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Recent studies indicate that relatively few genomic regions are repeatedly involved in the evolution of  Heliconius  butterfly wing patterns. Although this work demonstrates a number of cases where homologous loci underlie both convergent and divergent wing pattern change among different  Heliconius  species, it is still unclear exactly how many loci underlie pattern variation across the genus. To address this question for  Heliconius erato , we created fifteen independent crosses utilizing the four most distinct color pattern races and analyzed color pattern segregation across a total of 1271 F2 and backcross offspring. Additionally, we used the most variable brood, an F2 cross between  H. himera  and the east Ecuadorian  H. erato notabilis , to perform a quantitative genetic analysis of color pattern variation and produce a detailed map of the loci likely involved in the  H. erato  color pattern radiation. Using AFLP and gene based markers, we show that fewer major genes than previously envisioned control the color pattern variation in  H. erato . We describe for the first time the genetic architecture of  H. erato  wing color pattern by assessing quantitative variation in addition to traditional linkage mapping. In particular, our data suggest three genomic intervals modulate the bulk of the observed variation in color. Furthermore, we also identify several modifier loci of moderate effect size that contribute to the quantitative wing pattern variation. Our results are consistent with the two-step model for the evolution of mimetic wing patterns in  Heliconius  and support a growing body of empirical data demonstrating the importance of major effect loci in adaptive change.

Multi-Allelic Major Effect Genes Interact with Minor Effect QTLs to Control Adaptive Color Pattern Variation in Heliconius erato