, Ayse" uniqKey="Tenger Trolander A" first="Ayse" last="Tenger-Trolander">Ayse Tenger-Trolander</name><affiliation><nlm:aff id="A13">FAS Center for Systems Biology, Harvard University, Cambridge, MA 02138, USA.</nlm:aff></affiliation></author><author><name sortKey="Vogel, Heiko" sort="Vogel, Heiko" uniqKey="Vogel H" first="Heiko" last="Vogel">Heiko Vogel</name><affiliation><nlm:aff id="A11">Max Planck Institute for Chemical Ecology, D-07745 Jena, Germany.</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="A33">School of Biological Sciences, University of Bristol, Bristol, BS8 1UG, UK.</nlm:aff></affiliation></author><author><name sortKey="Wilson, Derek" sort="Wilson, Derek" uniqKey="Wilson D" first="Derek" last="Wilson">Derek Wilson</name><affiliation><nlm:aff id="A7">European Bioinformatics Institute, Hinxton, CB10 1SD, UK.</nlm:aff></affiliation></author><author><name sortKey="Yorke, James A" sort="Yorke, James A" uniqKey="Yorke J" first="James A." last="Yorke">James A. Yorke</name><affiliation><nlm:aff id="A6">Institute for Physical Science and Technology, University of Maryland, College Park, MD 20742, USA.</nlm:aff></affiliation></author><author><name sortKey="Yuan, Furong" sort="Yuan, Furong" uniqKey="Yuan F" first="Furong" last="Yuan">Furong Yuan</name><affiliation><nlm:aff id="A3">Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA.</nlm:aff></affiliation></author><author><name sortKey="Balmuth, Alexi L" sort="Balmuth, Alexi L" uniqKey="Balmuth A" first="Alexi L." last="Balmuth">Alexi L. Balmuth</name><affiliation><nlm:aff id="A34">The GenePool, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation></author><author><name sortKey="Eland, Cathlene" sort="Eland, Cathlene" uniqKey="Eland C" first="Cathlene" last="Eland">Cathlene Eland</name><affiliation><nlm:aff id="A34">The GenePool, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation></author><author><name sortKey="Gharbi, Karim" sort="Gharbi, Karim" uniqKey="Gharbi K" first="Karim" last="Gharbi">Karim Gharbi</name><affiliation><nlm:aff id="A34">The GenePool, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation></author><author><name sortKey="Thomson, Marian" sort="Thomson, Marian" uniqKey="Thomson M" first="Marian" last="Thomson">Marian Thomson</name><affiliation><nlm:aff id="A34">The GenePool, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation></author><author><name sortKey="Gibbs, Richard A" sort="Gibbs, Richard A" uniqKey="Gibbs R" first="Richard A." last="Gibbs">Richard A. Gibbs</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Han, Yi" sort="Han, Yi" uniqKey="Han Y" first="Yi" last="Han">Yi Han</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Jayaseelan, Joy C" sort="Jayaseelan, Joy C" uniqKey="Jayaseelan J" first="Joy C." last="Jayaseelan">Joy C. Jayaseelan</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Kovar, Christie" sort="Kovar, Christie" uniqKey="Kovar C" first="Christie" last="Kovar">Christie Kovar</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Mathew, Tittu" sort="Mathew, Tittu" uniqKey="Mathew T" first="Tittu" last="Mathew">Tittu Mathew</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Muzny, Donna M" sort="Muzny, Donna M" uniqKey="Muzny D" first="Donna M." last="Muzny">Donna M. Muzny</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Ongeri, Fiona" sort="Ongeri, Fiona" uniqKey="Ongeri F" first="Fiona" last="Ongeri">Fiona Ongeri</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Pu, Ling Ling" sort="Pu, Ling Ling" uniqKey="Pu L" first="Ling-Ling" last="Pu">Ling-Ling Pu</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Qu, Jiaxin" sort="Qu, Jiaxin" uniqKey="Qu J" first="Jiaxin" last="Qu">Jiaxin Qu</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Thornton, Rebecca L" sort="Thornton, Rebecca L" uniqKey="Thornton R" first="Rebecca L." last="Thornton">Rebecca L. Thornton</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Worley, Kim C" sort="Worley, Kim C" uniqKey="Worley K" first="Kim C." last="Worley">Kim C. Worley</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Wu, Yuan Qing" sort="Wu, Yuan Qing" uniqKey="Wu Y" first="Yuan-Qing" last="Wu">Yuan-Qing Wu</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Linares, Mauricio" sort="Linares, Mauricio" uniqKey="Linares M" first="Mauricio" last="Linares">Mauricio Linares</name><affiliation><nlm:aff id="A36">Facultad de Ciencias Naturales y Matemáticas, Universidad del Rosario and Instituto de Genética, Universidad de los Andes, Bogotá, Colombia.</nlm:aff></affiliation></author><author><name sortKey="Blaxter, Mark L" sort="Blaxter, Mark L" uniqKey="Blaxter M" first="Mark L." last="Blaxter">Mark L. Blaxter</name><affiliation><nlm:aff id="A4">Institute of Evolutionary Biology, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation><affiliation><nlm:aff id="A34">The GenePool, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</nlm:aff></affiliation></author><author><name sortKey="Constant, Richard H Ffrench" sort="Constant, Richard H Ffrench" uniqKey="Constant R" first="Richard H. Ffrench" last="Constant">Richard H. Ffrench Constant</name><affiliation><nlm:aff id="A14">Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Penryn, TR10 9EZ, UK.</nlm:aff></affiliation></author><author><name sortKey="Joron, Mathieu" sort="Joron, Mathieu" uniqKey="Joron M" first="Mathieu" last="Joron">Mathieu Joron</name><affiliation><nlm:aff id="A5">CNRS UMR 7205, Muséum National d’Histoire Naturelle, 45 rue Buffon, Paris 75005, France.</nlm:aff></affiliation></author><author><name sortKey="Kronforst, Marcus R" sort="Kronforst, Marcus R" uniqKey="Kronforst M" first="Marcus R." last="Kronforst">Marcus R. Kronforst</name><affiliation><nlm:aff id="A13">FAS Center for Systems Biology, Harvard University, Cambridge, MA 02138, USA.</nlm:aff></affiliation></author><author><name sortKey="Mullen, Sean P" sort="Mullen, Sean P" uniqKey="Mullen S" first="Sean P." last="Mullen">Sean P. Mullen</name><affiliation><nlm:aff id="A37">Department of Biology, Boston University, 5 Cummington Street, Boston, MA 02215, USA.</nlm:aff></affiliation></author><author><name sortKey="Reed, Robert D" sort="Reed, Robert D" uniqKey="Reed R" first="Robert D." last="Reed">Robert D. Reed</name><affiliation><nlm:aff id="A3">Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA.</nlm:aff></affiliation></author><author><name sortKey="Scherer, Steven E" sort="Scherer, Steven E" uniqKey="Scherer S" first="Steven E." last="Scherer">Steven E. Scherer</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Richards, Stephen" sort="Richards, Stephen" uniqKey="Richards S" first="Stephen" last="Richards">Stephen Richards</name><affiliation><nlm:aff id="A35">Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</nlm:aff></affiliation></author><author><name sortKey="Mallet, James" sort="Mallet, James" uniqKey="Mallet J" first="James" last="Mallet">James Mallet</name><affiliation><nlm:aff id="A1">Department of Genetics, Evolution and Environment, University College London, Gower Street, London, WC1E 6BT, UK.