The beautiful color patterns decorating buttery wings have been and continue to be inspirational for studies of a variety of biological issues, ranging from systematics and evolution to developmental genetics and biochemistry of pigmentation. The amazing diversity of these patterns, together with knowledge on their adaptive value and underlying developmental basis, make them a favorite system in the relatively new eld of evolutionary developmental biology, or evo-devo (Nijhout 1991; Beldade and Brakeeld 2002; McMillan, Monteiro, and Kapan 2002; Joron et al. 2006; Parchem, Perry, and Patel 2007; Wittkopp and Beldade 2009).