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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">A major gene controls mimicry and crypsis in butterflies and moths</title><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="A1">Department of Animal and Plant Sciences, University of Sheffield, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation></author><author><name sortKey="Pardo Diaz, Carolina" sort="Pardo Diaz, Carolina" uniqKey="Pardo Diaz C" first="Carolina" last="Pardo-Diaz">Carolina Pardo-Diaz</name><affiliation><nlm:aff id="A3">Biology Program, Faculty of Natural Sciences and Mathematics. Universidad del Rosario. Cra. 24 No 63C-69, Bogotá D.C., 111221, Colombia</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="A4">Institut de Systématique, Evolution et Biodiversité (UMR 7205 CNRS, MNHN, UPMC, EPHE, Sorbonne Université), Museum National d’Histoire Naturelle, CP50, 57 rue Cuvier, 75005 PARIS, France</nlm:aff></affiliation><affiliation><nlm:aff id="A5">Cell and Developmental Biology, John Innes Centre, Norwich, UK, NR4 7UH</nlm:aff></affiliation></author><author><name sortKey="Supple, Megan" sort="Supple, Megan" uniqKey="Supple M" first="Megan" last="Supple">Megan Supple</name><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation><affiliation><nlm:aff id="A6">The Australian National University, ACT, Australia</nlm:aff></affiliation></author><author><name sortKey="Saenko, Suzanne V" sort="Saenko, Suzanne V" uniqKey="Saenko S" first="Suzanne V." last="Saenko">Suzanne V. Saenko</name><affiliation><nlm:aff id="A4">Institut de Systématique, Evolution et Biodiversité (UMR 7205 CNRS, MNHN, UPMC, EPHE, Sorbonne Université), Museum National d’Histoire Naturelle, CP50, 57 rue Cuvier, 75005 PARIS, France</nlm:aff></affiliation></author><author><name sortKey="Wallbank, Richard W R" sort="Wallbank, Richard W R" uniqKey="Wallbank R" first="Richard W. R." last="Wallbank">Richard W. R. Wallbank</name><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation><affiliation><nlm:aff id="A7">Department of Zoology, University of Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Wu, Grace C" sort="Wu, Grace C" uniqKey="Wu G" first="Grace C." last="Wu">Grace C. Wu</name><affiliation><nlm:aff id="A8">Energy and Resources Group, University of California at Berkeley, CA, USA</nlm:aff></affiliation></author><author><name sortKey="Maroja, Luana" sort="Maroja, Luana" uniqKey="Maroja L" first="Luana" last="Maroja">Luana Maroja</name><affiliation><nlm:aff id="A9">Department of Biology, Williams College, MA, USA</nlm:aff></affiliation></author><author><name sortKey="Ferguson, Laura" sort="Ferguson, Laura" uniqKey="Ferguson L" first="Laura" last="Ferguson">Laura Ferguson</name><affiliation><nlm:aff id="A10">Department of Zoology, University of Oxford, UK</nlm:aff></affiliation></author><author><name sortKey="Hanly, Joseph J" sort="Hanly, Joseph J" uniqKey="Hanly J" first="Joseph J." last="Hanly">Joseph J. Hanly</name><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation><affiliation><nlm:aff id="A7">Department of Zoology, University of Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Hines, Heather" sort="Hines, Heather" uniqKey="Hines H" first="Heather" last="Hines">Heather Hines</name><affiliation><nlm:aff id="A11">Penn State University, 517 Mueller, University Park, PA 16802</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="A3">Biology Program, Faculty of Natural Sciences and Mathematics. Universidad del Rosario. Cra. 24 No 63C-69, Bogotá D.C., 111221, Colombia</nlm:aff></affiliation></author><author><name sortKey="Merrill, Richard" sort="Merrill, Richard" uniqKey="Merrill R" first="Richard" last="Merrill">Richard Merrill</name><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation><affiliation><nlm:aff id="A7">Department of Zoology, University of Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Dowling, Andrea" sort="Dowling, Andrea" uniqKey="Dowling A" first="Andrea" last="Dowling">Andrea Dowling</name><affiliation><nlm:aff id="A12">School of Biosciences, University of Exeter in Cornwall, Penryn, UK TR10 9EZ</nlm:aff></affiliation></author><author><name sortKey="Ffrench Constant, Richard" sort="Ffrench Constant, Richard" uniqKey="Ffrench Constant R" first="Richard" last="Ffrench-Constant">Richard Ffrench-Constant</name><affiliation><nlm:aff id="A12">School of Biosciences, University of Exeter in Cornwall, Penryn, UK TR10 9EZ</nlm:aff></affiliation></author><author><name sortKey="Llaurens, Violaine" sort="Llaurens, Violaine" uniqKey="Llaurens V" first="Violaine" last="Llaurens">Violaine Llaurens</name><affiliation><nlm:aff id="A4">Institut de Systématique, Evolution et Biodiversité (UMR 7205 CNRS, MNHN, UPMC, EPHE, Sorbonne Université), Museum National d’Histoire Naturelle, CP50, 57 rue Cuvier, 75005 PARIS, France</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="A4">Institut de Systématique, Evolution et Biodiversité (UMR 7205 CNRS, MNHN, UPMC, EPHE, Sorbonne Université), Museum National d’Histoire Naturelle, CP50, 57 rue Cuvier, 75005 PARIS, France</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="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation></author><author><name sortKey="Jiggins, Chris D" sort="Jiggins, Chris D" uniqKey="Jiggins C" first="Chris D." last="Jiggins">Chris D. Jiggins</name><affiliation><nlm:aff id="A7">Department of Zoology, University of Cambridge, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">27251285</idno><idno type="pmc">5094491</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC5094491</idno><idno type="RBID">PMC:5094491</idno><idno type="doi">10.1038/nature17961</idno><date when="2016">2016</date><idno type="wicri:Area/Pmc/Corpus">002D74</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">002D74</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">A major gene controls mimicry and crypsis in butterflies and moths</title><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="A1">Department of Animal and Plant Sciences, University of Sheffield, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation></author><author><name sortKey="Pardo Diaz, Carolina" sort="Pardo Diaz, Carolina" uniqKey="Pardo Diaz C" first="Carolina" last="Pardo-Diaz">Carolina Pardo-Diaz</name><affiliation><nlm:aff id="A3">Biology Program, Faculty of Natural Sciences and Mathematics. Universidad del Rosario. Cra. 24 No 63C-69, Bogotá D.C., 111221, Colombia</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="A4">Institut de Systématique, Evolution et Biodiversité (UMR 7205 CNRS, MNHN, UPMC, EPHE, Sorbonne Université), Museum National d’Histoire Naturelle, CP50, 57 rue Cuvier, 75005 PARIS, France</nlm:aff></affiliation><affiliation><nlm:aff id="A5">Cell and Developmental Biology, John Innes Centre, Norwich, UK, NR4 7UH</nlm:aff></affiliation></author><author><name sortKey="Supple, Megan" sort="Supple, Megan" uniqKey="Supple M" first="Megan" last="Supple">Megan Supple</name><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation><affiliation><nlm:aff id="A6">The Australian National University, ACT, Australia</nlm:aff></affiliation></author><author><name sortKey="Saenko, Suzanne V" sort="Saenko, Suzanne V" uniqKey="Saenko S" first="Suzanne V." last="Saenko">Suzanne V. Saenko</name><affiliation><nlm:aff id="A4">Institut de Systématique, Evolution et Biodiversité (UMR 7205 CNRS, MNHN, UPMC, EPHE, Sorbonne Université), Museum National d’Histoire Naturelle, CP50, 57 rue Cuvier, 75005 PARIS, France</nlm:aff></affiliation></author><author><name sortKey="Wallbank, Richard W R" sort="Wallbank, Richard W R" uniqKey="Wallbank R" first="Richard W. R." last="Wallbank">Richard W. R. Wallbank</name><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation><affiliation><nlm:aff id="A7">Department of Zoology, University of Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Wu, Grace C" sort="Wu, Grace C" uniqKey="Wu G" first="Grace C." last="Wu">Grace C. Wu</name><affiliation><nlm:aff id="A8">Energy and Resources Group, University of California at Berkeley, CA, USA</nlm:aff></affiliation></author><author><name sortKey="Maroja, Luana" sort="Maroja, Luana" uniqKey="Maroja L" first="Luana" last="Maroja">Luana Maroja</name><affiliation><nlm:aff id="A9">Department of Biology, Williams College, MA, USA</nlm:aff></affiliation></author><author><name sortKey="Ferguson, Laura" sort="Ferguson, Laura" uniqKey="Ferguson L" first="Laura" last="Ferguson">Laura Ferguson</name><affiliation><nlm:aff id="A10">Department of Zoology, University of Oxford, UK</nlm:aff></affiliation></author><author><name sortKey="Hanly, Joseph J" sort="Hanly, Joseph J" uniqKey="Hanly J" first="Joseph J." last="Hanly">Joseph J. Hanly</name><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation><affiliation><nlm:aff id="A7">Department of Zoology, University of Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Hines, Heather" sort="Hines, Heather" uniqKey="Hines H" first="Heather" last="Hines">Heather Hines</name><affiliation><nlm:aff id="A11">Penn State University, 517 Mueller, University Park, PA 16802</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="A3">Biology Program, Faculty of Natural Sciences and Mathematics. Universidad del Rosario. Cra. 24 No 63C-69, Bogotá D.C., 111221, Colombia</nlm:aff></affiliation></author><author><name sortKey="Merrill, Richard" sort="Merrill, Richard" uniqKey="Merrill R" first="Richard" last="Merrill">Richard Merrill</name><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation><affiliation><nlm:aff id="A7">Department of Zoology, University of Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Dowling, Andrea" sort="Dowling, Andrea" uniqKey="Dowling A" first="Andrea" last="Dowling">Andrea Dowling</name><affiliation><nlm:aff id="A12">School of Biosciences, University of Exeter in Cornwall, Penryn, UK TR10 9EZ</nlm:aff></affiliation></author><author><name sortKey="Ffrench Constant, Richard" sort="Ffrench Constant, Richard" uniqKey="Ffrench Constant R" first="Richard" last="Ffrench-Constant">Richard Ffrench-Constant</name><affiliation><nlm:aff id="A12">School of Biosciences, University of Exeter in Cornwall, Penryn, UK TR10 9EZ</nlm:aff></affiliation></author><author><name sortKey="Llaurens, Violaine" sort="Llaurens, Violaine" uniqKey="Llaurens V" first="Violaine" last="Llaurens">Violaine Llaurens</name><affiliation><nlm:aff id="A4">Institut de Systématique, Evolution et Biodiversité (UMR 7205 CNRS, MNHN, UPMC, EPHE, Sorbonne Université), Museum National d’Histoire Naturelle, CP50, 57 rue Cuvier, 75005 PARIS, France</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="A4">Institut de Systématique, Evolution et Biodiversité (UMR 7205 CNRS, MNHN, UPMC, EPHE, Sorbonne Université), Museum National d’Histoire Naturelle, CP50, 57 rue Cuvier, 75005 PARIS, France</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="A2">Smithsonian Tropical Research Institute, Panama</nlm:aff></affiliation></author><author><name sortKey="Jiggins, Chris D" sort="Jiggins, Chris D" uniqKey="Jiggins C" first="Chris D." last="Jiggins">Chris D. Jiggins</name><affiliation><nlm:aff id="A7">Department of Zoology, University of Cambridge, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Smithsonian Tropical Research Institute, Panama</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="2016">2016</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p id="P1">The wing patterns of butterflies and moths (Lepidoptera) are diverse and striking examples of evolutionary diversification by natural selection<xref rid="R1" ref-type="bibr">1</xref>,<xref rid="R2" ref-type="bibr">2</xref>. Lepidopteran wing colour patterns are a key innovation, consisting of arrays of coloured scales. We still lack a general understanding of how these patterns are controlled and if there is any commonality across the 160,000 moth and 17,000 butterfly species. Here, we identify a gene, <italic>cortex,</italic> through fine-scale mapping using population genomics and gene expression analyses, which regulates pattern switches in multiple species across the mimetic radiation in <italic>Heliconius</italic> butterflies. <italic>cortex</italic> belongs to a fast evolving subfamily of the otherwise highly conserved fizzy family of cell cycle regulators<xref rid="R3" ref-type="bibr">3</xref>, suggesting that it most likely regulates pigmentation patterning through regulation of scale cell development. In parallel with findings in the peppered moth (<italic>Biston betularia</italic>)<xref rid="R4" ref-type="bibr">4</xref>, our results suggest that this mechanism is common within Lepidoptera and that <italic>cortex</italic> has become a major target for natural selection acting on colour and pattern variation in this group of insects.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Cook, Lm" uniqKey="Cook L">LM Cook</name></author><author><name sortKey="Grant, Bs" uniqKey="Grant B">BS Grant</name></author><author><name sortKey="Saccheri, Ij" uniqKey="Saccheri I">IJ Saccheri</name></author><author><name sortKey="Mallet, J" uniqKey="Mallet J">J Mallet</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Jiggins, Cd" uniqKey="Jiggins C">CD Jiggins</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Dawson, Ia" uniqKey="Dawson I">IA Dawson</name></author><author><name 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<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">27251285</article-id><article-id pub-id-type="pmc">5094491</article-id><article-id pub-id-type="doi">10.1038/nature17961</article-id><article-id pub-id-type="manuscript">EMS68288</article-id><article-categories><subj-group subj-group-type="heading"><subject>Article</subject></subj-group></article-categories><title-group><article-title>A major gene controls mimicry and crypsis in butterflies and moths</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Nadeau</surname><given-names>Nicola J.</given-names></name><xref ref-type="aff" rid="A1">1</xref><xref ref-type="aff" rid="A2">2</xref></contrib><contrib contrib-type="author"><name><surname>Pardo-Diaz</surname><given-names>Carolina</given-names></name><xref ref-type="aff" rid="A3">3</xref></contrib><contrib contrib-type="author"><name><surname>Whibley</surname><given-names>Annabel</given-names></name><xref ref-type="aff" rid="A4">4</xref><xref ref-type="aff" rid="A5">5</xref></contrib><contrib contrib-type="author"><name><surname>Supple</surname><given-names>Megan</given-names></name><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A6">6</xref></contrib><contrib contrib-type="author"><name><surname>Saenko</surname><given-names>Suzanne V.</given-names></name><xref ref-type="aff" rid="A4">4</xref></contrib><contrib contrib-type="author"><name><surname>Wallbank</surname><given-names>Richard W. R.</given-names></name><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A7">7</xref></contrib><contrib contrib-type="author"><name><surname>Wu</surname><given-names>Grace C.</given-names></name><xref ref-type="aff" rid="A8">8</xref></contrib><contrib contrib-type="author"><name><surname>Maroja</surname><given-names>Luana</given-names></name><xref ref-type="aff" rid="A9">9</xref></contrib><contrib contrib-type="author"><name><surname>Ferguson</surname><given-names>Laura</given-names></name><xref ref-type="aff" rid="A10">10</xref></contrib><contrib contrib-type="author"><name><surname>Hanly</surname><given-names>Joseph J.