</nlm:aff></affiliation><affiliation><nlm:aff id="A38">Department of Organismic and Evolutionary Biology, Harvard University, 16 Divinity Avenue, Cambridge, MA 02138, USA.</nlm:aff></affiliation></author><author><name sortKey="Mcmillan, W Owen" sort="Mcmillan, W Owen" uniqKey="Mcmillan W" first="W. Owen" last="Mcmillan">W. Owen Mcmillan</name><affiliation><nlm:aff id="A9">Smithsonian Tropical Research Institute, Naos Island, Causeway Amador, Panamá, Republic de Panamá.</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="A2">Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, UK.</nlm:aff></affiliation><affiliation><nlm:aff id="A9">Smithsonian Tropical Research Institute, Naos Island, Causeway Amador, Panamá, Republic de Panamá.</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="2012">2012</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p id="P2">The evolutionary importance of hybridization and introgression has long been debated<sup><xref ref-type="bibr" rid="R1">1</xref></sup>. We used genomic tools to investigate introgression in <italic>Heliconius</italic>, a rapidly radiating genus of neotropical butterflies widely used in studies of ecology, behaviour, mimicry and speciation<sup><xref ref-type="bibr" rid="R2">2</xref>-<xref ref-type="bibr" rid="R5">5</xref></sup> . We sequenced the genome of <italic>Heliconius melpomene</italic> and compared it with other taxa to investigate chromosomal evolution in Lepidoptera and gene flow among multiple <italic>Heliconius</italic> species and races. Among 12,657 predicted genes for <italic>Heliconius</italic>, biologically important expansions of families of chemosensory and <italic>Hox</italic> genes are particularly noteworthy. Chromosomal organisation has remained broadly conserved since the Cretaceous, when butterflies split from the silkmoth lineage. Using genomic resequencing, we show hybrid exchange of genes between three co-mimics, <italic>H. melpomene</italic>, <italic>H. timareta</italic>, and <italic>H. elevatus</italic>, especially at two genomic regions that control mimicry pattern. Closely related <italic>Heliconius</italic> species clearly exchange protective colour pattern genes promiscuously, implying a major role for hybridization in adaptive radiation.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Seehausen, O" uniqKey="Seehausen O">O Seehausen</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Bates, Hw" uniqKey="Bates H">HW Bates</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Turner, Jrg" uniqKey="Turner J">JRG Turner</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Brown, Ks" uniqKey="Brown K">KS Brown</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Jiggins, Cd" uniqKey="Jiggins C">CD Jiggins</name></author><author><name sortKey="Naisbit, Re" uniqKey="Naisbit R">RE Naisbit</name></author><author><name sortKey="Coe, Rl" uniqKey="Coe R">RL Coe</name></author><author><name sortKey="Mallet, J" uniqKey="Mallet J">J Mallet</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Briscoe, Ad" uniqKey="Briscoe A">AD Briscoe</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Mallet, J" uniqKey="Mallet J">J Mallet</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kronforst, Mr" uniqKey="Kronforst M">MR Kronforst</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Salazar, C" uniqKey="Salazar C">C Salazar</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Pringle, Eg" uniqKey="Pringle E">EG Pringle</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Robinson, R" uniqKey="Robinson R">R Robinson</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Deng, Q" uniqKey="Deng Q">Q Deng</name></author><author><name sortKey="Zeng, Q" uniqKey="Zeng Q">Q Zeng</name></author><author><name sortKey="Qian, Y" uniqKey="Qian Y">Y Qian</name></author><author><name sortKey="Li, C" uniqKey="Li C">C Li</name></author><author><name sortKey="Yang, Y" uniqKey="Yang Y">Y Yang</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kemkemer, C" uniqKey="Kemkemer C">C Kemkemer</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="D Lencon, E" uniqKey="D Lencon E">E d’Alençon</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Vane Wright, Ri" uniqKey="Vane Wright R">RI Vane-Wright</name></author><author><name sortKey="Boppre, M" uniqKey="Boppre M">M Boppré</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Zhan, S" uniqKey="Zhan S">S Zhan</name></author><author><name sortKey="Merlin, C" uniqKey="Merlin C">C Merlin</name></author><author><name sortKey="Boore, Jl" uniqKey="Boore J">JL Boore</name></author><author><name sortKey="Reppert, Sm" uniqKey="Reppert S">SM Reppert</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Vieira, Fg" uniqKey="Vieira F">FG Vieira</name></author><author><name sortKey="Rozas, J" uniqKey="Rozas J">J Rozas</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Chai, Cl" uniqKey="Chai C">CL Chai</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Sackton, Tb" uniqKey="Sackton T">TB Sackton</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Green, Re" uniqKey="Green R">RE Green</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Durand, Ey" uniqKey="Durand E">EY Durand</name></author><author><name sortKey="Patterson, N" uniqKey="Patterson N">N Patterson</name></author><author><name sortKey="Reich, D" uniqKey="Reich D">D Reich</name></author><author><name sortKey="Slatkin, M" uniqKey="Slatkin M">M Slatkin</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Reed, Rd" uniqKey="Reed R">RD Reed</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Nadeau, Nj" uniqKey="Nadeau N">NJ Nadeau</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kim, M" uniqKey="Kim M">M Kim</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Song, Y" uniqKey="Song Y">Y Song</name></author></analytic></biblStruct></listBibl></div1></back></TEI><pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<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">22722851</article-id><article-id pub-id-type="pmc">3398145</article-id><article-id pub-id-type="doi">10.1038/nature11041</article-id><article-id pub-id-type="manuscript">UKMS47290</article-id><article-categories><subj-group subj-group-type="heading"><subject>Article</subject></subj-group></article-categories><title-group><article-title>Butterfly genome reveals promiscuous exchange of mimicry adaptations among species</article-title></title-group><contrib-group><contrib contrib-type="author"><collab>The <italic>Heliconius</italic> Genome Consortium</collab></contrib><contrib contrib-type="author"><name><surname>Dasmahapatra</surname><given-names>Kanchon K</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>Walters</surname><given-names>James R.