Butterfly speciation and the distribution of gene effect sizes fixed during adaptation.

Mimicry has had a significant historical influence as a tractable system for studying adaptation and is known to play a role in speciation. Here, we discuss recent theoretical treatment of adaptive walks to local adaptive peaks and contrast this with the adaptive landscape of mimicry. Evolution of novel Müllerian mimicry patterns almost certainly involves substitution of a major mutation to provide an initial similarity to the model, such that major gene effects are expected to an even greater degree than for other adaptive traits. The likelihood of large adaptive peak shifts in mimicry evolution may therefore promote speciation. In addition, mimicry adaptive peaks are determined by the local abundance of particular patterns and may be more fluid than the case for other traits. It will therefore be of considerable interest to test empirically the distribution of effect sizes fixed during mimicry evolution. Here, we show the feasibility of this by presenting a preliminary quantitative trait locus (QTL) analysis of Heliconius colour patterns. This shows that a number of modifier loci of different effect sizes influence forewing band morphology. We also show multiple pleiotropic effects of major Heliconius patterning loci and discuss the likelihood of multiple substitutions at the same loci in pattern evolution, which would inflate the importance of major loci in QTL analysis of the gene effect sizes. Analyses such as these have the potential to uncover the genetic architecture of both within and between species adaptive differences.