Polymorphic mimicry in Papilio dardanus: mosaic dominance, big effects, and origins

The mocker swallowtail, Papilio dardanus, has a female-limited polymorphic mimicry. This polymorphism is controlled by allelic variation at a single locus with at least 11 alleles. Many of the alternative morphs are accurate mimics of different species of distasteful butterflies. Geneticists have long been interested in the mechanism by which a single gene can have such diverse and profound effects on the phenotype and in the process by which these complex phenotypic effects could have evolved. Here we present the results of a morpho-metric analysis of the pleiotropic effects of the mimicry gene on the array of elements that makes up the overall pattern. We show that the patterns controlled by mimicking alleles are more variable and less internally correlated than those controlled by nonmimicking alleles, suggesting the two are subject to different degrees of selection and mutational variance. Analysis of the pleiotropic dominance of the alleles reveals a consistent pattern of dominance within a coevolved genetic background and a mosaic pattern of dominant and recessive effects (including overdominance) in a heterologous genetic background. The alleles of the mimicry gene have big effects on some pattern elements and small effects on others. When the array of big phenotypic effects of the mimicry gene is applied to the presumptive ancestral color pattern, it produces a reasonable resemblance to distasteful models and suggests the initial steps that may have produced the mimicry as well as the polymorphism.