</given-names></name><xref ref-type="aff" rid="A2">2</xref><xref ref-type="author-notes" rid="FN1">*</xref></contrib><contrib contrib-type="author"><name><surname>Briscoe</surname><given-names>Adriana D.</given-names></name><xref ref-type="aff" rid="A3">3</xref></contrib><contrib contrib-type="author"><name><surname>Davey</surname><given-names>John W.</given-names></name><xref ref-type="aff" rid="A4">4</xref></contrib><contrib contrib-type="author"><name><surname>Whibley</surname><given-names>Annabel</given-names></name><xref ref-type="aff" rid="A5">5</xref></contrib><contrib contrib-type="author"><name><surname>Nadeau</surname><given-names>Nicola J.</given-names></name><xref ref-type="aff" rid="A2">2</xref></contrib><contrib contrib-type="author"><name><surname>Zimin</surname><given-names>Aleksey V.</given-names></name><xref ref-type="aff" rid="A6">6</xref></contrib><contrib contrib-type="author"><name><surname>Hughes</surname><given-names>Daniel S. 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contrib-type="author"><name><surname>Han</surname><given-names>Yi</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Jayaseelan</surname><given-names>Joy C.</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Kovar</surname><given-names>Christie</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Mathew</surname><given-names>Tittu</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Muzny</surname><given-names>Donna M.</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Ongeri</surname><given-names>Fiona</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Pu</surname><given-names>Ling-Ling</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Qu</surname><given-names>Jiaxin</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Thornton</surname><given-names>Rebecca L.</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Worley</surname><given-names>Kim C.</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Wu</surname><given-names>Yuan-Qing</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Linares</surname><given-names>Mauricio</given-names></name><xref ref-type="aff" rid="A36">36</xref></contrib><contrib contrib-type="author"><name><surname>Blaxter</surname><given-names>Mark L.</given-names></name><xref ref-type="aff" rid="A4">4</xref><xref ref-type="aff" rid="A34">34</xref></contrib><contrib contrib-type="author"><name><surname>Constant</surname><given-names>Richard H. ffrench</given-names></name><xref ref-type="aff" rid="A14">14</xref></contrib><contrib contrib-type="author"><name><surname>Joron</surname><given-names>Mathieu</given-names></name><xref ref-type="aff" rid="A5">5</xref></contrib><contrib contrib-type="author"><name><surname>Kronforst</surname><given-names>Marcus R.</given-names></name><xref ref-type="aff" rid="A13">13</xref></contrib><contrib contrib-type="author"><name><surname>Mullen</surname><given-names>Sean P.</given-names></name><xref ref-type="aff" rid="A37">37</xref></contrib><contrib contrib-type="author"><name><surname>Reed</surname><given-names>Robert D.</given-names></name><xref ref-type="aff" rid="A3">3</xref></contrib><contrib contrib-type="author"><name><surname>Scherer</surname><given-names>Steven E.</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Richards</surname><given-names>Stephen</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Mallet</surname><given-names>James</given-names></name><xref ref-type="aff" rid="A1">1</xref><xref ref-type="aff" rid="A38">38</xref><xref ref-type="corresp" rid="CR1">§</xref></contrib><contrib contrib-type="author"><name><surname>McMillan</surname><given-names>W. Owen</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="A2">2</xref><xref ref-type="aff" rid="A9">9</xref></contrib></contrib-group><aff id="A1"><label>1</label>Department of Genetics, Evolution and Environment, University College London, Gower Street, London, WC1E 6BT, UK.</aff><aff id="A2"><label>2</label>Department of Zoology, Downing Street, University of Cambridge, Cambridge, CB2 3EJ, UK.</aff><aff id="A3"><label>3</label>Department of Ecology and Evolutionary Biology, University of California, Irvine, CA 92697, USA.</aff><aff id="A4"><label>4</label>Institute of Evolutionary Biology, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</aff><aff id="A5"><label>5</label>CNRS UMR 7205, Muséum National d’Histoire Naturelle, 45 rue Buffon, Paris 75005, France.</aff><aff id="A6"><label>6</label>Institute for Physical Science and Technology, University of Maryland, College Park, MD 20742, USA.</aff><aff id="A7"><label>7</label>European Bioinformatics Institute, Hinxton, CB10 1SD, UK.</aff><aff id="A8"><label>8</label>Department of Zoology, University of Oxford, South Parks Road, Oxford, OX1 3PS, UK.</aff><aff id="A9"><label>9</label>Smithsonian Tropical Research Institute, Naos Island, Causeway Amador, Panamá, Republic de Panamá.</aff><aff id="A10"><label>10</label>Institut für Mathematik und Informatik, Universität Greifswald, 17487 Greifswald, Germany.</aff><aff id="A11"><label>11</label>Max Planck Institute for Chemical Ecology, D-07745 Jena, Germany.</aff><aff id="A12"><label>12</label>Ecology and Evolution, Imperial College London, London, UK.</aff><aff id="A13"><label>13</label>FAS Center for Systems Biology, Harvard University, Cambridge, MA 02138, USA.</aff><aff id="A14"><label>14</label>Centre for Ecology and Conservation, School of Biosciences, University of Exeter, Penryn, TR10 9EZ, UK.</aff><aff id="A15"><label>15</label>Department of Biology, Mississippi State University, Mississippi State, Mississippi 39762, USA.</aff><aff id="A16"><label>16</label>School of Biological Sciences, University of East Anglia, Norwich Research Park, Norwich, NR4 7TJ, UK.</aff><aff id="A17"><label>17</label>Section of Integrative Biology and Brackenridge Field Laboratory, University of Texas, Austin, TX 78712, USA.</aff><aff id="A18"><label>18</label>Black Mountain Laboratories, CSIRO Ecosystem Sciences, Clunies Ross Street, Canberra, ACT 2601, Australia.</aff><aff id="A19"><label>19</label>Department of Genetics, North Carolina State University, Raleigh, NC 27695, USA.</aff><aff id="A20"><label>20</label>Inra, UMR-A 1272 Inra-Université Pierre et Marie Curie, Physiologie de l’Insecte: Signalisation et Communication, Route de Saint-Cyr, Versailles, France.</aff><aff id="A21"><label>21</label>Department of Genetics, Downing Street, University of Cambridge, Cambridge, CB2 3EH, UK.</aff><aff id="A22"><label>22</label>Department of Entomology, Center for Comparative Genomics, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, California 94118, USA.