Multiple approaches to study color pattern evolution in butterflies

The myriad colors and patterns on butterfly wings have caught the attention of biologists for well over a century. Today, with the advent of more sophisticated genetic and developmental tools, it is possible to identify and study the evolution of genes, gene networks, and the effect of the environment on the networks underlying wing color patterning. In addition, using molecular phylogenies and the comparative approach, it is possible to infer ancestral wing patterns, direction of evolutionary change, and occurrence of parallelism and convergence. Finally, the driving forces behind wing pattern evolution can be estimated using bioassay studies such as predator-prey and mate choice experiments. Here we review the different approaches to answer both proximate and ultimate questions about butterfly wing pattern evolution, and we highlight future research directions in a field that has the potential to become truly integrative.