</given-names></name><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A7">7</xref></contrib><contrib contrib-type="author"><name><surname>Hines</surname><given-names>Heather</given-names></name><xref ref-type="aff" rid="A11">11</xref></contrib><contrib contrib-type="author"><name><surname>Salazar</surname><given-names>Camilo</given-names></name><xref ref-type="aff" rid="A3">3</xref></contrib><contrib contrib-type="author"><name><surname>Merrill</surname><given-names>Richard</given-names></name><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A7">7</xref></contrib><contrib contrib-type="author"><name><surname>Dowling</surname><given-names>Andrea</given-names></name><xref ref-type="aff" rid="A12">12</xref></contrib><contrib contrib-type="author"><name><surname>ffrench-Constant</surname><given-names>Richard</given-names></name><xref ref-type="aff" rid="A12">12</xref></contrib><contrib contrib-type="author"><name><surname>Llaurens</surname><given-names>Violaine</given-names></name><xref ref-type="aff" rid="A4">4</xref></contrib><contrib contrib-type="author"><name><surname>Joron</surname><given-names>Mathieu</given-names></name><xref ref-type="aff" rid="A4">4</xref></contrib><contrib contrib-type="author"><name><surname>McMillan</surname><given-names>W. Owen</given-names></name><xref ref-type="aff" rid="A2">2</xref></contrib><contrib contrib-type="author"><name><surname>Jiggins</surname><given-names>Chris D.</given-names></name><xref ref-type="aff" rid="A7">7</xref><xref ref-type="aff" rid="A2">2</xref></contrib></contrib-group><aff id="A1"><label>1</label>Department of Animal and Plant Sciences, University of Sheffield, UK</aff><aff id="A2"><label>2</label>Smithsonian Tropical Research Institute, Panama</aff><aff id="A3"><label>3</label>Biology Program, Faculty of Natural Sciences and Mathematics. Universidad del Rosario. Cra. 24 No 63C-69, Bogotá D.C., 111221, Colombia</aff><aff id="A4"><label>4</label>Institut de Systématique, Evolution et Biodiversité (UMR 7205 CNRS, MNHN, UPMC, EPHE, Sorbonne Université), Museum National d’Histoire Naturelle, CP50, 57 rue Cuvier, 75005 PARIS, France</aff><aff id="A5"><label>5</label>Cell and Developmental Biology, John Innes Centre, Norwich, UK, NR4 7UH</aff><aff id="A6"><label>6</label>The Australian National University, ACT, Australia</aff><aff id="A7"><label>7</label>Department of Zoology, University of Cambridge, UK</aff><aff id="A8"><label>8</label>Energy and Resources Group, University of California at Berkeley, CA, USA</aff><aff id="A9"><label>9</label>Department of Biology, Williams College, MA, USA</aff><aff id="A10"><label>10</label>Department of Zoology, University of Oxford, UK</aff><aff id="A11"><label>11</label>Penn State University, 517 Mueller, University Park, PA 16802</aff><aff id="A12"><label>12</label>School of Biosciences, University of Exeter in Cornwall, Penryn, UK TR10 9EZ</aff><author-notes><corresp id="CR1">Correspondence and requests for materials should be addressed to <email>n.nadeau@sheffield.ac.uk</email> or <email>c.jiggins@zoo.cam.ac.uk</email></corresp></author-notes><pub-date pub-type="nihms-submitted"><day>8</day><month>9</month><year>2016</year></pub-date><pub-date pub-type="epub"><day>1</day><month>6</month><year>2016</year></pub-date><pub-date pub-type="collection"><day>1</day><month>6</month><year>2016</year></pub-date><pub-date pub-type="pmc-release"><day>01</day><month>12</month><year>2016</year></pub-date><volume>534</volume><issue>7605</issue><fpage>106</fpage><lpage>110</lpage><pmc-comment>elocation-id from pubmed: 10.1038/nature17961</pmc-comment>
<permissions><license><license-p>Users may view, print, copy, and download text and data-mine the content in such documents, for the purposes of academic research, subject always to the full Conditions of use:<ext-link ext-link-type="uri" xlink:href="http://www.nature.com/authors/editorial_policies/license.html#terms">http://www.nature.com/authors/editorial_policies/license.html#terms</ext-link></license-p></license></permissions><abstract><p id="P1">The wing patterns of butterflies and moths (Lepidoptera) are diverse and striking examples of evolutionary diversification by natural selection<xref rid="R1" ref-type="bibr">1</xref>,<xref rid="R2" ref-type="bibr">2</xref>. Lepidopteran wing colour patterns are a key innovation, consisting of arrays of coloured scales. We still lack a general understanding of how these patterns are controlled and if there is any commonality across the 160,000 moth and 17,000 butterfly species. Here, we identify a gene, <italic>cortex,</italic> through fine-scale mapping using population genomics and gene expression analyses, which regulates pattern switches in multiple species across the mimetic radiation in <italic>Heliconius</italic> butterflies. <italic>cortex</italic> belongs to a fast evolving subfamily of the otherwise highly conserved fizzy family of cell cycle regulators<xref rid="R3" ref-type="bibr">3</xref>, suggesting that it most likely regulates pigmentation patterning through regulation of scale cell development. In parallel with findings in the peppered moth (<italic>Biston betularia</italic>)<xref rid="R4" ref-type="bibr">4</xref>, our results suggest that this mechanism is common within Lepidoptera and that <italic>cortex</italic> has become a major target for natural selection acting on colour and pattern variation in this group of insects.</p></abstract></article-meta></front><body><p id="P2">In <italic>Heliconius</italic>, there is a major effect locus, <italic>Yb</italic>, that controls a diversity of colour pattern elements across the genus. It is the only locus in <italic>Heliconius</italic> that regulates all scale types and colours, including the diversity of white and yellow pattern elements in the two co-mimics <italic>H. melpomene</italic> (<italic>Hm</italic>) and <italic>H. erato</italic> (<italic>He</italic>), but also whole wing variation in black, yellow, white, and orange/red elements in <italic>H. numata</italic> (<italic>Hn</italic>)<xref rid="R5" ref-type="bibr">5</xref>–<xref rid="R7" ref-type="bibr">7</xref>. In addition, genetic variation underlying the <italic>Bigeye</italic> wing pattern mutation in <italic>Bicyclus anynana</italic>, melanism in the peppered moth, <italic>Biston betularia</italic>, and melanism and patterning differences in the silkmoth, <italic>Bombyx mori</italic>, have all been localised to homologous genomic regions<xref rid="R8" ref-type="bibr">8</xref>–<xref rid="R10" ref-type="bibr">10</xref> (<xref ref-type="fig" rid="F1">Fig 1</xref>). Therefore, this genomic region appears to contain one or more genes that act as major regulators of wing pigmentation and patterning across the Lepidoptera.</p><p id="P3">Previous mapping of this locus in <italic>He, Hm</italic> and <italic>Hn</italic> identified a genomic interval of ~1Mb<xref rid="R11" ref-type="bibr">11</xref>–<xref rid="R13" ref-type="bibr">13</xref> (<xref ref-type="table" rid="T1">Extended Data Table 1</xref>), which also overlaps with the 1.4Mb region containing the <italic>carbonaria</italic> locus in <italic>B. betularia</italic><xref rid="R9" ref-type="bibr">9</xref> and a 100bp non-coding region containing the <italic>Ws</italic> mutation in <italic>B. mori</italic><xref rid="R10" ref-type="bibr">10</xref> (<xref ref-type="fig" rid="F1">Fig 1</xref>). We took a population genomics approach to identify single nucleotide polymorphisms (SNPs) most strongly associated with phenotypic variation within the ~1Mb <italic>Heliconius</italic> interval. The diversity of wing patterning in <italic>Heliconius</italic> arises from divergence at wing pattern loci<xref rid="R7" ref-type="bibr">7</xref>, while convergent patterns generally involve the same loci and sometimes even the same alleles<xref rid="R14" ref-type="bibr">14</xref>–<xref rid="R16" ref-type="bibr">16</xref>. We used this pattern of divergence and sharing to identify SNPs associated with colour pattern elements across many individuals from a wide diversity of colour pattern phenotypes (<xref ref-type="fig" rid="F2">Fig 2</xref>).</p><p id="P4">In three separate <italic>Heliconius</italic> species, our analysis consistently implicated the gene <italic>cortex</italic> as being involved in adaptive differences in wing colour pattern. In <italic>He</italic> the strongest associations with the presence of a yellow hindwing bar were centred around the genomic region containing <italic>cortex</italic> (<xref ref-type="fig" rid="F2">Fig 2A</xref>). We identified 108 SNPs that were fixed for one allele in <italic>He favorinus</italic>, and fixed for the alternative allele in all individuals lacking the yellow bar, the majority of which were in introns of <italic>cortex</italic> (<xref ref-type="table" rid="T2">Extended Data Table 2</xref>). 15 SNPs showed a similar fixed pattern for <italic>He demophoon</italic>, which also has a yellow bar. These were non-overlapping with those in <italic>He favorinus</italic>, consistent with the hypothesis that this phenotype evolved independently in the two disjunct populations<xref rid="R17" ref-type="bibr">17</xref>.</p><p id="P5">Previous work has suggested that alleles at the <italic>Yb</italic> locus are shared between <italic>Hm</italic> and the closely related species <italic>H. timareta,</italic> and also the more distantly related species <italic>H. elevatus</italic>, resulting in mimicry between these species<xref rid="R18" ref-type="bibr">18</xref>. Across these species, the strongest associations with the yellow hindwing bar phenotype were again found at <italic>cortex</italic> (<xref ref-type="fig" rid="F2">Fig 2D</xref>, <xref ref-type="fig" rid="F5">Extended Data Fig 1A</xref> and <xref ref-type="table" rid="T3">Table 3</xref>). Similarly, the strongest associations with the yellow forewing band were found around the 5’ UTRs of <italic>cortex</italic> and gene <italic>HM00036,</italic> an orthologue of <italic>D. melanogaster washout</italic> gene. A single SNP ~17kb upstream of <italic>cortex</italic> (the closest gene) was perfectly associated with the yellow forewing band across all <italic>Hm, H. timareta</italic> and <italic>H. elevatus</italic> individuals (<xref ref-type="fig" rid="F5">Extended Data Fig 1A</xref>, <xref ref-type="fig" rid="F6">Fig 2</xref> and <xref ref-type="table" rid="T3">Table 3</xref>). We found no fixed coding sequence variants at <italic>cortex</italic> in a larger sample (43-61 individuals) of <italic>Hm aglaope</italic> and <italic>Hm amaryllis</italic> (<xref ref-type="fig" rid="F7">Extended Data Figure 3</xref>, <xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>), which differ in <italic>Yb</italic> controlled phenotypes<xref rid="R19" ref-type="bibr">19</xref>, suggesting that functional variants are likely to be regulatory rather than coding. We found extensive transposable element variation around <italic>cortex</italic> but it is unclear if any of these associate with phenotype (<xref ref-type="fig" rid="F7">Extended Data Figure 3</xref> and <xref ref-type="table" rid="T4">Table 4</xref>; <xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>).</p><p id="P6">Finally, in <italic>Hn</italic> large inversions at the <italic>P</italic> supergene locus (<xref ref-type="fig" rid="F1">Fig 1</xref>) are associated with different morphs<xref rid="R13" ref-type="bibr">13</xref>. There is a steep increase in genotype-by-phenotype association at the breakpoint of inversion 1, consistent with the role of these inversions in reducing recombination (<xref ref-type="fig" rid="F2">Fig 2E</xref>). However, the <italic>bicoloratus</italic> morph can recombine with all other morphs across one or the other inversion, permitting finer-scale association mapping of this region. As in <italic>He</italic> and <italic>Hm</italic>, this analysis showed a narrow region of associated SNPs corresponding exactly to the <italic>cortex</italic> gene (<xref ref-type="fig" rid="F2">Fig 2E</xref>), again with the majority of SNPs in introns (<xref ref-type="table" rid="T2">Extended Data Table 2</xref>). This associated region does not correspond to any other known genomic feature, such as an inversion or inversion breakpoint.</p><p id="P7">To determine whether sequence variants around <italic>cortex</italic> were regulating its expression we investigated gene expression across the <italic>Yb</italic> locus. We used a custom designed microarray including probes from all predicted genes in the <italic>H. melpomene</italic> genome<xref rid="R18" ref-type="bibr">18</xref>, as well as probes tiled across the central portion of the <italic>Yb</italic> locus, focussing on two naturally hybridising <italic>Hm</italic> races (<italic>plesseni</italic> and <italic>malleti</italic>) that differ in <italic>Yb</italic> controlled phenotypes<xref rid="R7" ref-type="bibr">7</xref>. <italic>cortex</italic> was the only gene across the entire interval to show significant expression differences both between races with different wing patterns and between wing sections with different pattern elements (<xref ref-type="fig" rid="F3">Fig 3</xref>). This finding was reinforced in the tiled probe set, where we observed strong differences in expression of <italic>cortex</italic> exons and introns but few differences outside this region (<xref ref-type="table" rid="T2">Extended Data Table 2</xref>). <italic>cortex</italic> expression was higher in <italic>Hm malleti</italic> than <italic>Hm plesseni</italic> in all three wing sections used (but not eyes) (<xref ref-type="fig" rid="F3">Fig 3C</xref>; <xref ref-type="fig" rid="F8">Extended Data Fig 4C</xref>). When different wing sections were compared within each race, <italic>cortex</italic> expression in <italic>Hm malleti</italic> was higher in the distal section that contains the <italic>Yb</italic> controlled yellow forewing band, consistent with <italic>cortex</italic> producing this band. In contrast, <italic>Hm plesseni</italic>, which lacks the yellow band, had higher <italic>cortex</italic> expression in the proximal forewing section (<xref ref-type="fig" rid="F3">Fig 3F</xref>; <xref ref-type="fig" rid="F8">Extended Data Fig 4J</xref>). Expression differences were found only in day 1 and day 3 pupal wings rather than day 5 or day 7 (<xref ref-type="fig" rid="F8">Extended Data Fig 4</xref>), similar to the pattern observed previously for the transcription factor <italic>optix</italic><xref rid="R20" ref-type="bibr">20</xref>.</p><p id="P8">Differential expression was not confined to the exons of <italic>cortex</italic>; the majority of differentially expressed probes in the tiling array corresponded to <italic>cortex</italic> introns (<xref ref-type="fig" rid="F3">Fig 3</xref>). This does not appear to be due to transposable element variation (<xref ref-type="table" rid="T2">Extended Data Table 2</xref>), but may be due to elevated background transcription and unidentified splice variants. RT-PCR revealed a diversity of splice variants (<xref ref-type="fig" rid="F9">Extended Data Fig 5</xref>), and sequenced products revealed 8 non-constitutive exons and 6 variable donor/acceptor sites, but this was not exhaustive (<xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>). We cannot rule out the possibility that some of the differentially expressed intronic regions could be distinct non-coding RNAs. However, qRT-PCR in other hybridising races with divergent <italic>Yb</italic> alleles (<italic>aglaope/amaryllis</italic> and <italic>rosina/melpomene)</italic> also identified expression differences at <italic>cortex</italic> and allele-specific splicing differences between both pairs of races (<xref ref-type="fig" rid="F5">Extended Data Figs 1</xref> and <xref ref-type="fig" rid="F9">5</xref>, <xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>).</p><p id="P9">Finally, <italic>in situ</italic> hybridisation of <italic>cortex</italic> in final instar larval hindwing discs showed expression in wing regions fated to become black in the adult wing, most strikingly in their correspondence to the black patterns on adult <italic>Hn</italic> wings (<xref ref-type="fig" rid="F4">Fig 4</xref>). In contrast, the array results from pupal wings were suggestive of higher expression in non-melanic regions. This may suggest that <italic>cortex</italic> is upregulated at different time-points in wing regions fated to become different colours.