</aff><aff id="A23"><label>23</label>Center for Conservation and Research Training, Pacific Biosciences Research Center, University of Hawaii at Manoa, 3050 Maile Way, Gilmore 406, Honolulu, HI 96822, USA.</aff><aff id="A24"><label>24</label>Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Apartado 14-0434, Lima, Peru.</aff><aff id="A25"><label>25</label>School of Computing Sciences, University of East Anglia, Norwich Research Park, Norwich, NR4 7TJ, UK.</aff><aff id="A26"><label>26</label>Department of Biology, Williams College, Williamstown, MA 01267, USA.</aff><aff id="A27"><label>27</label>Department of Genetics, Yale University School of Medicine, 333 Cedar Street, New Haven, CT 06520, USA.</aff><aff id="A28"><label>28</label>Department of Biology and Center for Applied Tropical Ecology and Conservation, University of Puerto Rico, Rio Piedras, Puerto Rico.</aff><aff id="A29"><label>29</label>Department of Biochemistry, Molecular Biology, Entomology and Plant Pathology, Mississippi State University, Mississippi State, Mississippi 39762, USA.</aff><aff id="A30"><label>30</label>Institute for Genomics, Biocomputing and Biotechnology, Mississippi State University, Mississippi State, Mississippi 39759, USA.</aff><aff id="A31"><label>31</label>McKusick-Nathans Institute of Genetic Medicine, Johns Hopkins University, Baltimore, MD 21205, USA.</aff><aff id="A32"><label>32</label>Biomathematics Program, North Carolina State University, Raleigh, NC 27695, USA.</aff><aff id="A33"><label>33</label>School of Biological Sciences, University of Bristol, Bristol, BS8 1UG, UK.</aff><aff id="A34"><label>34</label>The GenePool, Ashworth Laboratories, University of Edinburgh, West Mains Road, Edinburgh, EH9 3JT, UK.</aff><aff id="A35"><label>35</label>Human Genome Sequencing Center, Baylor College of Medicine, One Baylor Plaza, Houston, TX 77030, USA.</aff><aff id="A36"><label>36</label>Facultad de Ciencias Naturales y Matemáticas, Universidad del Rosario and Instituto de Genética, Universidad de los Andes, Bogotá, Colombia.</aff><aff id="A37"><label>37</label>Department of Biology, Boston University, 5 Cummington Street, Boston, MA 02215, USA.</aff><aff id="A38"><label>38</label>Department of Organismic and Evolutionary Biology, Harvard University, 16 Divinity Avenue, Cambridge, MA 02138, USA.</aff><author-notes><fn fn-type="equal" id="FN1"><label>*</label><p id="P1">These authors contributed equally to this work.</p></fn><corresp id="CR1"><label>§</label>Correspondence and requests for materials should be addressed to J.M. (<email>jmallet@oeb.harvard.edu</email>).</corresp></author-notes><pub-date pub-type="nihms-submitted"><day>21</day><month>3</month><year>2012</year></pub-date><pub-date pub-type="ppub"><day>5</day><month>7</month><year>2012</year></pub-date><pub-date pub-type="pmc-release"><day>05</day><month>1</month><year>2013</year></pub-date><volume>487</volume><issue>7405</issue><fpage>94</fpage><lpage>98</lpage><permissions><license><license-p>Users may view, print, copy, download and text and data- mine the content in such documents, for the purposes of academic research, subject always to the full Conditions of use:
<uri xlink:type="simple" xlink:href="http://www.nature.com/authors/editorial_policies/license.html#terms">http://www.nature.com/authors/editorial_policies/license.html#terms</uri></license-p></license></permissions><abstract><p id="P2">The evolutionary importance of hybridization and introgression has long been debated<sup><xref ref-type="bibr" rid="R1">1</xref></sup>. We used genomic tools to investigate introgression in <italic>Heliconius</italic>, a rapidly radiating genus of neotropical butterflies widely used in studies of ecology, behaviour, mimicry and speciation<sup><xref ref-type="bibr" rid="R2">2</xref>-<xref ref-type="bibr" rid="R5">5</xref></sup> . We sequenced the genome of <italic>Heliconius melpomene</italic> and compared it with other taxa to investigate chromosomal evolution in Lepidoptera and gene flow among multiple <italic>Heliconius</italic> species and races. Among 12,657 predicted genes for <italic>Heliconius</italic>, biologically important expansions of families of chemosensory and <italic>Hox</italic> genes are particularly noteworthy. Chromosomal organisation has remained broadly conserved since the Cretaceous, when butterflies split from the silkmoth lineage. Using genomic resequencing, we show hybrid exchange of genes between three co-mimics, <italic>H. melpomene</italic>, <italic>H. timareta</italic>, and <italic>H. elevatus</italic>, especially at two genomic regions that control mimicry pattern. Closely related <italic>Heliconius</italic> species clearly exchange protective colour pattern genes promiscuously, implying a major role for hybridization in adaptive radiation.</p></abstract></article-meta></front><body><p id="P3">The butterfly genus <italic>Heliconius</italic> (Nymphalidae: Heliconiinae) is associated with a suite of derived life-history and ecological traits, including pollen-feeding, extended life-span, augmented ultraviolet colour vision, ‘trap-lining’ foraging behavior, gregarious roosting and complex mating behaviours, and provides outstanding opportunities for genomic studies of adaptive radiation and speciation<sup><xref ref-type="bibr" rid="R4">4</xref>, <xref ref-type="bibr" rid="R6">6</xref></sup>. The genus is best known for the hundreds of different colour pattern races seen among its 43 species, with repeated examples of both convergent evolution among distantly related species and divergent evolution between closely related taxa<sup><xref ref-type="bibr" rid="R3">3</xref></sup>. Geographic mosaics of multiple colour pattern races, such as in <italic>Heliconius melpomene</italic> (<xref ref-type="fig" rid="F1">Fig. 1</xref>), converge to similar mosaics in other species, and this led to the hypothesis of mimicry<sup><xref ref-type="bibr" rid="R2">2</xref></sup>. <italic>Heliconius</italic> are unpalatable and Müllerian mimicry of warning colour patterns enables species to share the cost of educating predators<sup><xref ref-type="bibr" rid="R3">3</xref></sup>. Divergence in wing pattern is also associated with speciation and adaptive radiation due to a dual role in mimicry and mate selection<sup><xref ref-type="bibr" rid="R3">3</xref>, <xref ref-type="bibr" rid="R5">5</xref></sup>. A particularly recent radiation is the <italic>melpomene</italic>-silvaniform clade, where mimetic patterns often appear polyphyletic (<xref ref-type="fig" rid="F1">Fig. 1a</xref>). Most species in this clade occasionally hybridise in the wild with other clade members<sup><xref ref-type="bibr" rid="R7">7</xref></sup>. Gene genealogies at a small number of loci indicate introgression between species<sup><xref ref-type="bibr" rid="R8">8</xref></sup>, and one non-mimetic species, <italic>H. heurippa</italic>, has a hybrid origin<sup><xref ref-type="bibr" rid="R9">9</xref></sup>. Adaptive introgression of mimicry loci is therefore a plausible explanation for parallel evolution of multiple mimetic patterns in the <italic>melpomene</italic>-silvaniform clade.