Letter | Published:

doublesex is a mimicry supergene

Nature volume 507, pages 229232 (13 March 2014) | Download Citation


One of the most striking examples of sexual dimorphism is sex-limited mimicry in butterflies, a phenomenon in which one sex—usually the female—mimics a toxic model species, whereas the other sex displays a different wing pattern1. Sex-limited mimicry is phylogenetically widespread in the swallowtail butterfly genus Papilio, in which it is often associated with female mimetic polymorphism1,2,3. In multiple polymorphic species, the entire wing pattern phenotype is controlled by a single Mendelian ‘supergene’4. Although theoretical work has explored the evolutionary dynamics of supergene mimicry5,6,7,8,9, there are almost no empirical data that address the critical issue of what a mimicry supergene actually is at a functional level. Using an integrative approach combining genetic and association mapping, transcriptome and genome sequencing, and gene expression analyses, we show that a single gene, doublesex, controls supergene mimicry in Papilio polytes. This is in contrast to the long-held view that supergenes are likely to be controlled by a tightly linked cluster of loci4. Analysis of gene expression and DNA sequence variation indicates that isoform expression differences contribute to the functional differences between dsx mimicry alleles, and protein sequence evolution may also have a role. Our results combine elements from different hypotheses for the identity of supergenes, showing that a single gene can switch the entire wing pattern among mimicry phenotypes but may require multiple, tightly linked mutations to do so.

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We thank W. Wang for sharing genome sequence data, C. Robinett for providing the Dsx-DM monoclonal antibody, and E. Westerman, S. Nallu, M. Zhang, G. Garcia and N. Pierce for assistance and discussion. This project was funded by National Science Foundation grant DEB-1316037 to M.R.K.

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Author notes

    • K. Kunte
    •  & W. Zhang

    These authors contributed equally to this work.


  1. National Center for Biological Sciences, Tata Institute of Fundamental Research, Bengaluru 560065, India

    • K. Kunte
  2. Department of Ecology and Evolution, University of Chicago, Chicago, Illinois 60637, USA

    • W. Zhang
    • , A. Tenger-Trolander
    •  & M. R. Kronforst
  3. Committee on Evolutionary Biology, University of Chicago, Chicago, Illinois 60637, USA

    • D. H. Palmer
    •  & M. R. Kronforst
  4. Department of Ecology and Evolutionary Biology, Cornell University, Ithaca, New York 14853, USA

    • A. Martin
    •  & R. D. Reed
  5. Department of Biology, Boston University, Boston, Massachusetts 02215, USA

    • S. P. Mullen


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K.K. conceived the project and helped design the study, reared mapping families and samples for gene expression analysis and genome sequencing, performed bulk-segregant analysis and RAD mapping, and contributed to drafting the manuscript. W.Z. generated the reference genome sequences and transcriptome assemblies, performed association mapping, GWAS analysis, HKA tests, structural variant detection and linkage disequilibrium analyses, analysis of protein structure and synonymous/non-synonymous calculations, and contributed to drafting the manuscript. A.T.-T. assisted with butterfly husbandry, performed fine mapping, cDNA sequencing and qRT–PCR analyses. D.H.P. performed qRT–PCR analyses. A.M. and R.D.R. performed Dsx immunohistochemistry. S.P.M. helped design the project and contributed to drafting the manuscript. M.R.K. designed and directed the project, analysed data and wrote the manuscript.

Competing interests

The authors declare no competing financial interests.

Corresponding authors

Correspondence to K. Kunte or M. R. Kronforst.

Sequence data are available from NCBI SRA (SRP035394) and GenBank (KJ150616KJ150623).

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