</p><p id="P10">Overall, <italic>cortex</italic> shows significant differential expression and is the only gene in the candidate region to be consistently differentially expressed in multiple race comparisons and between differently patterned wing regions. Coupled with the strong genotype-by-phenotype associations across multiple independent lineages (<xref ref-type="table" rid="T1">Extended Data Table 1</xref>), this strongly implicates <italic>cortex</italic> as a major regulator of colour and pattern. However, we have not excluded the possibility that other genes in this region also influence pigmentation patterning. A prominent role for <italic>cortex</italic> is also supported by studies in other taxa; our identification of distant 5’ untranslated exons of <italic>cortex</italic> (<xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>) suggests that the 100bp interval containing the <italic>Ws</italic> mutation in <italic>B. mori</italic> is likely to be within an intron of <italic>cortex</italic> and not in intergenic space as previously thought<xref rid="R10" ref-type="bibr">10</xref>. In addition, fine-mapping and gene expression also implicate <italic>cortex</italic> as controlling melanism in the peppered moth<xref rid="R4" ref-type="bibr">4</xref>.</p><p id="P11">It seems likely that <italic>cortex</italic> controls pigmentation patterning through control of scale cell development. The <italic>cortex</italic> gene falls in an insect specific lineage within the fizzy/CDC20 family of cell cycle regulators (<xref ref-type="fig" rid="F10">Extended Data Fig 6A</xref>). The phylogenetic tree of the gene family highlighted three major orthologous groups, two of which have highly conserved functions in cell cycle regulation mediated through interaction with the anaphase promoting complex/cyclosome (APC/C)<xref rid="R3" ref-type="bibr">3</xref>,<xref rid="R21" ref-type="bibr">21</xref>. The third group, cortex, is evolving rapidly, with low amino acid identity between <italic>D. melanogaster</italic> and <italic>Hm</italic> cortex (14.1%), contrasting with much higher identities for orthologues between these species in the other two groups (fzy, 47.8% and rap/fzr,47.2%, <xref ref-type="fig" rid="F10">Extended Data Fig 6A</xref>). <italic>Drosophila melanogaster</italic> cortex acts through a similar mechanism to fzy in order to control meiosis in the female germ line<xref rid="R22" ref-type="bibr">22</xref>–<xref rid="R24" ref-type="bibr">24</xref>. <italic>Hm</italic> cortex also has some conservation of the fizzy family C-box and IR elements (<xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>) that mediate binding to the APC/C<xref rid="R23" ref-type="bibr">23</xref>, suggesting that it may have retained a cell cycle function, although we found that expressing <italic>Hm cortex</italic> in <italic>D. melanogaster</italic> wings produced no detectable effect (<xref ref-type="fig" rid="F10">Extended Data Fig 6</xref>, <xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>).</p><p id="P12">Previously identified butterfly wing patterning genes have been transcription factors or signalling molecules<xref rid="R20" ref-type="bibr">20</xref>,<xref rid="R25" ref-type="bibr">25</xref>. Developmental rate has long been thought to play a role in lepidopteran patterning<xref rid="R26" ref-type="bibr">26</xref>,<xref rid="R27" ref-type="bibr">27</xref>, but <italic>cortex</italic> was not a likely <italic>a priori</italic> candidate, because its <italic>Drosophila</italic> orthologue has a highly specific function in meiosis<xref rid="R23" ref-type="bibr">23</xref>. The recruitment of <italic>cortex</italic> to wing patterning appears to have occurred before the major diversification of the Lepidoptera and this gene has repeatedly been targeted by natural selection<xref rid="R1" ref-type="bibr">1</xref>,<xref rid="R7" ref-type="bibr">7</xref>,<xref rid="R9" ref-type="bibr">9</xref>,<xref rid="R28" ref-type="bibr">28</xref> to generate both cryptic<xref rid="R4" ref-type="bibr">4</xref> and aposematic patterns.</p><sec sec-type="methods" id="S1" specific-use="web-only"><title>Methods</title><sec id="S2"><title>He Cr reference</title><p id="P13"><italic>Cr</italic> is the homologue of <italic>Yb</italic> in <italic>He</italic> (<xref ref-type="fig" rid="F1">Fig 1</xref>). An existing reference for this region was available in 3 pieces (467,734bp, 114,741bp and 161,149bp, GenBank: KC469893.1)<xref rid="R31" ref-type="bibr">31</xref>. We screened the same BAC library used previously<xref rid="R11" ref-type="bibr">11</xref>,<xref rid="R31" ref-type="bibr">31</xref> using described procedures<xref rid="R11" ref-type="bibr">11</xref> with probes designed to the ends of the existing BAC sequences and the <italic>HmYb</italic> BAC reference sequence. Two BACs (04B01 and 10B14) were identified as spanning one of the gaps and sequenced using Illumina 2x250 bp paired-end reads collected on the Illumina MiSeq. The raw reads were screened to remove vector and <italic>E.</italic> coli bases. The first 50k read pairs were taken for each BAC and assembled individually with the Phrap<xref rid="R32" ref-type="bibr">32</xref> software and manually edited with consed<xref rid="R33" ref-type="bibr">33</xref>. Contigs with discordant read pairs were manually broken and properly merged using concordant read data. Gaps between contig ends were filled using an in-house finishing technique where the terminal 200bp of the contig ends were extracted and queried against the unused read data for spanning pairs, which were added using the addSolexaReads.perl script in the consed package. Finally, a single reference contig was generated by identifying and merging overlapping regions of the two consensus BAC sequences.</p><p id="P14">In order to fill the remaining gap (between positions 800,387 and 848,446) we used the overhanging ends to search the scaffolds from a preliminary <italic>He</italic> genome assembly of five Illumina paired end libraries with different insert sizes (250, 500, 800, 4300 and 6500bp) from two related <italic>He demophoon</italic> individuals. We identified two scaffolds (scf1869 and scf1510) that overlapped and spanned the gap (using 12,257bp of the first scaffold and 35,803bp of the second).</p><p id="P15">The final contig was 1,009,595bp in length of which 2,281bp were unknown (N’s). The <italic>HeCr</italic> assembly was verified by aligning to the <italic>HmYb</italic> genome scaffold (HE667780) with mummer and blast. The <italic>HeCr</italic> contig was annotated as described previously<xref rid="R32" ref-type="bibr">32</xref>, with some minor modifications. Briefly this involved first generating a reference based transcriptome assembly with existing <italic>H. erato</italic> RNA-seq wing tissue (GenBank accession SRA060220). We used Trimmomatic<xref rid="R34" ref-type="bibr">34</xref> (v0.22), and FLASh<xref rid="R35" ref-type="bibr">35</xref> (v1.2.2) to prepare the raw sequencing reads, checking the quality with FastQC<xref rid="R36" ref-type="bibr">36</xref> (v0.10.0). We then used the Bowtie/TopHat/Cufflinks<xref rid="R37" ref-type="bibr">37</xref>–<xref rid="R39" ref-type="bibr">39</xref> pipeline to generate transcripts for the unmasked reference sequence. We generated gene predictions with the MAKER pipeline<xref rid="R40" ref-type="bibr">40</xref> (v2.31). Homology and synteny in gene content with the <italic>Hm Yb</italic> reference were identified by aligning the <italic>Hm</italic> coding sequences to the <italic>He</italic> reference with BLAST. Homologous genes were present in the same order and orientation in <italic>He</italic> and <italic>Hm</italic> (<xref ref-type="fig" rid="F2">Fig 2B,C</xref>). Annotations were manually adjusted if genes had clearly been merged or split in comparison to <italic>H. melpomene</italic> (which has been extensively manually curated<xref rid="R12" ref-type="bibr">12</xref>). In addition <italic>He cortex</italic> was manually curated from the RNA-seq data and using <italic>Exonerate</italic><xref rid="R41" ref-type="bibr">41</xref> alignments of the <italic>H. melpomene</italic> protein and mRNA transcripts, including the 5’ UTRs.</p></sec><sec id="S3"><title>Genotype-by-phenotype association analyses</title><p id="P16">Information on the individuals used and ENA accessions for sequence data are given in <xref ref-type="supplementary-material" rid="SD4">Supplementary Table 1</xref>. We used shotgun Illumina sequence reads from 45 <italic>He</italic> individuals from 7 races that were generated as part of a previous study<xref rid="R31" ref-type="bibr">31</xref> (<xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>). Reads were aligned to an <italic>He</italic> reference containing the <italic>Cr</italic> contig and other sequenced <italic>He</italic> BACs<xref rid="R11" ref-type="bibr">11</xref>,<xref rid="R31" ref-type="bibr">31</xref> with BWA<xref rid="R42" ref-type="bibr">42</xref> , which has previously been found to work better than Stampy<xref rid="R43" ref-type="bibr">43</xref> (which was used for the alignments in the other species) with an incomplete reference sequence<xref rid="R31" ref-type="bibr">31</xref>. The parameters used were as follows: Maximum edit distance (n), 8; maximum number of gap opens (o), 2; maximum number of gap extensions (e), 3; seed (l), 35; maximum edit distance in seed (k), 2. We then used Picard tools to remove PCR and optical duplicate sequence reads and GATK<xref rid="R44" ref-type="bibr">44</xref> to re-align indels and call SNPs using all individuals as a single population. Expected heterozygosity was set to 0.2 in GATK. 132,397 SNPs were present across <italic>Cr</italic>. A further 52,698 SNPs not linked to colour pattern loci were used to establish background association levels.</p><p id="P17">For the <italic>Hm / Hn</italic> clade we used previously published sequence data from 19 individuals from enrichment sequencing targeting of the <italic>Yb</italic> region, the unlinked <italic>HmB/D</italic> region that controls the presence/absence of red colour pattern elements, and ~1.8Mb of non-colour pattern genomic regions<xref rid="R45" ref-type="bibr">45</xref>, as well as 9 whole genome shotgun sequenced individuals<xref rid="R18" ref-type="bibr">18</xref>,<xref rid="R46" ref-type="bibr">46</xref>. We added targeted sequencing and shotgun whole genome sequencing of an additional 47 individuals (<xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>). Alignments were performed using Stampy<xref rid="R43" ref-type="bibr">43</xref> with default parameters except for substitution rate which was set to 0.01. We again removed duplicates and used GATK to re-align indels and call SNPs with expected heterozygosity set to 0.1.</p><p id="P18">The analysis of the <italic>Hm/timareta</italic>/silvaniform included 49 individuals, which were aligned to v1.1 of the <italic>Hm</italic> reference genome with the scaffolds containing <italic>Yb</italic> and <italic>HmB/D</italic> swapped with reference BAC sequences<xref rid="R18" ref-type="bibr">18</xref>, which contained fewer gaps of unknown sequence than the genome scaffolds. 232,631 SNPs were present in the <italic>Yb</italic> region and a further 370,079 SNPs were used to establish background association levels.</p><p id="P19">The <italic>Hn</italic> analysis included 26 individuals aligned to unaltered v1.1 of the <italic>Hm</italic> reference genome, because the genome scaffold containing <italic>Yb</italic> is longer than the BAC reference making it easier to compare the inverted and non-inverted regions present in this species. We tested for associations at 262,137 SNPs on the <italic>Yb</italic> scaffold with the <italic>Hn bicoloratus</italic> morph, which had a sample of 5 individuals.</p><p id="P20">We measured associations between genotype and phenotype using a score test (qtscore) in the GenABEL package in R<xref rid="R47" ref-type="bibr">47</xref>. This was corrected for background population structure using a test specific inflation factor, λ, calculated from the SNPs unlinked to the major colour pattern controlling loci (described above), as the colour pattern loci are known to have different population structure to the rest of the genome<xref rid="R14" ref-type="bibr">14</xref>,<xref rid="R15" ref-type="bibr">15</xref>,<xref rid="R18" ref-type="bibr">18</xref>. We used a custom perl script to convert GATK vcf files to Illumina SNP format for input to genABEL<xref rid="R47" ref-type="bibr">47</xref>. genABEL does not accept multiallelic sites, so the script also converted the genotype of any individuals for which a third (or fourth) allele was present to a missing genotype (with these defined as the lowest frequency alleles). Custom R scripts were used to identify sites showing perfect associations with calls for >75% of individuals.</p></sec><sec id="S4"><title>Microarray Gene Expression Analyses</title><p id="P21">We designed a Roche NimbleGen microarray (12x135K format) with probes for all annotated <italic>Hm</italic> genes<xref rid="R18" ref-type="bibr">18</xref> and tiling the central portion of the <italic>Yb</italic> BAC sequence contig that was previously identified as showing the strongest differentiation between <italic>Hm</italic> races<xref rid="R45" ref-type="bibr">45</xref>. In addition to the <italic>HmYb</italic> tilling array probes there were 6,560 probes tiling <italic>HmAc</italic> (a third unlinked colour pattern locus) and 10,716 probes tiling <italic>HmB/D</italic>, again distanced on average at 10bp intervals. The whole-genome gene expression array contained 107,898 probes in total.</p><p id="P22">This was interrogated with Cy3 labelled double stranded cDNA generated from total RNA (with a SuperScript double-stranded cDNA synthesis kit, Invitrogen, and a one-colour DNA labelling kit, Niblegen) from four pupal developmental stages of <italic>Hm plesseni</italic> and <italic>malleti.</italic> Pupae were from captive stocks maintained in insectary facilities in Gamboa, Panama. Tissue was stored in RNA later at -80°C prior to RNA extraction. RNA was extracted using TRIzol (Invitrogen) followed by purification with RNeasy (Qiagen) and DNase treated with DNA-free (Ambion). Quantification was performed using a Qubit 2.0 fluorometer (Invitrogen) and purity and integrity assessed using a Bioanalyzer 2100 (Agilent). Samples were randomised and each hybridised to a separate array. The <italic>HmYb</italic> probe array contained 9,979 probes distanced on average at 10bp. The whole-genome expression array contained on average 9 probes per annotated gene in the genome (v1.1<xref rid="R18" ref-type="bibr">18</xref>) as well as any transcripts not annotated but predicted from RNA-seq evidence.</p><p id="P23">Background corrected expression values for each probe were extracted using NimbleScan software (version 2.3). Analyses were performed with the LIMMA package implemented in R/Bioconductor<xref rid="R48" ref-type="bibr">48</xref>. The tiling array and whole-genome data sets were analysed separately. Expression values were extracted and quantile-normalised, log<sub>2</sub>-transformed, quality controlled and analysed for differences in expression between individuals and wing regions. P-values were adjusted for multiple hypotheses testing using the False Discovery Rate (FDR) method <xref rid="R49" ref-type="bibr">49</xref>.</p><p id="P24">We detected isoform-specific expression differences between <italic>Hm aglaope</italic>/<italic>amaryllis</italic> and <italic>Hm rosina/melpomene</italic> using RT-PCR and qRT-PCR on RNA extracted from developing hind-wing tissue (further details in <xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>). Previously published RNAseq data was also used to assess gene expression differences between <italic>Hm aglaope</italic> and <italic>amaryllis</italic><xref rid="R18" ref-type="bibr">18</xref> (further details in <xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>).