</p><p id="P4">A <italic>Heliconius melpomene melpomene</italic> stock from Darién, Panama (<xref ref-type="fig" rid="F1">Fig. 1</xref>) was inbred via five generations of sib mating. A single male was sequenced to 38x coverage (after quality filtering) using combined 454 and Illumina technologies (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 1</xref>-<xref ref-type="supplementary-material" rid="SD1">8</xref>). The complete draft genome assembly of 269 Mb consists of 3,807 scaffolds with an N50 of 277 kb and contains 12,657 predicted protein-coding genes. RAD linkage mapping was used to assign and order 83% of the sequenced genome onto the 21 chromosomes (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 4</xref>). These data permit a considerably improved genome-wide chromosomal synteny comparison with the silkmoth <italic>Bombyx mori</italic><sup><xref ref-type="bibr" rid="R10">10</xref>, <xref ref-type="bibr" rid="R11">11</xref></sup>. Using 6,010 orthologues identified between <italic>H. melpomene</italic> and <italic>B. mori</italic> we found that 11 of 21 <italic>H. melpomene</italic> linkage groups show homology to single <italic>B. mori</italic> chromosomes and ten linkage groups have major contributions from two <italic>B. mori</italic> chromosomes (<xref ref-type="fig" rid="F2">Fig. 2a</xref> and <xref ref-type="supplementary-material" rid="SD1">Supplementary Information 8</xref>), revealing several previously unidentified chromosomal fusions. These fusions on the <italic>Heliconius</italic> lineage most likely occurred after divergence from the sister genus <italic>Eueides</italic><sup><xref ref-type="bibr" rid="R4">4</xref></sup>, which has the lepidopteran modal karyotype of <italic>n</italic>=31<sup><xref ref-type="bibr" rid="R12">12</xref></sup>. Three chromosomal fusions are evident in <italic>Bombyx</italic> (<xref ref-type="fig" rid="F2">Fig. 2a</xref>, <italic>B. mori</italic> chromosomes 11, 23 and 24), as required for evolution of the <italic>Bombyx n</italic>=28 karyotype from the ancestral <italic>n</italic>=31 karyotype. <italic>Heliconius</italic> and <italic>Bombyx</italic> lineages diverged in the Cretaceous >100 MYA<sup><xref ref-type="bibr" rid="R11">11</xref></sup>, so the chromosomal structures of Lepidoptera genomes have remained highly conserved compared to those of flies or vertebrates<sup><xref ref-type="bibr" rid="R13">13</xref>, <xref ref-type="bibr" rid="R14">14</xref></sup>. In contrast, small-scale rearrangements were frequent. In the comparison with <italic>Bombyx</italic>, we estimate 0.05-0.13 breaks/Mb/MY, and with the Monarch butterfly, <italic>Danaus plexippus</italic>, 0.04-0.29 breaks/Mb/MY. Although lower than previously suggested for Lepidoptera<sup><xref ref-type="bibr" rid="R15">15</xref></sup>, these rates are comparable to <italic>Drosophila</italic> (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 8</xref>).</p><p id="P5">The origin of butterflies was associated with a switch from nocturnal to diurnal behaviour, and a corresponding increase in visual communication<sup><xref ref-type="bibr" rid="R16">16</xref></sup>. <italic>Heliconius</italic> have increased visual complexity through expression of a duplicate UV opsin<sup><xref ref-type="bibr" rid="R6">6</xref></sup>, in addition to the long wavelength, blue, and UV-sensitive opsins in <italic>Bombyx</italic>. We might therefore predict reduced complexity of olfactory genes, but in fact <italic>Heliconius</italic> and <italic>Danaus</italic><sup><xref ref-type="bibr" rid="R17">17</xref></sup> genomes have more chemosensory proteins (CSPs) than any other insect genome: 33 and 34 CSPs respectively (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 9</xref>), versus 24 in <italic>Bombyx</italic> and 3- 4 in <italic>Drosophila</italic><sup><xref ref-type="bibr" rid="R18">18</xref></sup>. Lineage-specific CSP expansions were evident in both <italic>Danaus</italic> and <italic>Heliconius</italic> (<xref ref-type="fig" rid="F2">Fig. 2b</xref>). In contrast, all three lepidopteran genomes possess similar numbers of odorant binding proteins and olfactory receptors (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 9</xref>). <italic>Hox</italic> genes are involved in body plan development and show strong conservation across animals. We identified four additional <italic>Hox</italic> genes located between the canonical <italic>Hox</italic> genes <italic>pb</italic> and <italic>zen</italic>, orthologous to <italic>shx</italic> genes in <italic>B. mori</italic> (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 10</xref>)<sup><xref ref-type="bibr" rid="R19">19</xref></sup>. These <italic>Hox</italic> gene duplications in the butterflies and <italic>Bombyx</italic> share a common origin, and are independent of the two tandem duplications known in dipterans (<italic>zen2</italic>, <italic>bcd</italic>). Immunity-related gene families are similar across all three lepidopterans (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 11</xref>), contrasting with extensive duplications and losses within dipterans<sup><xref ref-type="bibr" rid="R20">20</xref></sup>.</p><p id="P6">The <italic>Heliconius</italic> reference genome enabled rigorous tests for introgression among <italic>melpomene</italic>-silvaniform clade species. We used RAD resequencing to reconstruct a robust phylogenetic tree based on 84 individuals of <italic>H. melpomene</italic> and its relatives,sampling on average 12 Mb, or 4% of the genome (<xref ref-type="fig" rid="F1">Fig 1a</xref>, <xref ref-type="supplementary-material" rid="SD1">Supplementary Information 12</xref>, <xref ref-type="supplementary-material" rid="SD1">13</xref>, <xref ref-type="supplementary-material" rid="SD1">18</xref>). We then tested for introgression between the sympatric co-mimetic postman races of <italic>H. melpomene aglaope</italic> and <italic>H. timareta</italic> ssp. nov. (<xref ref-type="fig" rid="F1">Fig. 1</xref>) in Peru, employing ‘ABBA-BABA’ single nucleotide sites and Patterson’s <italic>D</italic>-statistics (<xref ref-type="fig" rid="F3">Fig. 3a</xref>), originally developed to test for admixture between Neanderthals and modern humans<sup><xref ref-type="bibr" rid="R21">21</xref>, <xref ref-type="bibr" rid="R22">22</xref></sup> (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 12</xref>). Genome-wide