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Heliconius  butterflies are an excellent system for understanding the genetic basis of phenotypic change. Here we document surprising diversity in the genetic control of a common phenotype. Two disjunct  H. erato  populations have each recruited the  Cr  and/or  Sd  loci that control similar yellow hindwing patterns, but the alleles involved partially complement one another indicating either multiple origins for the patterning alleles or developmental drift in genetic control of similar patterns. We show that in these  H. erato  populations  cr  and  sd  are epistatically interacting and that the parental origin of alleles can explain phenotypes of backcross individuals. In contrast, mimetic  H. melpomene  populations with identical phenotypes ( H. m. rosina  and  H. m. amaryllis ) do not show genetic complementation (F 1 s and F 2 s are phenotypically identical to parentals). Finally, we report hybrid female inviability in  H. m. melpomene  ×  H. m. rosina  crosses (previously only female infertility had been reported) and presence of standing genetic variation for alternative color alleles at the  Yb  locus in true breeding  H. melpomene melpomene  populations (expressed when in a different genomic background) that could be an important source of variation for the evolution of novel phenotypes or a result of developmental drift. Although recent work has emphasized the simple genetic control of wing pattern in  Heliconius , we show there is underlying complexity in the allelic variation and epistatic interactions between major patterning loci.

Partial Complementarity of the Mimetic Yellow Bar Phenotype in Heliconius Butterflies