</p></sec><sec id="S5"><title><italic>In situ</italic> hybridisations</title><p id="P25"><italic>Hn</italic> and <italic>Hm</italic> larvae were reared in a greenhouse at 25-30°C and sampled at the last instar. In situ hybridizations were performed according to previously described methods<xref rid="R25" ref-type="bibr">25</xref> with a <italic>cortex</italic> riboprobe synthesized from a 831-bp cDNA amplicon from <italic>Hn</italic>. Wing discs were incubated in a standard hybridization buffer containing the probe for 20-24 h at 60°C. For secondary detection of the probe, wing discs were incubated in a 1:3000 dilution of anti-digoxigenin alkaline phosphatase Fab fragments and stained with BM Purple for 3-6 h at room temperature. Stained wing discs were photographed with a Leica DFC420 digital camera mounted on a Leica Z6 APO stereomicroscope.</p></sec><sec id="S6"><title><italic>De novo</italic> assembly of short read data in <italic>Hm</italic> and related taxa</title><p id="P26">In order to better characterise indel variation from the short-read sequence data used for the genotype-by-phenotype association analysis, we performed <italic>de novo</italic> assemblies of a subset of <italic>Hm</italic> individuals and related taxa with a diversity of phenotypes (<xref ref-type="fig" rid="F6">Extended Data Figure 2</xref>). Assemblies were performed using the <italic>de novo</italic> assembly function of CLCGenomics Workbench v.6.0 under default parameters. The assembled contigs were then BLASTed against the <italic>Yb</italic> region of the <italic>Hm melpomene</italic> genome<xref rid="R18" ref-type="bibr">18</xref>, using Geneious v.8.0. The contigs identified by BLAST were then concatenated to generate an allele sequence for each individual. Occasionally two unphased alleles were generated when two contigs were matched to a given region. If more than two contigs of equal length matched then this was considered an unresolvable repeat region and replaced with Ns. The assembled alleles were then aligned using the MAFFT alignment plugin in Geneious v.8.0.</p></sec><sec id="S7"><title>Long-range PCR targeted sequencing of cortex in <italic>Hm aglaope</italic> and <italic>Hm amaryllis</italic></title><p id="P27">We generated two long-range PCR products covering 88.8% of the 1,344bp coding region of <italic>cortex</italic> (excluding 67bp at the 5’ end and 83bp at the 3’ end, further details in <xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>). A product spanning coding exons 5 to 9 (the final exon) was obtained from 29 <italic>Hm amaryllis</italic> individuals and 29 <italic>Hm aglaope</italic> individuals; a product spanning coding exons 2 to 5 was obtained from 32 <italic>Hm amaryllis</italic> individuals and 14 <italic>Hm aglaope</italic>. In addition, a product spanning exons 4 to 6 was obtained from 6 <italic>Hm amaryllis</italic> and 5 <italic>Hm aglaope</italic> that failed to amplify one or both of the larger products. Long-range PCR was performed using Extensor long-range PCR mastermix (Thermo Scientific) following manufacturers guidelines with a 60°C annealing temperature in a 10-20µl volume. The product spanning coding exons 5 to 9 was obtained with primers HM25_long_F1 and HM25_long_R4 (see <xref ref-type="supplementary-material" rid="SD5">Supplementary Table 2</xref> for primer sequences); the product spanning coding exons 2 to 5 was obtained with primers HM25_long_F4 and HM25_long_R2; the product spanning exons 4 to 6 was obtained with primers 25_ex5-ex7_r1 and 25_ex5-ex7_f1. Products were pooled for each individual, including 5 additional products from the <italic>Yb</italic> locus and 7 products in the region of the <italic>HmB/D</italic> locus. They were then cleaned using QIAquick PCR purification kit (QIAgen) before being quantified with a Qubit Fluorometer (Life Technologies) and pooled in equimolar amounts for each individual, taking into account variation in the length and number of PCR products included for each individual (because of some PCR failures, ie. proportionally less DNA was included if some PCR products were absent for a given individual).</p><p id="P28">Products were pooled within individuals (including additional products for other genes not analysed here) and then quantified and pooled in equimolar amounts for each individual within each race. The pooled products for each race (<italic>Hm aglaope</italic> and <italic>amaryllis)</italic> were then prepared as two separate libraries with molecular identifiers and sequenced on a single lane of an Illumina GAIIx. Analysis was performed using Galaxy and the history is available at <ext-link ext-link-type="uri" xlink:href="https://usegalaxy.org/u/njnadeau/h/long-pcr-final">https://usegalaxy.org/u/njnadeau/h/long-pcr-final</ext-link>. Reads were quality filtered with a minimum quality of 20 required over 90% of the read, which resulted in 5% of reads being discarded. Reads were then quality trimmed to remove bases with quality less than 20 from the ends. They were then aligned to the target regions using the fosmid sequences from known races<xref rid="R45" ref-type="bibr">45</xref> with sequence from the <italic>Yb</italic> BAC walk<xref rid="R12" ref-type="bibr">12</xref> used to fill any gaps. Alignments were performed with BWA v0.5.6<xref rid="R42" ref-type="bibr">42</xref> and converted to pileup format using Samtools v0.1.12 before being filtered based on quality (≥20) and coverage (≥10). BWA alignment parameters were as follows: fraction of missing alignments given 2% uniform base error rate (aln -n) 0.01; maximum number of gap opens (aln -o) 2; maximum number of gap extensions (aln -e) 12; disallow long deletion within 12 bp towards the 3'-end (aln -d); number of first subsequences to take as seed (aln -l) 100. We then calculated coverage and minor allele frequencies for each race and the difference between these using custom scripts in R<xref rid="R50" ref-type="bibr">50</xref>.</p></sec><sec id="S8"><title>Sequencing and analysis of <italic>Hm</italic> fosmid clones</title><p id="P29">Fosmid libraries had previously been made from single individuals of 3 <italic>Hm</italic> races (<italic>rosina</italic>, <italic>amaryllis</italic> and <italic>aglaope</italic>) and several clones overlapping the <italic>Yb</italic> interval had been sequenced<xref rid="R45" ref-type="bibr">45</xref>. We extended the sequencing of this region, particularly the region overlapping <italic>cortex</italic> by sequencing an additional 4 clones from <italic>Hm rosina</italic> (1051_83D21, accession KU514430; 1051_97A3, accession KU514431; 1051_65N6, accession KU514432; 1051_93D23, accession KU514433) 2 clones from <italic>Hm amaryllis</italic> (1051_13K4, accession KU514434; 1049_8P23, accession KU514435) and 3 clones from <italic>Hm aglaope</italic> (1048_80B22, accession KU514437; 1049_19P15, accession KU514436; 1048_96A7, accession KU514438). These were sequenced on a MiSeq 2000, and assembled using the <italic>de novo</italic> assembly function of CLCGenomcs Workbench v.6.0. The individual clones (including existing clones 1051-143B3, accession FP578990; 1049-27G11, accession FP700055; 1048-62H20, accession FP565804) were then aligned to the BAC and genome scaffold<xref rid="R18" ref-type="bibr">18</xref> references using the MAFFT alignment plugin of Geneious v.8.0. Regions of general sequence similarity were identified and visualised using MAUVE<xref rid="R51" ref-type="bibr">51</xref>. We merged overlapping clones from the same individual if they showed no sequence differences, indicating that they came from the same allele. We identified transposable elements (TEs) using nBLAST with an insect TE list downloaded from Repbase Update<xref rid="R52" ref-type="bibr">52</xref> including known <italic>Heliconius</italic> specific TEs<xref rid="R53" ref-type="bibr">53</xref>.</p></sec><sec id="S9"><title>5’ RACE, RT-PCR and qRT-PCR</title><p id="P30">All tissues used for gene expression analyses were dissected from individuals from captive stocks derived from wild caught individuals of various races of <italic>Hm (aglaope, amaryllis, melpomene, rosina, plesseni, malleti</italic>) and F2 individuals from a <italic>Hm rosina</italic> (female) x <italic>Hm melpomene</italic> (male) cross. Experimental individuals were reared at 28°C-31°C. Developing wings were dissected and stored in RNAlater (Ambion Life Technologies). RNA was extracted using a QIAgen RNeasy Mini kit following the manufacturer’s guidelines and treated with TURBO DNA-free DNase kit (Ambion Life Technologies) to remove remaining genomic DNA. RNA quantification was performed with a Nanodrop spectrophotometer, and the RNA integrity was assessed using the Bioanalyzer 2100 system (Agilent).</p><p id="P31">Total RNA was thoroughly checked for DNA contamination by performing PCR for EF1α (using primers ef1-a_RT_for and ef1-a_RT_rev, <xref ref-type="supplementary-material" rid="SD5">Table S2</xref>) with 0.5µl of RNA extract (50ng-1µg of RNA) in a 20µl reaction using a polymerase enzyme that is not functional with RNA template (BioScript, Bioline Reagents Ltd.). If a product amplified within 45 cycles then the RNA sample was re-treated with DNase.</p><p id="P32">Single stranded cDNA was synthesised using BioScript MMLV Reverse Transcriptase (Bioline Reagents Ltd.) with random hexamer (N6) primers and 1µg of template RNA from each sample in a 20 µl reaction volume following the manufacturer’s protocol. The resulting cDNA samples were then diluted 1:1 with nuclease free water and stored at -80°C.</p><p id="P33">5’ RACE was performed using RNA from hind-wing discs from one <italic>Hm aglaope</italic> and one <italic>Hm amaryllis</italic> final instar larvae with a SMARTer RACE kit from Clonetech (California, USA). The gene specific primer used for the first round of amplification was anchored in exon 4 (fzl_raceex5_R1, <xref ref-type="supplementary-material" rid="SD5">Supplementary Table 2</xref>). Secondary PCR of these products was then performed using a primer in exon 2 (HM25_long_F2, <xref ref-type="supplementary-material" rid="SD5">Supplementary Table 2</xref>) and the nested universal primer A. Other isoforms were detected by RT-PCR using primers within exons 2 and 9 (gene25_for_full1 and gene25_rev_ex3). We identified isoforms from 5’ RACE and RT-PCR products by cutting individual bands from agarose gels and if necessary by cloning products before Sanger sequencing. Cloning of products was performed using TOPO TA (Invitrogen) or pGEM-T (Promega) cloning kits. Sanger sequencing was performed using BigDye terminator v3.1 (Applied Biosystems) run on an ABI13730 capillary sequencer. Primers fzl_ex1a_F1 and fzl_ex4_R1 were used to confirm expression of the furthest 5’ UTR. For isoforms that appeared to show some degree of race specificity we designed isoform specific PCR primers spanning specific exon junctions (<xref ref-type="fig" rid="F6">Extended Data Fig 2</xref>, <xref ref-type="fig" rid="F8">4</xref>, <xref ref-type="supplementary-material" rid="SD5">Supplementary Table 2</xref>) and used these to either qualitatively (RT-PCR) or quantitatively (qRT-PCR) assess differences in expression between races.</p><p id="P34">We performed qRT-PCR using SensiMix SYBR green (Bioline Reagents Ltd.) with 0.2-0.25µM of each primer and 1μl of the diluted product from the cDNA reactions. Reactions were performed in an Opticon 2 DNA engine (MJ Research), with the following cycling parameters: 95°C for 10min, 35-50 x: (95°C for 15sec, 55-60°C for 30sec, 72° for 30sec), 72°C for 5min. Melting curves were generated between 55°C and 90°C with readings taken every 0.2°C for each of the products to check that a single product was generated. At least one product from each set of primers was also run on a 1% agarose gel to check that a single product of the expected size was produced and the identity of the product confirmed by direct sequencing (See <xref ref-type="supplementary-material" rid="SD5">Supplementary Table 2</xref> for details of primers for each gene). We used two housekeeping genes (<italic>EF1α</italic> and <italic>Ribosomal Protein S3A)</italic> for normalisation and all results were taken as averages of triplicate PCR reactions for each sample.</p><p id="P35"><italic>C<sub>t</sub></italic> values were defined as the point at which fluorescence crossed a threshold (<italic>R<sub>Ct</sub></italic>) adjusted manually to be the point at which fluorescence rose above the background level. Amplification efficiencies (<italic>E</italic>) were calculated using a dilution series of clean PCR product. Starting fluorescence, which is proportional to the starting template quantity, was calculated as <italic>R<sub>0</sub></italic> = <italic>R<sub>Ct</sub></italic> (1+<italic>E</italic>)<sup><italic>-Ct</italic></sup>. Normalized values were then obtained by dividing <italic>R<sub>0</sub></italic> values for the target loci by <italic>R<sub>0</sub></italic> values for EF1α and RPS3A. Results from both of these controls were always very similar, therefore the results presented are normalized to the mean of EF1α and RPS3A. All results were taken as averages of triplicate PCR reactions. If one of the triplicate values was more than one cycle away from the mean then this replicate was excluded. Similarly any individuals that were more than two standard deviations away from the mean of all individuals for the target or normalization genes were excluded (these are not included in the numbers of individuals reported). Statistical significance was assessed by Wilcoxon rank sum tests performed in R<xref rid="R50" ref-type="bibr">50</xref>.</p></sec><sec id="S10"><title>RNAseq analysis of <italic>Hm amaryllis/aglaope</italic></title><p id="P36">RNA-seq data for hind-wings from three developmental stages had previously been obtained for two individuals of each race at each stage (12 individuals in total) and used in the annotation of the <italic>Hm</italic> genome<xref rid="R18" ref-type="bibr">18</xref> (deposited in ENA under study accessions ERP000993 and PRJEB7951). Four samples were multiplexed on each sequencing lane with the fifth instar larval and day 2 pupal samples sequenced on a GAIIx sequencer and the day 3 pupal wings sequenced on a Hiseq 2000 sequencer.</p><p id="P37">Two methods were used for alignment of reads to the reference genome and inferring read counts, Stampy<xref rid="R43" ref-type="bibr">43</xref> and RSEM (RNAseq by Expectation Maximisation)<xref rid="R54" ref-type="bibr">54</xref>. In addition we used two different R/Bioconductor packages for estimation of differential gene expression, DESeq<xref rid="R55" ref-type="bibr">55</xref> and BaySeq <xref rid="R56" ref-type="bibr">56</xref>. Read bases with quality scores < 20 were trimmed with FASTX-Toolkit (<ext-link ext-link-type="uri" xlink:href="http://hannonlab.cshl.edu/fastx_toolkit/index.html">http://hannonlab.cshl.edu/fastx_toolkit/index.html</ext-link>). Stampy was run with default parameters except for mean insert size, which was set to 500, SD 100 and substitution rate, which was set to 0.01. Alignments were filtered to exclude reads with mapping quality <30 and sorted using Samtools<xref rid="R57" ref-type="bibr">57</xref>. We used the HT seq-count script in with HTseq<xref rid="R58" ref-type="bibr">58</xref> to infer counts per gene from the BAM files.</p><p id="P38">RSEM<xref rid="R54" ref-type="bibr">54</xref> was run with default parameters to infer a transcriptome and then map RNAseq reads against this using Bowtie<xref rid="R37" ref-type="bibr">37</xref> as an aligner. This was run with default parameters except maximum number of mismatches, which was set to 3.