we found an excess of ABBA sites, giving a significantly positive Patterson’s <italic>D</italic> = 0.037 ± 0.003 (two tailed <italic>Z</italic>-test for <italic>D</italic> = 0, <italic>P</italic> = 1 × 10<sup>−40</sup>), indicating greater genome-wide introgression between the sympatric mimetic taxa <italic>H. m. amaryllis</italic> and <italic>H. timareta</italic> ssp. nov., than between <italic>H. m. aglaope</italic> and <italic>H. timareta</italic> ssp. nov., which do not overlap spatially (<xref ref-type="fig" rid="F1">Fig. 1b</xref>). These <italic>D</italic>-statistics yield an estimate of 2-5% of the genome exchanged<sup><xref ref-type="bibr" rid="R21">21</xref></sup> between the two taxa (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 12</xref>). Eleven of the 21 chromosomes have significantly positive <italic>D</italic>-statistics (<xref ref-type="fig" rid="F3">Fig. 3b</xref>,); interestingly, the strongest signals of introgressions were found on two chromosomes containing the known mimicry loci <italic>B</italic>/<italic>D</italic> and <italic>N</italic>/<italic>Yb</italic> (<xref ref-type="fig" rid="F3">Fig. 3b</xref>, <xref ref-type="supplementary-material" rid="SD1">Supplementary Information 15</xref>).</p><p id="P7">Perhaps the best known case of Müllerian mimicry is the geographic mosaic of ~30 bold postman and rayed colour pattern races of <italic>H. melpomene</italic> (<xref ref-type="fig" rid="F1">Fig. 1b</xref>, <xref ref-type="supplementary-material" rid="SD1">Supplementary Information 22</xref>), which mimic a near-identical colour pattern mosaic in <italic>H. erato</italic> (<xref ref-type="fig" rid="F1">Fig. 1a</xref>), among other <italic>Heliconius</italic>. Mimicry variation is generally controlled by a few loci with major effects. Mimetic pattern differences between the postman <italic>H. melpomene amaryllis</italic> and rayed <italic>H. melpomene aglaope</italic> races studied here (<xref ref-type="fig" rid="F1">Fig 1a</xref>) are controlled by the <italic>B</italic>/<italic>D</italic> (red pattern) and <italic>N</italic>/<italic>Yb</italic> (yellow pattern) loci<sup><xref ref-type="bibr" rid="R23">23</xref>, <xref ref-type="bibr" rid="R24">24</xref></sup>. These loci are located on the same two chromosomes showing the strongest <italic>D</italic>- statistics in our RAD analysis (<xref ref-type="fig" rid="F3">Fig. 3b</xref>). To test whether mimicry loci might be introgressed between co-mimetic <italic>H. timareta</italic> and <italic>H. melpomene</italic> (<xref ref-type="fig" rid="F1">Fig. 1a</xref>)<sup><xref ref-type="bibr" rid="R7">7</xref></sup>, we resequenced the colour pattern regions <italic>B</italic>/<italic>D</italic> (0.7 Mb) and <italic>N</italic>/<italic>Yb</italic> (1.2 Mb), and 1.8 Mb of unlinked regions across the genome, from both postman and ray-patterned <italic>H. melpomene</italic> and <italic>H. timareta</italic> from Peru and Colombia, and six silvaniform outgroup taxa (<xref ref-type="fig" rid="F1">Fig. 1a</xref>, <xref ref-type="supplementary-material" rid="SD1">Supplementary Information 12</xref>). To test for introgression at the <italic>B/D</italic> mimicry locus we compared rayed <italic>H. m. aglaope</italic> and postman <italic>H. m. amaryllis</italic> as the ingroup with postman <italic>H. timareta</italic> ssp. nov. (as in <xref ref-type="fig" rid="F3">Fig. 3a</xref>) and found large, significant peaks of shared fixed ABBA nucleotide sites combined with an almost complete lack of BABA sites (<xref ref-type="fig" rid="F4">Fig. 4b</xref>). This provides evidence that blocks of shared sequence variation in the <italic>B</italic>/<italic>D</italic> region were exchanged between postman <italic>H. timareta</italic> and postman <italic>H. melpomene</italic>, in the genomic region known to determine red mimicry patterns between races of <italic>H. melpomene</italic><sup><xref ref-type="bibr" rid="R23">23</xref>, <xref ref-type="bibr" rid="R24">24</xref></sup> (<xref ref-type="fig" rid="F4">Fig. 4a</xref>).</p><p id="P8">For a reciprocal test, we used the same <italic>H. melpomene</italic> races as the ingroup to compare with rayed <italic>H. timareta florencia</italic> at the <italic>B</italic>/<italic>D</italic> region. In this case, correspondingly large and significant peaks of BABA nucleotide sites are accompanied by virtual absence of ABBA sites (<xref ref-type="fig" rid="F4">Fig. 4c</xref>) indicating that variation at the same mimicry locus was also shared between rayed <italic>H. timareta</italic> and rayed <italic>H. melpomene</italic>. Equivalent results in the <italic>N</italic>/<italic>Yb</italic> colour pattern region, controlling yellow colour pattern differences, are in the expected directions for introgression and highly significant for the test using postman <italic>H. timareta</italic> ssp. nov. (<italic>P</italic> = 6 × 10<sup>−34</sup>), although not significant with rayed <italic>H. timareta florencia</italic> (<italic>P</italic> = 0.13, <xref ref-type="supplementary-material" rid="SD1">Supplementary Information 17</xref>). In contrast hardly any ABBA or BABA sites are present in either comparison across 1.8 Mb in 55 genomic scaffolds unlinked to the colour pattern regions (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 21</xref>). These concordant, but reciprocal patterns, where fixed ABBA and BABA substitutions occur almost exclusively within large genomic blocks at two different colour pattern loci (449 and 99 sites for <italic>B</italic>/<italic>D</italic> and <italic>N</italic>/<italic>Yb</italic> respectively, <xref ref-type="fig" rid="F4">Figs. 4b,c</xref> and <xref ref-type="supplementary-material" rid="SD1">Supplementary Information 17</xref>) would be very hard to explain via convergent functional site evolution or under coalescent fluctuations. Instead, our results imply that derived colour pattern elements have introgressed recently between both rayed and postman forms of <italic>H. timareta</italic> and <italic>H. melpomene</italic>.