Wing coloration and pigment gradients in scales of pierid butterflies

Depending on the species, the individual scales of butterfly wings have a longitudinal gradient in structure and reflectance properties, as shown by scanning electron microscopy and microspectrophotometry. White scales of the male Small White, Pieris rapae crucivora, show a strong gradient in both the density in pigment granules and the reflectance. After pigment extraction by aqueous ammonia, scales of male P. r. crucivora closely resemble the unpigmented scales of female P. r. crucivora. Only a minor gradient exists in the white and orange scales of the male Orange Tip, Anthocharis cardamines. Pigment extraction of orange scales of A. cardamines causes bleaching. Partial bleaching transforms the scales so that they resemble certain scales of Phoebis philea that have a natural extreme gradient. Reflectance measurements on an artificial stack of two overlapping scales as well as on the scale stacks existing on intact and partially denuded wings of the Large White, Pieris brassicae, quantitatively demonstrate the reflectance enhancement by scale stacking.