</p></sec><sec id="S11"><title>Annotation and alignment of fizzy family proteins</title><p id="P39">In the arthropod genomes, some fizzy family proteins were found to be poorly annotated based on alignments to other family members. In these cases annotations were improved using well annotated proteins from other species as references in the program Exonerate<xref rid="R41" ref-type="bibr">41</xref> and the outputs were manually curated. Specifically, the annotation of <italic>B. mori fzr</italic> was extended based on alignment of <italic>D. plexippus fzr</italic>; the annotation of <italic>B. mori fzy</italic> was altered based on alignment of <italic>Drosophila melanogaster</italic> and <italic>D. plexippus fzy</italic>; <italic>H. melpomene fzy</italic> was identified as part of the annotated gene HMEL017486 on scaffold HE671623 (Hmel v1.1) based on alignment of <italic>D. plexippus fzy;</italic> the <italic>Apis mellifera fzr</italic> annotation was altered based on alignment of <italic>D. melanogaster fzr</italic>; the annotation of <italic>Acyrthosiphon pisum fzr</italic> was altered based on alignment of <italic>D. melanogaster fzr.</italic> No one-to-one orthologues of <italic>D. melanogaster fzr2</italic> were found in any of the other arthropod genera, suggesting that this gene is <italic>Drosophila</italic> specific. Multiple sequence alignment of all the fizzy family proteins was then performed using the Expresso server<xref rid="R59" ref-type="bibr">59</xref> within T-coffee<xref rid="R60" ref-type="bibr">60</xref>, and this alignment was used to generate a neighbour joining tree in Geneious v8.1.7.</p></sec><sec id="S12"><title>Expression of <italic>H. melpomene cortex</italic> in <italic>D. melanogaster</italic> wings</title><p id="P40"><italic>D. melanogaster</italic> Cortex is known to generate an irregular microchaete phenotype when ectopically expressed in the posterior compartment of the adult fly wing<xref rid="R24" ref-type="bibr">24</xref>. We performed the same assay using <italic>H. melpomene cortex</italic> in order to test if this functionality was conserved. Following the methods of Swan and Schüpbach<xref rid="R24" ref-type="bibr">24</xref> a UAS-GAL4 construct was created using the coding region for the long isoform of <italic>Hm cortex</italic>, plus a <italic>Drosophila cortex</italic> version to act as positive control. The HA-tagged <italic>H. melpomene</italic> UAS<italic>-cortex</italic> expression construct was generated using cDNA reverse transcribed (Revert-Aid, Thermo-Scientific) from RNA extracted (Qiagen RNeasy) from pre-ommochrome pupal wing material. An HA-tagged <italic>D.melanogaster</italic> UAS-<italic>cortex</italic> version was also constructed, following the methods of Swan and Schüpbach, (2007). Expression was driven by hsp70 promoter. Constructs were injected into ϕC31-attP40 flies (#25709, Bloomington stock centre, Indiana; Cambridge University Genetics Department, UK, fly injection service) by site directed insertion into CII via an attB site in the construct. Homozygous transgenic flies were crossed with w,y’;en-GAL4;UAS-GFP (gift of M. Landgraf lab, Cambridge University Zoology Department) to drive expression in the engrailed posterior domain of the wing, and adult offspring wings photographed (<xref ref-type="fig" rid="F10">Extended Data Fig 6B-D</xref>). Expression of the construct was confirmed by IHC (standard <italic>Drosophila</italic> protocol) of final instar larval wing discs using mouse anti-HA and goat anti-mouse alexa-fluor 568 secondary antibodies (Abcam), imaged by Leica SP5 confocal. Successful expression of <italic>Hm_Cortex</italic> was confirmed by IHC against an HA tag inserted at the N terminal of either protein (<xref ref-type="fig" rid="F10">Extended Data Fig 6E</xref>).</p></sec></sec><sec sec-type="extended-data" id="S13"><title>Extended Data</title><fig id="F5" orientation="portrait" position="anchor"><label>Extended Data Figure 1</label><caption><p id="P41">A) Exons and splice variants of <italic>cortex</italic> in <italic>Hm</italic>. Orientation is reversed with respect to <xref ref-type="fig" rid="F2">figures 2</xref> and <xref ref-type="fig" rid="F4">4</xref>, with transcription going from left to right. SNPs showing the strongest associations with phenotype are shown with stars. B) Differential expression of two regions of <italic>cortex</italic> between <italic>Hm amaryllis</italic> and <italic>Hm aglaope</italic> whole hindwings (N=11 and N=10 respectively). Boxplots are standard (median; 75<sup>th</sup> and 25<sup>th</sup> percentiles; maximum and minimum excluding outliers – shown as discrete points) C) Expression of a <italic>cortex</italic> isoform lacking exon 3 is found in <italic>Hm aglaope</italic> but not <italic>Hm amaryllis</italic> hindwings. D) Expression of an isoform lacking exon 5 is found in <italic>Hm rosina</italic> but not <italic>Hm melpomene</italic> hindwings. Green triangles indicate predicted start codons and red triangles predicted stop codons, with usage dependent on which exons are present in the isoform. Schematics of the targeted exons are shown for each (q)RT-PCR product, black triangles indicate the position of the primers used in the assay.</p></caption><graphic xlink:href="emss-68288-f005"></graphic></fig><fig id="F6" orientation="portrait" position="anchor"><label>Extended Data Figure 2</label><caption><p id="P42">Alignments of <italic>de novo</italic> assembled fragments containing the top associated SNPs from <italic>Hm</italic> and related taxa short-read data. Identified indels do not show stronger associations with phenotype that those seen at SNPs (as shown in <xref ref-type="table" rid="T2">Extended Data Table 2</xref>), although some near-perfect associations are seen in fragment C. Black regions = missing data; yellow box = individuals with a hindwing yellow bar; blue box = individuals with a yellow forewing band.</p></caption><graphic xlink:href="emss-68288-f006"></graphic></fig><fig id="F7" orientation="portrait" position="anchor"><label>Extended Data Figure 3</label><caption><p id="P43">Sequencing of long-range PCR products and fosmids spanning <italic>cortex.</italic> A) Sequence read coverage from long-range PCR products across the <italic>cortex</italic> coding region from 2 <italic>Hm</italic> races. B) Minor allele frequency difference from these reads between <italic>Hm aglaope</italic> and <italic>Hm amaryllis</italic>. Exons of <italic>cortex</italic> are indicated by boxes, numbered as in <xref ref-type="fig" rid="F6">Extended Data Figure 2</xref>. C) Alignments of sequenced fosmids overlapping <italic>cortex</italic> from 3 <italic>Hm</italic> individuals of difference races. No major rearrangements are observed, nor any major differences in transposable element (TE) content between closely related races with different colour patterns (<italic>melpomene/rosina</italic> or <italic>amaryllis/aglaope</italic>). <italic>Hm amaryllis</italic> and <italic>rosina</italic> have the same phenotype, but do not share any TEs that are not present in the other races. Hm_BAC = BAC reference sequence, Hm_mel = <italic>melpomene</italic> from new unpublished assembly of <italic>Hm</italic> genome<xref rid="R51" ref-type="bibr">51</xref>, Hm_ros = <italic>rosina</italic> (2 different alleles were sequenced from this individual), Hm_ama = <italic>amaryllis</italic> (2 non-overlapping clones were sequenced in this individual), Hm_agla = <italic>aglaope</italic> (4 clones were sequenced in this individual 2 of which represent alternative alleles). Alignments were performed with Mauve: coloured bars represent homologous genomic regions. <italic>cortex</italic> is annotated in black above each clone. Variable TEs are shown as coloured bars below each clone: red = Metulj-like non-LTR, yellow = Helitron-like DNA, grey = other.</p></caption><graphic xlink:href="emss-68288-f007"></graphic></fig><fig id="F8" orientation="portrait" position="anchor"><label>Extended Data Figure 4</label><caption><p id="P44">Expression array results for additional stages, related to <xref ref-type="fig" rid="F4">Figure 4</xref>. A-G: comparisons between races (<italic>H. m. plesseni</italic> and <italic>H. m. malleti</italic>) for 3 wing regions. H-N: comparisons between proximal and distal forewing regions for each race. Significance values (-log10(p-value)) are shown separately for genes in the <italic>HmYb</italic> region from the gene array (A,D,F,H,K,M) and for the <italic>HmYb</italic> tiling array (B,E,G,I,L,N) for day 1 (A,B,H,I), day 5 (D,E,K,L) and day 7 (F,G,M,N) after pupation. The level of expression difference (log fold change) for tiling probes showing significant differences (p≤0.05) is shown for day 1 (C and J) with probes in known <italic>cortex</italic> exons shown in dark colours and probes elsewhere shown as pale colours.</p></caption><graphic xlink:href="emss-68288-f008"></graphic></fig><fig id="F9" orientation="portrait" position="anchor"><label>Extended Data Figure 5</label><caption><p id="P45">Alternative splicing of <italic>cortex</italic>. A) Amplification of the whole <italic>cortex</italic> coding region, showing the diversity of isoforms and variation between individuals. B) Differences in splicing of exon 3 between <italic>H. m. aglaope</italic> and <italic>H. m. amaryllis</italic>. Products amplified with a primer spanning the exon 2/4 junction at 3 developmental stages. The lower panel shows verification of this assay by amplification between exons 2 and 4 for the same final instar larval samples (replicated in <xref ref-type="fig" rid="F6">Extended Data Figure 2C</xref>) C) Lack of consistent differences between <italic>H. m. melpomene</italic> and <italic>H. m. rosina</italic> in splicing of exon 3. Top panel shows products amplified with a primer spanning the exon 2/4 junction, lower panel is the same samples amplified between exons 2 and 4. D) Differences in splicing of exon 5 between <italic>H. m. melpomene</italic> and <italic>H. m. rosina</italic>. Products amplified with a primer spanning the exon 4/6 junction at 3 developmental stages. E) Subset of samples from D amplified with primers between exons 4 and 6 for verification (middle, 24hr pupae samples are replicated in <xref ref-type="fig" rid="F6">Extended Data Figure 2D</xref>). F) Lack of consistent differences between <italic>H. m. aglaope</italic> and <italic>H. m. amaryllis</italic> in splicing of exon 5. Products amplified with a primer spanning the exon 4/6 junction. G) <italic>H. m. cythera</italic> also expresses the isoform lacking exon 5, while a pool of 6 <italic>H. m. malleti</italic> individuals do not. H) Expression of the isoform lacking exon 5 from an F2 <italic>H. m. melpomene</italic> x <italic>H. m. rosina</italic> cross. Individuals homozygous or heterozygous for the <italic>H. m. rosina HmYb</italic> allele express the isoform while those homozygous for the <italic>H. m. melpomene HmYb</italic> allele do not. I) Allele specific expression of isoforms with and without exon 5. Heterozygous individuals (indicated with blue and red stars) express only the <italic>H. m. rosina</italic> allele in the isoform lacking exon 5 (G at highlighted position), while they express both alleles in the isoform containing exon 5 (G/A at this position).</p></caption><graphic xlink:href="emss-68288-f009"></graphic></fig><fig id="F10" orientation="portrait" position="anchor"><label>Extended Data Figure 6</label><caption><p id="P46">Phylogeny of fizzy family proteins and effects of expressing <italic>cortex</italic> in the Drosophila wing. A) Neighbour joining phylogeny of Fizzy family proteins including functionally characterised proteins (in bold) from <italic>Saccharomyces cerevisiae, Homo sapiens</italic> and <italic>Drosophila melanogaster</italic> as well as copies from the basal metazoan <italic>Trichoplax adhaerens</italic> and a range of annotated arthropod genomes (<italic>Daphnia pulex, Acyrthosiphon pisum, Pediculus humanus, Apis mellifica, Nasonia vitripennis, Anopheles gambiae, Tribolium castaneum</italic>) including the lepidoptera <italic>H. melpomene</italic> (in blue), <italic>Danaus plexippus</italic> and <italic>Bombyx mori</italic>. Branch colours: dark blue, CDC20/fzy; light blue, CDH1/fzr/rap; red, lepidoptran cortex. B-E) Ectopic expression of <italic>cortex</italic> in <italic>Drosophila melanogaster</italic>. <italic>Drosophila cortex</italic> produces an irregular microchaete phenotype when expressed in the posterior compartment of the fly wing (C) whereas <italic>Heliconius cortex</italic> does not (D), when compared to no expression (B). A, anterior; P, posterior. Successful <italic>Heliconius cortex</italic> expression was confirmed by anti-HA IHC in the last instar <italic>Drosophila</italic> larva wing imaginal disc (D, red), with DAPI staining in blue.</p></caption><graphic xlink:href="emss-68288-f010"></graphic></fig><table-wrap id="T1" position="anchor" orientation="landscape"><label>Extended Data Table 1</label><caption><p id="P47">Genes in the <italic>Yb</italic> region and evidence for wing patterning control in <italic>Heliconius</italic></p><p id="P48">Yb<sup>I</sup>, within the previously mapped <italic>Yb</italic> interval<xref rid="R12" ref-type="bibr">12</xref>. Sb<sup>I</sup>, within the previously mapped <italic>Sb</italic> interval<xref rid="R12" ref-type="bibr">12</xref>. <italic>Sb</italic> controls a white/yellow hindwing margin and is not investigated in this study. The <italic>N</italic> locus has not been fine-mapped previously. A<sup>Yb</sup>, number of above background SNPs associated with the hindwing yellow bar in this study. A<sup>N</sup>, number of above background SNPs associated with the forewing yellow band in this study. E<sup>1</sup>, detected as differentially expressed between <italic>Hm aglaope</italic> and <italic>amaryllis</italic> from RNAseq data in this study (<xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>). E<sup>gw</sup>, detected as differentially expressed between forewing regions in the gene array in this study. E<sup>gr</sup>, detected as differentially expressed between <italic>Hm plesseni</italic> and <italic>malleti</italic> in in the gene array in this study. E<sup>tw</sup>, numbers of probes showing differential expression between forewing regions in the tilling array in this study. E<sup>tr</sup>, numbers of probes showing differential expression between <italic>Hm plesseni</italic> and <italic>malleti</italic> in in the tiling array in this study. Cr<sup>I</sup>, within the previously mapped <italic>HeCr</italic> interval<xref rid="R11" ref-type="bibr">11</xref>. A<sup><italic>pet</italic></sup>, number of SNPs fixed for the alternative allele in <italic>He demophoon</italic>. A<sup><italic>fav</italic></sup>, number of SNPs fixed for the alternative allele in <italic>He favorinus</italic>. P<sup>I</sup>, within the previously mapped P interval<xref rid="R13" ref-type="bibr">13</xref>. A<sup><italic>bic</italic></sup>, number of above background SNPs associated with the <italic>Hn bicoloratus</italic> phenotype in this study.