</p><p id="P9">To test whether colour pattern loci might be shared more broadly across the clade, we used sliding-window phylogenetic analyses along the colour pattern regions. For regions flanking and unlinked to colour pattern loci, tree topologies are similar to the overriding signal recovered from the genome as a whole (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 18</xref>). Races of <italic>H. melpomene</italic> and <italic>H. timareta</italic> each form separate monophyletic sister groups and both are separated from the more distantly related silvaniform species (<xref ref-type="fig" rid="F4">Fig. 4d</xref>). By contrast, within the region of peak ABBA/BABA differences, the topologies switch dramatically. Races of <italic>H. melpomene</italic> and <italic>H. timareta</italic> group according to wing pattern, while the species themselves become polyphyletic (<xref ref-type="fig" rid="F4">Figs. 4e,f</xref>, <xref ref-type="supplementary-material" rid="SD1">Supplementary Information 19</xref>, <xref ref-type="supplementary-material" rid="SD1">20</xref>). Remarkably, the rayed <italic>H. elevatus</italic>, a member of the silvaniform clade according to genome average relationships (<xref ref-type="fig" rid="F1">Fig. 1a</xref>, <xref ref-type="supplementary-material" rid="SD1">Supplementary Information 18</xref>), groups with rayed races of unrelated <italic>H. melpomene</italic> and <italic>H. timareta</italic> in small sections within both <italic>B</italic>/<italic>D</italic> and <italic>N</italic>/<italic>Yb</italic> colour pattern loci (<xref ref-type="fig" rid="F4">Fig. 4e</xref>, <xref ref-type="supplementary-material" rid="SD1">Supplementary Information 19</xref>, <xref ref-type="supplementary-material" rid="SD1">20</xref>). These results are again most readily explained by introgression and fixation of mimicry genes.</p><p id="P10">We have developed a <italic>de novo</italic> reference genome sequence that will facilitate evolutionary and ecological studies in this key group of butterflies. We have demonstrated repeated exchange of small (~100 kb) adaptive genome regions among multiple species in an adaptive radiation. Our genome-scale analysis provides considerably greater power than previous tests of introgression <sup><xref ref-type="bibr" rid="R8">8</xref>, <xref ref-type="bibr" rid="R25">25</xref>, <xref ref-type="bibr" rid="R26">26</xref></sup>. As with <italic>H. heurippa</italic><sup><xref ref-type="bibr" rid="R9">9</xref></sup>, our evidence suggests that <italic>H. elevatus</italic> was formed during a hybrid speciation event. The main genomic signal from this rayed species places it closest to <italic>H. pardalinus butleri</italic> (<xref ref-type="fig" rid="F1">Fig. 1a</xref>), but colour pattern genomic regions resemble those of rayed races of <italic>H. melpomene</italic> (<xref ref-type="fig" rid="F4">Fig. 4e</xref> and <xref ref-type="supplementary-material" rid="SD1">Supplementary Information 18</xref>-<xref ref-type="supplementary-material" rid="SD1">20</xref>). Colour pattern is important in mating behaviour in <italic>Heliconius</italic><sup><xref ref-type="bibr" rid="R5">5</xref></sup>, and the transfer of mimetic pattern may have enabled the divergent sibling species <italic>H. elevatus</italic> to coexist with <italic>H. pardalinus</italic> across the Amazon. Although it was long suspected that introgression might be important in adaptive radiation<sup><xref ref-type="bibr" rid="R1">1</xref></sup>, our results from the most diverse terrestrial biome on the planet suggest that adaptive introgression is more pervasive than previously realized.</p><sec sec-type="methods" id="S1"><title>Methods summary</title><p id="P11">A full description of methods can be found in the <xref ref-type="supplementary-material" rid="SD1">Supplementary Information</xref>.</p></sec><sec sec-type="supplementary-material" id="SM"><title>Supplementary Material</title><supplementary-material content-type="local-data" id="SD1"><label>1</label><media xlink:href="NIHMS47290-supplement-1.pdf" orientation="portrait" xlink:type="simple" id="d35e1708" position="anchor" mimetype="application" mime-subtype="pdf"></media><p id="P12"><bold>Supplementary Information</bold> is linked to the online version of the paper at <ext-link ext-link-type="uri" xlink:href="http://www.nature.com/nature">www.nature.com/nature</ext-link>.</p></supplementary-material></sec></body><back><ack id="S2"><title>Acknowledgements</title><p>We thank the governments of Colombia, Peru and Panama for permission to collect and export butterflies. Sequencing was funded by contributions from consortium members. We thank Moisés Abanto for assistance in raising the inbred line. Individual laboratories were funded by the Leverhulme Trust (CDJ), John Fell Fund and Christ Church College, Oxford (LCF), The Royal Society (MJ, CDJ), NSF (WOM, MK, RR, SM, ADB), NIH (MK, SLS, JY), CNRS (MJ), ERC (MJ, PWHH), Banco de la República and COLCIENCAS (ML), and the BBSRC (JM, CDJ, MB, R ff-C).</p></ack><fn-group><title>Author contributions</title><fn id="FN3"><p id="P14"><bold>Consortium leaders</bold>:</p><p id="P15">Chris D. Jiggins, W. Owen McMillan</p></fn><fn id="FN7"><p id="P16"><bold>Heliconius Genome Consortium PIs</bold>:</p><p id="P17">Richard ffrench-Constant, Marcus Kronforst, Mathieu Joron, James Mallet, Sean Mullen, Robert Reed, Mark Blaxter, Larry Gilbert, Mauricio Linares, Gerardo Lamas</p></fn><fn id="FN9"><p id="P18"><bold>Introgression study leader & corresponding author</bold>:</p><p id="P19">James Mallet</p></fn><fn id="FN10"><p id="P20"><bold>Lead investigators</bold>:</p><p id="P21">Kanchon Dasmahapatra, James Walters, Nicola Nadeau, Annabel Whibley, John Davey, Adriana Briscoe, Laura Ferguson, Daniel Hughes, Simon Martin, Camilo Salazar, James Lewis</p></fn><fn id="FN11"><p id="P22"><bold>Sequencing</bold>:</p><p id="P13">Stephen Richards, Steve Scherer, Alexi Balmuth, Marian Thomson, Karim Gharbi, Cathlene Eland, Mark Blaxter, Richard Gibbs, Yi Han, Joy Christina Jayaseelan, Christie Kovar, Tittu Mathew, Donna Marie Muzny, Fiona Ongeri, Ling-Ling Pu, Jiaxin Qu, Rebecca Lynn Thornton, Kim C. Worley, Yuan-Qing Wu</p></fn><fn id="FN12"><p id="P23"><bold>Assembly</bold>:</p><p id="P24">Aleksey Zimin, James Yorke, Steven Salzberg, Alexie Papanicolaou, Karl Gordon</p></fn><fn id="FN13"><p id="P25"><bold>RAD map and assembly verification</bold>:</p><p id="P50">John Davey, Simon Baxter, Mark Blaxter, Luana Maroja, Durrell D. Kapan, James Walters, Paul Wilkinson</p></fn><fn id="FN14"><p id="P26"><bold>Geographic distribution map</bold>:</p><p id="P27">Neil Rosser</p></fn><fn id="FN15"><p id="P28"><bold>Annotation</bold>:</p><p id="P29">James Walters, Daniel Hughes, Derek Wilson, Daniel Lawson, Katharina Hoff, Sebastian Adler, Paul Wilkinson</p></fn><fn id="FN16"><p id="P30"><bold>Genome browser and databases</bold>:</p><p id="P31">Daniel Hughes, James Lewis</p></fn><fn id="FN8"><p id="P32"><bold>Manual annotation and evolutionary analyses</bold>:</p><p id="P33"><bold>Olfactory proteins:</bold> Adriana Briscoe, Emmanuelle Jacquin-Joly, Furong Yuan</p><p id="P34"><bold>Hox genes:</bold> Laura Ferguson, Peter W. H. Holland, James Walters</p><p id="P35"><bold>Micro-RNAs:</bold> Alison Surridge, Tamas Dalmay, Daniel Mapleson, Simon Moxon</p><p id="P36"><bold>Immune genes:</bold> William Palmer, Francis Jiggins</p><p id="P37"><bold>P450 genes:</bold> Robert Jones and Ritika Chauhan</p><p id="P38"><bold>UGT