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Background  The ability for serially homologous structures to acquire a separate identity has been primarily investigated for structures dependent on Hox gene input but is still incompletely understood in other systems. The fore and hindwings of butterflies are serially homologous structures as are the serially homologous eyespots that can decorate each of these wings. Eyespots can vary in number between fore and hindwings of the same individual and mutations of large effect can control the total number of eyespots that each of the wings displays. Here we investigate the genetics of a new spontaneous color pattern mutation,  Missing , that alters eyespot number in the nymphalid butterfly,  Bicyclus anynana . We further test the interaction of  Missing  with a previously described mutation,  Spotty , describe the developmental stage affected by  Missing , and test whether  Missing  is a mutant variant of the gene  Distal-less  via a linkage association study.    Results   Missing  removes or greatly reduces the size of two of the hindwing eyespots from the row of seven eyespots, with no detectable effect on the rest of the wing pattern. Offspring carrying a single  Missing  allele display intermediate sized eyespots at these positions.  Spotty  has the opposite effect of  Missing , i.e., it introduces two extra eyespots in homologous wing positions to those affected by  Missing , but on the forewing. When  Missing  is combined with  Spotty  the size of the two forewing eyespots decreases but the size of the hindwing spots stays the same, suggesting that these two mutations have a combined effect on the forewing such that  Missing  reduces eyespot size when in the presence of a  Spotty  mutant allele, but that  Spotty  has no effect on the hindwing.  Missing  prevents the complete differentiation of two of the eyespot foci on the hindwing. We found no evidence for any linkage between the  Distal-less  and  Missing  genes.    Conclusion  The spontaneous mutation  Missing  controls the differentiation of the signaling centers of a subset of the serial homologous eyespots present on both the fore and the hindwing in a dose-dependent fashion. The effect of  Missing  on the forewing, however, is only observed when the mutation  Spotty  introduces additional eyespots on this wing.  Spotty , on the other hand, controls the differentiation of eyespot centers only on the forewing.  Spotty , unlike  Missing , may be under Ubx gene regulation, since it affects a subset of eyespots on only one of the serially homologous wings.

The combined effect of two mutations that alter serially homologous color pattern elements on the fore and hindwings of a butterfly