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1"></th><th align="left" rowspan="1" colspan="1"></th><th align="left" rowspan="1" colspan="1"></th><th align="left" colspan="9" rowspan="1"><italic>Heliconius melpomene</italic></th><th align="left" colspan="3" rowspan="1"><italic>H. erato</italic></th><th align="left" colspan="2" rowspan="1"><italic>Hn</italic></th></tr><tr><th align="left" rowspan="1" colspan="1"></th><th align="left" rowspan="1" colspan="1"></th><th align="left" rowspan="1" colspan="1"></th><th align="left" colspan="14" rowspan="1"><hr></hr></th></tr><tr><th align="left" rowspan="1" colspan="1"><italic>Hm</italic> gene ID</th><th align="left" rowspan="1" colspan="1"><italic>He</italic> gene ID</th><th align="left" rowspan="1" colspan="1">Putative gene name</th><th align="left" rowspan="1" colspan="1">Yb<sup>I</sup></th><th align="left" rowspan="1" colspan="1">Sb<sup>I</sup></th><th align="left" rowspan="1" colspan="1">A<sup>Yb</sup></th><th align="left" rowspan="1" colspan="1">A<sup>N</sup></th><th align="left" rowspan="1" colspan="1">E<sup>1</sup></th><th align="left" rowspan="1" colspan="1">E<sup>gw</sup></th><th align="left" rowspan="1" colspan="1">E<sup>gr</sup></th><th align="left" rowspan="1" colspan="1">E<sup>tw</sup></th><th align="left" rowspan="1" colspan="1">E<sup>tr</sup></th><th align="left" rowspan="1" colspan="1">Cr<sup>I</sup></th><th align="left" rowspan="1" colspan="1">A<sup><italic>pet</italic></sup></th><th align="left" rowspan="1" colspan="1">A<sup><italic>fav</italic></sup></th><th align="left" rowspan="1" colspan="1">P<sup>I</sup></th><th align="left" rowspan="1" colspan="1">A<sup><italic>bic</italic></sup></th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1">HM00002</td><td align="left" rowspan="1" colspan="1">HERA000036</td><td align="left" rowspan="1" colspan="1">Acylpeptide hydrolase</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00003</td><td align="left" rowspan="1" colspan="1">HERA000037</td><td align="left" rowspan="1" colspan="1">HM00003</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00004</td><td align="left" rowspan="1" colspan="1">HERA000038</td><td align="left" rowspan="1" colspan="1">Trehalase-1B</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00006</td><td align="left" rowspan="1" colspan="1">HERA000038.1</td><td align="left" rowspan="1" colspan="1">Trehalase-1A</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00007</td><td align="left" rowspan="1" colspan="1">HERA000039</td><td align="left" rowspan="1" colspan="1">B9 protein</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00008</td><td align="left" rowspan="1" colspan="1">HERA000040</td><td align="left" rowspan="1" colspan="1">HM00008</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00010</td><td align="left" rowspan="1" colspan="1">HERA000041</td><td align="left" rowspan="1" colspan="1">WD40 repeat domain 85</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00012</td><td align="left" rowspan="1" colspan="1">HERA000042</td><td align="left" rowspan="1" colspan="1">CG2519</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00013</td><td align="left" rowspan="1" colspan="1">HERA000045</td><td align="left" rowspan="1" colspan="1">Unkempt</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00014</td><td align="left" rowspan="1" colspan="1">HERA000046</td><td align="left" rowspan="1" colspan="1">Histone H3</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00015</td><td align="left" rowspan="1" colspan="1">HERA000047</td><td align="left" rowspan="1" colspan="1">HM00015</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00016</td><td align="left" rowspan="1" colspan="1">HERA000048</td><td align="left" rowspan="1" colspan="1">HM00016</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00017</td><td align="left" rowspan="1" colspan="1">HERA000049</td><td align="left" rowspan="1" colspan="1">RecQ Helicase</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00018</td><td align="left" rowspan="1" colspan="1">HERA000051</td><td align="left" rowspan="1" colspan="1">HM00018</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00019</td><td align="left" rowspan="1" colspan="1">HERA000052</td><td align="left" rowspan="1" colspan="1">BmSuc2</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00020</td><td align="left" rowspan="1" colspan="1">HERA000053</td><td align="left" rowspan="1" colspan="1">CG5796</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00021</td><td align="left" rowspan="1" colspan="1">HERA000054</td><td align="left" rowspan="1" colspan="1">HM00021</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00022</td><td align="left" rowspan="1" colspan="1">HERA000055</td><td align="left" rowspan="1" colspan="1">Enoyl-CoA hydratase</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00023</td><td align="left" rowspan="1" colspan="1">HERA000056</td><td align="left" rowspan="1" colspan="1">ATP binding protein</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00024</td><td align="left" rowspan="1" colspan="1">HERA000057</td><td align="left" rowspan="1" colspan="1">HM00024</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00025</td><td align="left" rowspan="1" colspan="1">HERA000059</td><td align="left" rowspan="1" colspan="1">cortex</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1">56</td><td align="left" rowspan="1" colspan="1">74</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1">603</td><td align="left" rowspan="1" colspan="1">1796</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">99</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1">51</td></tr><tr><td align="left" rowspan="1" colspan="1">HM00026</td><td align="left" rowspan="1" colspan="1">HERA000077</td><td align="left" rowspan="1" colspan="1">Poly(A)-specific ribonuclease (parn)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1">10</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">34</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00027</td><td align="left" rowspan="1" colspan="1">HERA000079</td><td align="left" rowspan="1" colspan="1">CG31320</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00028</td><td align="left" rowspan="1" colspan="1">HERA000080</td><td align="left" rowspan="1" colspan="1">ARP-like</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00029</td><td align="left" rowspan="1" colspan="1">HERA000081</td><td align="left" rowspan="1" colspan="1">CG4692</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00030</td><td align="left" rowspan="1" colspan="1">HERA000082</td><td align="left" rowspan="1" colspan="1">Proteasome 26S non ATPase subunit 4</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00031</td><td align="left" rowspan="1" colspan="1">HERA000083</td><td align="left" rowspan="1" colspan="1">HM00031</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00032</td><td align="left" rowspan="1" colspan="1">HERA000084</td><td align="left" rowspan="1" colspan="1">Zinc phosphodiesterase</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00033</td><td align="left" rowspan="1" colspan="1">HERA000085</td><td align="left" rowspan="1" colspan="1">Serine/threonine-protein kinase (LMTK1)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00034</td><td align="left" rowspan="1" colspan="1">HERA000086</td><td align="left" rowspan="1" colspan="1">WD repeat domain 13 (Wdr13)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">4</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00035</td><td align="left" rowspan="1" colspan="1">HERA000087</td><td align="left" rowspan="1" colspan="1">Domeless</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00036</td><td align="left" rowspan="1" colspan="1">HERA000061</td><td align="left" rowspan="1" colspan="1">WAS protein family homologue 1</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">36</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">37</td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00038</td><td align="left" rowspan="1" colspan="1">HERA000062</td><td align="left" rowspan="1" colspan="1">Lethal (2) k05819 CG3054</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00039</td><td align="left" rowspan="1" colspan="1">HERA000064</td><td align="left" rowspan="1" colspan="1">Mitogen-activated protein kinase (MAPKK)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00040</td><td align="left" rowspan="1" colspan="1">HERA000064.1</td><td align="left" rowspan="1" colspan="1">DNA excision repair protein ERCC-6</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00041</td><td align="left" rowspan="1" colspan="1">HERA000065</td><td align="left" rowspan="1" colspan="1">Penguin</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00042</td><td align="left" rowspan="1" colspan="1">HERA000066</td><td align="left" rowspan="1" colspan="1">Thymidylate kinase</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00043</td><td align="left" rowspan="1" colspan="1">HERA000067</td><td align="left" rowspan="1" colspan="1">Caspase-activated DNase</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00044</td><td align="left" rowspan="1" colspan="1">HERA000068</td><td align="left" rowspan="1" colspan="1">Regulator of ribosome biosynthesis</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00045</td><td align="left" rowspan="1" colspan="1">HERA000069</td><td align="left" rowspan="1" colspan="1">CG12659</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00046</td><td align="left" rowspan="1" colspan="1">HERA000070</td><td align="left" rowspan="1" colspan="1">CG33505</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00047</td><td align="left" rowspan="1" colspan="1">HERA000071</td><td align="left" rowspan="1" colspan="1">Sr protein</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00048</td><td align="left" rowspan="1" colspan="1">HERA000073</td><td align="left" rowspan="1" colspan="1">HM00048</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00049</td><td align="left" rowspan="1" colspan="1">HERA000073.1</td><td align="left" rowspan="1" colspan="1">HM00049</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00050</td><td align="left" rowspan="1" colspan="1">HERA000074</td><td align="left" rowspan="1" colspan="1">Shuttle craft</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00051</td><td align="left" rowspan="1" colspan="1">HERA000075</td><td align="left" rowspan="1" colspan="1">HM00051</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">HM00052</td><td align="left" rowspan="1" colspan="1">HERA000076</td><td align="left" rowspan="1" colspan="1">HM00052</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">x</td><td align="left" rowspan="1" colspan="1"></td></tr></tbody></table></table-wrap><table-wrap id="T2" position="anchor" orientation="landscape"><label>Extended Data Table 2</label><caption><p id="P49">Locations of fixed/above background SNPs and differentially expressed (DE) tiling array probes</p></caption><table frame="void" rules="none"><tbody><tr><td align="left" rowspan="1" colspan="1"></td><td colspan="10" align="left" valign="top" rowspan="1"><hr></hr></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" valign="top" colspan="2" rowspan="1"><bold>Positions of SNPs in the <italic>He</italic> and <italic>Hn</italic> cortex Other association analyses</bold></td><td align="left" valign="bottom" rowspan="1" colspan="1"><italic>cortex</italic> coding exons</td><td align="left" valign="bottom" rowspan="1" colspan="1"><italic>cortex</italic> UTR exons</td><td align="left" valign="bottom" rowspan="1" colspan="1"><italic>cortex</italic> introns (nonTE)</td><td align="left" valign="bottom" rowspan="1" colspan="1"><italic>cortex</italic> flanking intergenic (nonTE)</td><td align="left" valign="bottom" rowspan="1" colspan="1">TEs</td><td align="left" valign="bottom" rowspan="1" colspan="1">Other genes (exons or introns)</td><td align="left" valign="bottom" rowspan="1" colspan="1">Other intergenic</td><td align="left" valign="bottom" rowspan="1" colspan="1">Total</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td colspan="10" align="left" valign="top" rowspan="1"><hr></hr></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" colspan="2" rowspan="1"><italic>erato favorinus</italic> fixed</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">96</td><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">108</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" colspan="2" rowspan="1"><italic>erato demophoon</italic> fixed</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">6</td><td align="left" rowspan="1" colspan="1">15</td></tr><tr><td align="left" rowspan="1" colspan="1"></td><td align="left" colspan="2" rowspan="1"><italic>numata bicoloratus</italic> above background</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">3</td><td align="left" rowspan="1" colspan="1">47</td><td align="left" rowspan="1" colspan="1">16</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">69</td></tr><tr><td colspan="11" align="left" valign="top" rowspan="1"><hr></hr></td></tr><tr><td align="left" valign="top" colspan="3" rowspan="1"><bold>Positions of DE tiling array probes</bold></td><td align="left" valign="bottom" rowspan="1" colspan="1">Known <italic>cortex</italic> coding exons</td><td align="left" valign="bottom" rowspan="1" colspan="1"><italic>cortex</italic> UTR exons</td><td align="left" valign="bottom" rowspan="1" colspan="1"><italic>cortex</italic> introns (nonTE)</td><td align="left" valign="bottom" rowspan="1" colspan="1">miRNAs</td><td align="left" valign="bottom" rowspan="1" colspan="1">TEs</td><td align="left" valign="bottom" rowspan="1" colspan="1">Other gene exons</td><td align="left" valign="bottom" rowspan="1" colspan="1">Other introns/intergenic</td><td align="left" valign="bottom" rowspan="1" colspan="1">Total</td></tr><tr><td colspan="11" align="left" valign="top" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="6" style="=-webkit-transform:rotate(-90deg);-moz-transform:rotate(-90deg);-o-transform:rotate(-90deg);transform:rotate(-90deg);padding: 10px" colspan="1">Day3</td><td align="center" rowspan="3" style="=-webkit-transform:rotate(-90deg);-moz-transform:rotate(-90deg);-o-transform:rotate(-90deg);transform:rotate(-90deg);padding: 10px" colspan="1">malleti vs<break></break>plesseni</td><td align="right" rowspan="1" colspan="1">Forewing proximal</td><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">7</td><td align="left" rowspan="1" colspan="1">323</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">13</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">7</td><td align="left" rowspan="1" colspan="1">359</td></tr><tr><td align="right" rowspan="1" colspan="1">Forewing distal</td><td align="left" rowspan="1" colspan="1">12</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">327</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">8</td><td align="left" rowspan="1" colspan="1">357</td></tr><tr><td align="right" rowspan="1" colspan="1">Hindwing</td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">14</td><td align="left" rowspan="1" colspan="1">378</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">9</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">6</td><td align="left" rowspan="1" colspan="1">413</td></tr><tr><td colspan="10" align="left" valign="top" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="2" style="=-webkit-transform:rotate(-90deg);-moz-transform:rotate(-90deg);-o-transform:rotate(-90deg);transform:rotate(-90deg);padding: 10px" colspan="1">Proximal vs<break></break>distal</td><td align="right" rowspan="1" colspan="1">malleti</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">68</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">12</td><td align="left" rowspan="1" colspan="1">81</td></tr><tr><td align="right" rowspan="1" colspan="1">plesseni</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">4</td><td align="left" rowspan="1" colspan="1">222</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">10</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">4</td><td align="left" rowspan="1" colspan="1">242</td></tr><tr><td colspan="11" align="left" valign="top" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="6" style="=-webkit-transform:rotate(-90deg);-moz-transform:rotate(-90deg);-o-transform:rotate(-90deg);transform:rotate(-90deg);padding: 10px" colspan="1">Day1</td><td align="center" rowspan="3" style="=-webkit-transform:rotate(-90deg);-moz-transform:rotate(-90deg);-o-transform:rotate(-90deg);transform:rotate(-90deg);padding: 10px" colspan="1">malleti vs<break></break>plesseni</td><td align="right" rowspan="1" colspan="1">Forewing proximal</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">22</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">3</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">7</td><td align="left" rowspan="1" colspan="1">33</td></tr><tr><td align="right" rowspan="1" colspan="1">Forewing distal</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">3</td><td align="left" rowspan="1" colspan="1">116</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">9</td><td align="left" rowspan="1" colspan="1">5</td><td align="left" rowspan="1" colspan="1">112</td><td align="left" rowspan="1" colspan="1">248</td></tr><tr><td align="right" rowspan="1" colspan="1">Hindwing</td><td align="left" rowspan="1" colspan="1">9</td><td align="left" rowspan="1" colspan="1">10</td><td align="left" rowspan="1" colspan="1">500</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">20</td><td align="left" rowspan="1" colspan="1">2</td><td align="left" rowspan="1" colspan="1">80</td><td align="left" rowspan="1" colspan="1">622</td></tr><tr><td colspan="10" align="left" valign="top" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="2" style="=-webkit-transform:rotate(-90deg);-moz-transform:rotate(-90deg);-o-transform:rotate(-90deg);transform:rotate(-90deg);padding: 12px" colspan="1">Proximal vs<break></break>distal</td><td align="right" rowspan="1" colspan="1">malleti</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">12</td><td align="left" rowspan="1" colspan="1">95</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">108</td></tr><tr><td align="right" rowspan="1" colspan="1">plesseni</td><td align="left" rowspan="1" colspan="1">3</td><td align="left" rowspan="1" colspan="1">3</td><td align="left" rowspan="1" colspan="1">81</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">99</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">186</td></tr><tr><td colspan="12" align="left" valign="top" rowspan="1"><hr></hr></td></tr></tbody></table></table-wrap><table-wrap id="T3" position="anchor" orientation="landscape"><label>Extended Data Table 3</label><caption><p id="P50">SNPs showing the strongest phenotypic associations in the <italic>H. melpomene/timareta</italic>/silvaniform comparison.