The origin of new morphological characters is a long-standing problem in ev
olutionary biology. Novelties arise through changes in development, but the
nature of these changes is largely unknown. In butterflies, eyespots have
evolved as new pattern elements that develop from special organizers called
foci. Formation of these foci is associated with novel expression patterns
of the Hedgehog signaling protein, its receptor Patched, the transcription
factor Cubitus interruptus, and the engrailed target gene that break the c
onserved compartmental restrictions on this regulatory circuit in insect wi
ngs. Redeployment of preexisting regulatory circuits may be a general mecha
nism underlying the evolution of novelties.