genes:</bold> Heiko Vogel, Seung-Joon Ahn, David Heckel</p><p id="P39"><bold>Ribosomal Proteins:</bold> Yannick Pauchet</p><p id="P40"><bold>Manual annotation group:</bold> Simon Baxter, Mark Blaxter, Adriana Briscoe, Nicola Chamberlain, Brian Counterman, Laura Ferguson, Heather Hines, Chris Jiggins, Frank Jiggins, Mathieu Joron, Durrell Kapan, Marcus Kronforst, Jim Mallet, Arnaud Martin, Sean Mullen, Nicola Nadeau, William Palmer, Riccardo Papa, Megan Supple, Ayse Trolander, Annabel Whibley, Furong Yuan</p><p id="P41"><bold>Transposable elements:</bold> Brian Counterman, David Ray</p><p id="P42"><bold>Orthologue predictions:</bold> Dean Baker</p><p id="P43"><bold>Synteny:</bold> Annabel Whibley, John Davey, David Heckel, Karl Gordon</p><p id="P44"><bold>Introgression analysis:</bold> Kanchon Dasmahapatra, Nicola Nadeau, John Davey, Simon Martin, Camilo Salazar, Chris Jiggins, Mathieu Joron, James Mallet</p></fn></fn-group><fn-group><fn id="FN4"><p id="P45"><bold>Author Information</bold> The genome sequence has been submitted to European Nucleotide Archive with accession numbers HE667773-HE672081. The annotated genome is available on our genome browser at <ext-link ext-link-type="uri" xlink:href="http://butterflygenome.org/">http://butterflygenome.org/</ext-link> and will also be made available in the next release of ENSEMBL Genomes. Additional short read sequences have been submitted to the European Nucleotide Archive with accession numbers ERP000993 and ERP000991.</p></fn><fn id="FN5"><p id="P46">Reprints and permissions information is available at <ext-link ext-link-type="uri" xlink:href="http://www.nature.com/">www.nature.com/</ext-link> reprints.</p></fn><fn id="FN6"><p id="P47">The authors declare no competing financial interests. 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The <italic>H. timareta</italic> races investigated have limited distributions indicated by arrows (red) and mimic sympatric races of <italic>H. melpomene</italic>. <italic>H. elevatus</italic> and the other silvaniform species are distributed widely across the Amazon basin (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 22</xref>).</p></caption><graphic xlink:href="ukmss-47290-f0001"></graphic></fig><fig id="F2" orientation="portrait" position="float"><label>Figure 2</label><caption><title>Comparative analysis of synteny and expansion of the chemosensory genes</title><p><bold>a</bold> Maps of the 21 <italic>Heliconius</italic> chromosomes (above) and of the 28 <italic>Bombyx</italic> chromosomes (below in grey) based on positions of 6010 orthologue pairs demonstrate highly conserved synteny and a shared <italic>n</italic>=31 ancestor (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 8</xref>). Dotted lines within chromosomes indicate major chromosomal fusions. <bold>b</bold> Maximum likelihood tree showing expansion of the chemosensory protein genes (CSP) in two butterfly genomes.</p></caption><graphic xlink:href="ukmss-47290-f0002"></graphic></fig><fig id="F3" orientation="portrait" position="float"><label>Figure 3</label><caption><title>Four-taxon ABBA-BABA test of introgression</title><p><bold>a</bold> ABBA and BABA nucleotide sites employed in the test are derived (– – <bold>B</bold> –) in <italic>H. timareta</italic>, compared with the silvaniform outgroup (– – – <bold>A</bold>), but differ among <italic>H. melpomene amaryllis</italic> and <italic>H. melpomene aglaope</italic> (either <bold>AB</bold>BA or <bold>BA</bold>BA). As this almost exclusively restricts attention to sites polymorphic in the ancestor of <italic>H. timareta</italic> and <italic>H. melpomene</italic>, equal numbers of ABBA and BABA sites<sup><xref ref-type="bibr" rid="R22">22</xref></sup> are expected under a null hypothesis of no introgression, as depicted in the two gene genealogies. <bold>b</bold> Distribution among chromosomes of Patterson’s <italic>D</italic>-statistic ± s.e., which measures excess of ABBA vs. BABA sites<sup><xref ref-type="bibr" rid="R22">22</xref></sup>, here for the comparison <italic>H. m. aglaope</italic>; <italic>H. m. amaryllis</italic>; <italic>H. timareta</italic> ssp. nov.; silvaniform. Chromosomes containing the two colour pattern regions (<italic>B</italic>/<italic>D</italic> red; <italic>N</italic>/<italic>Yb</italic> yellow) have the two highest <italic>D</italic>-statistics; the combinatorial probability of this occurring by chance is 0.005. The excess of ABBA sites (0 < <italic>D</italic> < 1) indicates introgression between sympatric <italic>H. timareta</italic> and <italic>H. melpomene amaryllis</italic>.</p></caption><graphic xlink:href="ukmss-47290-f0003"></graphic></fig><fig id="F4" orientation="portrait" position="float"><label>Figure 4</label><caption><title>Evidence for adaptive introgression at the <italic>B</italic>/<italic>D</italic> mimicry locus</title><p><bold>a</bold> Genetic divergence between <italic>H. melpomene</italic> races <italic>aglaope</italic> (rayed) and <italic>amaryllis</italic> (postman) across a hybrid zone in N.E. Peru. Divergence, <italic>F<sub>ST</sub></italic>, is measured along the <italic>B/D</italic> region (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 14</xref>). <italic>F<sub>ST</sub></italic> peaks in the region known to control red wing pattern elements between the genes <italic>kinesin</italic> and <italic>optix</italic><sup><xref ref-type="bibr" rid="R23">23</xref></sup>. <bold>b, c</bold> Distribution of fixed ABBA and BABA sites (see Fig. 4a) along <italic>B</italic>/<italic>D</italic> for two comparisons. Excesses of ABBA in <bold>b</bold> and BABA in <bold>c</bold> are highly significant (two-tailed <italic>Z</italic>-tests for <italic>D</italic> = 0; <italic>D</italic> = 0.90 ± 0.13, <italic>P</italic> = 5 × 10<sup>−14</sup> and <italic>D</italic> = –0.91 ± 0.10, <italic>P</italic> = 9 × 10<sup>−24</sup> respectively), indicating introgression. <bold>d, e, f,</bold> Genealogical change along <italic>B</italic>/<italic>D</italic> investigated with maximum likelihood based on 50 kb windows. Three representative tree topologies are shown. Topology A, the species tree, is found within the white windows. In topologies B (dark green window) and C (light green windows) taxa group by colour pattern rather than species. Within striped windows <italic>H. melpomene</italic> and/or <italic>H. timareta</italic> are paraphyletic but the taxa do not group by colour pattern. Support is shown for nodes with > 50% bootstrap support (<xref ref-type="supplementary-material" rid="SD1">Supplementary Information 19</xref>).</p></caption><graphic xlink:href="ukmss-47290-f0004"></graphic></fig></floats-group></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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