</p><p id="P51">*downstream of <italic>cortex</italic>, †between exons 3 and 4 of <italic>cortex</italic>, ‡upstream of <italic>cortex</italic>, §between exons U4 and U3 of <italic>cortex</italic>. None of these SNPs are within known TEs. Colours show phenotypic associations: yellow = yellow hindwing bar; pink = no yellow hindwing bar; green = yellow forewing band; blue = no yellow forewing band; grey = allele does not match expected pattern.</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1">Species</th><th align="left" rowspan="1" colspan="1">Race</th><th align="left" rowspan="1" colspan="1">Sample code</th><th align="left" rowspan="1" colspan="1">HW bar</th><th align="left" rowspan="1" colspan="1">SNP pos 457083† (p=6.07E-10)</th><th align="left" rowspan="1" colspan="1">SNP pos 439063* (p=1.72E-09 )</th><th align="left" rowspan="1" colspan="1">SNP pos 602131‡ (p=2.42E-09)</th><th align="left" rowspan="1" colspan="1">SNP pos 457056† (p=2.42E-09)</th><th align="left" rowspan="1" colspan="1">FW band</th><th align="left" rowspan="1" colspan="1">SNP pos 584465§ (p=1.37E-07)</th><th align="left" rowspan="1" colspan="1">SNP pos 584418§ (p=1.41E-07)</th><th align="left" rowspan="1" colspan="1">SNP pos 584633§ (p=2.10E-07)</th><th align="left" rowspan="1" colspan="1">SNP pos 603344‡ (p=2.19E-07)</th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>aglaope</italic></td><td align="left" rowspan="1" colspan="1">09-246</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">A/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">A/A</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>aglaope</italic></td><td align="left" rowspan="1" colspan="1">09-267</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>aglaope</italic></td><td align="left" rowspan="1" colspan="1">09-268</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>aglaope</italic></td><td align="left" rowspan="1" colspan="1">09-357</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">G/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>aglaope</italic></td><td align="left" rowspan="1" colspan="1">aglaope.1</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" rowspan="1" colspan="1">N/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">C/T</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">T/A</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">T/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>amandus</italic></td><td align="left" rowspan="1" colspan="1">2221</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">A/A</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>amandus</italic></td><td align="left" rowspan="1" colspan="1">2228</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">A/A</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">C/T</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">T/A</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">T/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>amaryllis</italic></td><td align="left" rowspan="1" colspan="1">09-332</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>amaryllis</italic></td><td align="left" rowspan="1" colspan="1">09-333</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>amaryllis</italic></td><td align="left" rowspan="1" colspan="1">09-075</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>amaryllis</italic></td><td align="left" rowspan="1" colspan="1">09-079</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>amaryllis</italic></td><td align="left" rowspan="1" colspan="1">amaryllis.1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>bellula</italic></td><td align="left" rowspan="1" colspan="1">228</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" rowspan="1" colspan="1">NA</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>bellula</italic></td><td align="left" rowspan="1" colspan="1">231</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">G/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">C/T</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">T/A</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">T/C</td><td align="left" rowspan="1" colspan="1">NA</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>cythera</italic></td><td align="left" rowspan="1" colspan="1">2856</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>cythera</italic></td><td align="left" rowspan="1" colspan="1">2857</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" rowspan="1" colspan="1">NA</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>maIleti</italic></td><td align="left" rowspan="1" colspan="1">17162</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>melpomene</italic></td><td align="left" rowspan="1" colspan="1">18038</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>melpomene</italic></td><td align="left" rowspan="1" colspan="1">18097</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" rowspan="1" colspan="1">NA</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>melpomene</italic></td><td align="left" rowspan="1" colspan="1">m0.06</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>melpomene</italic></td><td align="left" rowspan="1" colspan="1">gen_ref</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>melpomene</italic></td><td align="left" rowspan="1" colspan="1">13435</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>melpomene</italic></td><td align="left" rowspan="1" colspan="1">9315</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>melpomene</italic></td><td align="left" rowspan="1" colspan="1">9316</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>melpomene</italic></td><td align="left" rowspan="1" colspan="1">9317</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>plesseni</italic></td><td align="left" rowspan="1" colspan="1">9156</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" rowspan="1" colspan="1">NA</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>plesseni</italic></td><td align="left" rowspan="1" colspan="1">16293</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" rowspan="1" colspan="1">NA</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>rosina</italic></td><td align="left" rowspan="1" colspan="1">rosina.1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>rosina</italic></td><td align="left" rowspan="1" colspan="1">2071</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>rosina</italic></td><td align="left" rowspan="1" colspan="1">531</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>rosina</italic></td><td align="left" rowspan="1" colspan="1">533</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" rowspan="1" colspan="1">NA</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>rosina</italic></td><td align="left" rowspan="1" colspan="1">546</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>thelixopeia</italic></td><td align="left" rowspan="1" colspan="1">13566</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">C/T</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">T/A</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">T/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>vulcanus</italic></td><td align="left" rowspan="1" colspan="1">14632</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" rowspan="1" colspan="1">NA</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. melpomene</italic></td><td align="left" rowspan="1" colspan="1"><italic>vulcanus</italic></td><td align="left" rowspan="1" colspan="1">519</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>florencia</italic></td><td align="left" rowspan="1" colspan="1">2403</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>florencia</italic></td><td align="left" rowspan="1" colspan="1">2406</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">A/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>florencia</italic></td><td align="left" rowspan="1" colspan="1">2407</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">A/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>florencia</italic></td><td align="left" rowspan="1" colspan="1">2410</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>timareta</italic></td><td align="left" rowspan="1" colspan="1">8533</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">C/T</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">T A</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">T/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>timareta</italic></td><td align="left" rowspan="1" colspan="1">9184</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>timareta</italic></td><td align="left" rowspan="1" colspan="1">8520</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>timareta</italic></td><td align="left" rowspan="1" colspan="1">8523</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>thelxinoe</italic></td><td align="left" rowspan="1" colspan="1">09-312</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>thelxinoe</italic></td><td align="left" rowspan="1" colspan="1">8624</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>thelxinoe</italic></td><td align="left" rowspan="1" colspan="1">8628</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. timareta</italic></td><td align="left" rowspan="1" colspan="1"><italic>thelxinoe</italic></td><td align="left" rowspan="1" colspan="1">8631</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FECCCB" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">A/A</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. elevatus</italic></td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1">09-343</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">A/T</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">G/G</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">1</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">C/T</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#99CDFF" rowspan="1" colspan="1">T/T</td></tr><tr><td align="left" rowspan="1" colspan="1"><italic>H. pardalinus</italic></td><td align="left" rowspan="1" colspan="1"><italic>sergestus</italic></td><td align="left" rowspan="1" colspan="1">09-326</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#DADADA" rowspan="1" colspan="1">A/A</td><td align="left" style="background-color:#FEFF99" rowspan="1" colspan="1">A/A</td><td align="left" rowspan="1" colspan="1">NA</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">0</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">C/C</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" style="background-color:#CDFFCC" rowspan="1" colspan="1">T/T</td><td align="left" rowspan="1" colspan="1">NA</td></tr></tbody></table></table-wrap><table-wrap id="T4" position="anchor" orientation="landscape"><label>Extended Data Table 4</label><caption><p id="P52">Transposable Elements (TEs) found within the <italic>Yb</italic> region.</p></caption><table frame="hsides" rules="none"><tbody><tr><td align="left" valign="top" colspan="5" rowspan="1">Unique Occurrences<hr></hr></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">BAC</td><td align="left" valign="top" rowspan="1" colspan="1">mel</td><td align="left" valign="top" rowspan="1" colspan="1">ros</td><td align="left" valign="top" rowspan="1" colspan="1">ama</td><td align="left" valign="top" rowspan="1" colspan="1">agl</td><td align="left" valign="top" rowspan="1" colspan="1">No.</td><td align="left" valign="top" rowspan="1" colspan="1">TE name</td><td align="left" valign="top" rowspan="1" colspan="1">Superfamily</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Type</td></tr><tr><td colspan="10" align="left" valign="top" rowspan="1"><hr></hr></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">BEL-1</td><td align="left" valign="top" rowspan="1" colspan="1">BEL</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">CR1-2</td><td align="left" valign="top" rowspan="1" colspan="1">Jockey</td><td align="left" valign="top" rowspan="1" colspan="1">LINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">Daphne-1</td><td align="left" valign="top" rowspan="1" colspan="1">Jockey</td><td align="left" valign="top" rowspan="1" colspan="1">LINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">Daphne-6</td><td align="left" valign="top" rowspan="1" colspan="1">Jockey</td><td align="left" valign="top" rowspan="1" colspan="1">LINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">DNA-like-8</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td colspan="10" align="left" valign="top" rowspan="1"><hr></hr></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-like-14</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_A</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">4</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-like-12</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_A</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">5</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-like-12b</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_A</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">7</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-like-4a</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_A</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-like-4b</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_A</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-N2</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_A</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">3</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-like-7</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_A</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">5</td><td align="left" valign="top" rowspan="1" colspan="1">3</td><td align="left" valign="top" rowspan="1" colspan="1">3</td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1">16</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-like-6a</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_B</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-like-6b</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_B</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-like-11</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_B</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">11</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-like-15</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_B</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">6</td><td align="left" valign="top" rowspan="1" colspan="1">5</td><td align="left" valign="top" rowspan="1" colspan="1">3</td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">18</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron-like-5</td><td align="left" valign="top" rowspan="1" colspan="1">Helitron_B</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td colspan="10" align="left" valign="top" rowspan="1"><hr></hr></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1">Hmel_Unknown_50</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1">Hmel_Unknown_174a/b</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">Hmel_Unknown_187b</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1">Hmel_Unknown_230</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">Hmel_Unknown_234a</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">Hmel_Unknown_236a</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">Jockey-4</td><td align="left" valign="top" rowspan="1" colspan="1">Jockey</td><td align="left" valign="top" rowspan="1" colspan="1">LINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">LTR-3_gypsy</td><td align="left" valign="top" rowspan="1" colspan="1">Gypsy</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">Mariner-4</td><td align="left" valign="top" rowspan="1" colspan="1">Mariner/Tc1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td colspan="10" align="left" valign="top" rowspan="1"><hr></hr></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">3</td><td align="left" valign="top" rowspan="1" colspan="1">29</td><td align="left" valign="top" rowspan="1" colspan="1">Metulj-0</td><td align="left" valign="top" rowspan="1" colspan="1">Metulj</td><td align="left" valign="top" rowspan="1" colspan="1">SINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Metulj-1</td><td align="left" valign="top" rowspan="1" colspan="1">Metulj</td><td align="left" valign="top" rowspan="1" colspan="1">SINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Metulj-2</td><td align="left" valign="top" rowspan="1" colspan="1">Metulj</td><td align="left" valign="top" rowspan="1" colspan="1">SINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Metulj-3</td><td align="left" valign="top" rowspan="1" colspan="1">Metulj</td><td align="left" valign="top" rowspan="1" colspan="1">SINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Metulj-4</td><td align="left" valign="top" rowspan="1" colspan="1">Metulj</td><td align="left" valign="top" rowspan="1" colspan="1">SINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Metulj-5</td><td align="left" valign="top" rowspan="1" colspan="1">Metulj</td><td align="left" valign="top" rowspan="1" colspan="1">SINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Metulj-6</td><td align="left" valign="top" rowspan="1" colspan="1">Metulj</td><td align="left" valign="top" rowspan="1" colspan="1">SINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">Metulj-7</td><td align="left" valign="top" rowspan="1" colspan="1">Metulj</td><td align="left" valign="top" rowspan="1" colspan="1">SINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">nTc3-4</td><td align="left" valign="top" rowspan="1" colspan="1">Mariner/Tc1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">SINE-1</td><td align="left" valign="top" rowspan="1" colspan="1">SINE</td><td align="left" valign="top" rowspan="1" colspan="1">SINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td colspan="10" align="left" valign="top" rowspan="1"><hr></hr></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1">nMar-3</td><td align="left" valign="top" rowspan="1" colspan="1">Mariner/Tc1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">nMar-16</td><td align="left" valign="top" rowspan="1" colspan="1">Mariner/Tc1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">nMar-12/20</td><td align="left" valign="top" rowspan="1" colspan="1">Mariner/Tc1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">nPIF-3</td><td align="left" valign="top" rowspan="1" colspan="1">PIF/Harbinger</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">nTc3-2</td><td align="left" valign="top" rowspan="1" colspan="1">Mariner/Tc1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1">nTc3-3</td><td align="left" valign="top" rowspan="1" colspan="1">Mariner/Tc1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">DNA transposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1">R4-1</td><td align="left" valign="top" rowspan="1" colspan="1">R2</td><td align="left" valign="top" rowspan="1" colspan="1">LINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">6</td><td align="left" valign="top" rowspan="1" colspan="1">Rep-1</td><td align="left" valign="top" rowspan="1" colspan="1">REP</td><td align="left" valign="top" rowspan="1" colspan="1">LINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">4</td><td align="left" valign="top" rowspan="1" colspan="1">RTE-3</td><td align="left" valign="top" rowspan="1" colspan="1">RTE</td><td align="left" valign="top" rowspan="1" colspan="1">LINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">2</td><td align="left" valign="top" rowspan="1" colspan="1">RTE-11</td><td align="left" valign="top" rowspan="1" colspan="1">RTE</td><td align="left" valign="top" rowspan="1" colspan="1">LINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td colspan="10" align="left" valign="top" rowspan="1"><hr></hr></td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">3</td><td align="left" valign="top" rowspan="1" colspan="1">Zenon-1</td><td align="left" valign="top" rowspan="1" colspan="1">Jockey</td><td align="left" valign="top" rowspan="1" colspan="1">LINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr><tr><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1"></td><td align="left" valign="top" rowspan="1" colspan="1">1</td><td align="left" valign="top" rowspan="1" colspan="1">Zenon-3</td><td align="left" valign="top" rowspan="1" colspan="1">Jockey</td><td align="left" valign="top" rowspan="1" colspan="1">LINE</td><td align="left" valign="top" rowspan="1" colspan="1">Non-LTR retrotransposon</td></tr></tbody></table></table-wrap></sec><sec sec-type="supplementary-material" id="SM"><title>Supplementary Material</title><p id="P53"><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 content-type="local-data" id="SD2"><label>Supplementary Guide</label><media xlink:href="NIHMS68288-supplement-Supplementary_Guide.docx" mimetype="application" mime-subtype="octet-stream" orientation="portrait" xlink:type="simple" id="d37e6092" position="anchor"></media></supplementary-material><supplementary-material content-type="local-data" id="SD1"><label>Supplementary Data 1</label><media xlink:href="NIHMS68288-supplement-Supplementary_Data_1.pdf" mimetype="application" mime-subtype="pdf" orientation="portrait" xlink:type="simple" id="d37e6096" position="anchor"></media></supplementary-material><supplementary-material content-type="local-data" id="SD3"><label>Supplementary Information</label><media xlink:href="NIHMS68288-supplement-Supplementary_Information.pdf" mimetype="application" mime-subtype="pdf" orientation="portrait" xlink:type="simple" id="d37e6100" position="anchor"></media></supplementary-material><supplementary-material content-type="local-data" id="SD4"><label>Supplementary Table 1</label><media xlink:href="NIHMS68288-supplement-Supplementary_Table_1.xlsx" mimetype="application" mime-subtype="octet-stream" orientation="portrait" xlink:type="simple" id="d37e6104" position="anchor"></media></supplementary-material><supplementary-material content-type="local-data" id="SD5"><label>Supplementary Table 2</label><media xlink:href="NIHMS68288-supplement-Supplementary_Table_2.xlsx" mimetype="application" mime-subtype="octet-stream" orientation="portrait" xlink:type="simple" id="d37e6108" position="anchor"></media></supplementary-material></sec></body><back><ack id="S14"><title>Acknowledgements</title><p>We thank Christopher Saski, Clemson University, for assembly of the <italic>He</italic> BACs. Moises Abanto and Adriana Tapia assisted with raising butterflies. Thanks to Mathieu Chouteau, Jake Morris and Kanchon Dasmahapatra for providing larvae for <italic>in situ</italic> hybridisations. Anna Morrison, Robert Tetley, Sarah Carl and Hanna Wegener assisted with lab work at the University of Cambridge. Simon Baxter made the <italic>Hm</italic> fosmid libraries. We thank the governments of Colombia, Ecuador, Panama and Peru for permission to collect butterflies. This work was funded by a Leverhulme Trust award and BBSRC grant (H01439X/1) to CDJ, NSF grants (DEB 1257689, IOS 1052541) to WOM, an ERC starting grant to MJ and a French National Agency for Research (ANR) grant to VL (ANR-13-JSV7-0003-01). NJN is funded by a NERC fellowship (NE/K008498/1).</p></ack><fn-group><fn fn-type="con" id="FN1"><p id="P54"><bold>Author Contributions</bold> NJN performed the association analyses, 5’ RACE, RT-PCR, qRT-PCR and prepared the manuscript. NJN and CDJ co-ordinated the research. CP-D performed and analysed the microarray and RNAseq experiments. AW performed the <italic>Hn</italic> association analysis. MS assembled and annotated the <italic>HeCr</italic> BAC reference and the <italic>He</italic> alignments. SVS performed <italic>in situ</italic> hybridizations. RWRW performed the transgenic experiments and analysis of <italic>de novo</italic> assembled sequences and fosmids together with JJH. GW and LF initially identified splicing variants of <italic>cortex</italic>. LM performed crosses between <italic>Hm</italic> races. HH screened the <italic>HeCr</italic> BAC library. CS and RM provided samples. AD contributed to the <italic>Hm</italic> BAC sequencing and annotation. R-fC, MJ, VL, WOM and CDJ are PIs who obtained funding and led the project elements. All authors commented on the manuscript.</p></fn><fn id="FN2"><p id="P55"><bold>Author Information</bold> Short read sequence data generated for this study are available from ENA (<ext-link ext-link-type="uri" xlink:href="http://www.ebi.ac.uk/ena">http://www.ebi.ac.uk/ena</ext-link>) under study accession PRJEB8011 and PRJEB12740 (see <xref ref-type="supplementary-material" rid="SD4">Supplementary Table 1</xref> for previously published data accessions). The updated Cr contig is deposited in Genbank with accession KC469893. The assembled <italic>Hm</italic> fosmid sequences are deposited in Genbank with accessions KU514430-KU514438. The microarray data are deposited in GEO with accessions GSM1563402- GSM1563497. 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Left: phylogenetic relationships<xref rid="R29" ref-type="bibr">29</xref>. Right: chromosome maps with colour pattern intervals in grey, coloured bars represent markers used to assign homology<xref rid="R5" ref-type="bibr">5</xref>,<xref rid="R8" ref-type="bibr">8</xref>–<xref rid="R10" ref-type="bibr">10</xref>, the first and last genes from <xref ref-type="fig" rid="F2">Fig 2</xref> shown in red. In <italic>He</italic> the <italic>HeCr</italic> locus controls the yellow hind-wing bar phenotype (grey boxed races). In <italic>Hm</italic> it controls both the yellow hind-wing bar (<italic>HmYb,</italic> pink box) and the yellow forewing band (<italic>HmN,</italic> blue box). In <italic>Hn</italic> it modulates black, yellow and orange elements on both wings (<italic>HnP</italic>), producing phenotypes that mimic butterflies in the genus <italic>Melinaea</italic>. Morphs/races of <italic>Heliconius</italic> species included in this study are shown with names.</p></caption><graphic xlink:href="emss-68288-f001"></graphic></fig><fig id="F2" orientation="portrait" position="float"><label>Figure 2</label><caption><p>Association analyses across the genomic region known to contain major colour pattern loci in <italic>Heliconius.</italic> A) Association in <italic>He</italic> with the yellow hind-wing bar (n=45). Coloured SNPs are fixed for a unique state in <italic>He demophoon</italic> (orange) or <italic>He favorinus</italic> (purple). B) Genes in <italic>He</italic> with direct homologs in <italic>Hm</italic>. Genes are in different colours with exons (coding and UTRs) connected by a line. Grey bars are transposable elements. C) <italic>Hm</italic> genes and transposable elements: colours correspond to homologous <italic>He</italic> genes; MicroRNAs<xref rid="R30" ref-type="bibr">30</xref> in black. D) Association in the <italic>Hm/timareta</italic>/silvaniform group with the yellow hind-wing bar (red) and yellow forewing band (blue) (n=49). E) Association in <italic>Hn</italic> with the <italic>bicoloratus</italic> morph (n=26); inversion positions<xref rid="R13" ref-type="bibr">13</xref> shown below. In all cases black/dark coloured points are above the strongest associations found outside the colour pattern scaffolds (<italic>He</italic> p=1.63e-05; <italic>Hm</italic> p=2.03e-05 and p=2.58e-05; <italic>Hn</italic> p=6.81e-06).</p></caption><graphic xlink:href="emss-68288-f002"></graphic></fig><fig id="F3" orientation="portrait" position="float"><label>Figure 3</label><caption><p>Differential gene expression across the genomic region known to contain major colour pattern loci in <italic>Heliconius melpomene.</italic> Expression differences in day 3 pupae, for all genes in the <italic>Yb</italic> interval (A,D) and tiling probes spanning the central portion of the interval (B,C,E,F). Expression is compared between races for each wing region (A,B,C) and between proximal and distal forewing sections for each race (D,E,F). C and F: magnitude and direction of expression difference (log<sub>2</sub> fold-change) for tiling probes showing significant differences (p≤0.05); probes in known <italic>cortex</italic> exons shown in dark colours. Gene <italic>HM00052</italic> was differentially expressed between other races in RNA sequence data (<xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>) but is not differentially expressed here.</p></caption><graphic xlink:href="emss-68288-f003"></graphic></fig><fig id="F4" orientation="portrait" position="float"><label>Figure 4</label><caption><p><italic>In situ</italic> hybridisations of <italic>cortex</italic> in hind-wings of final instar larvae. B) <italic>Hn tarapotensis</italic>; adult wing shown in A, coloured points indicate landmarks, yellow arrows highlight adult pattern elements corresponding to the <italic>cortex</italic> staining. D) <italic>Hm rosina</italic>; adult wing shown in C, staining patterns in other <italic>Hm</italic> races (<italic>meriana</italic> and <italic>aglaope</italic>) appeared similar. The probe used was complementary to the <italic>cortex</italic> isoform with the longest open reading frame (also the most common, <xref ref-type="supplementary-material" rid="SD3">Supplementary Information</xref>).</p></caption><graphic xlink:href="emss-68288-f004"></graphic></fig></floats-group></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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