Correlations between scale structure and pigmentation in butterfly wings.

We examined the correlation between color and structure of wing scales in the nymphalid butterflies Bicyclus anynana and Heliconius melpomene. All scales in B. anynana are rather similar in comparison to the clear structural differences of differently pigmented scales in H. melpomene. Where scale structural differences in H. melpomene are qualitative, they seem to be quantitative in B. anynana. There is a "gradient" in the density of some structural elements, the cross ribs, in the scales of B. anynana: black, gold, and brown scales show progressively lower cross rib density within an individual. There is, however, high individual variation in the absolute cross rib densities (i.e., scales with a particular color and cross rib density in one individual may have a different color but similar density in another individual). By ectopically inducing color pattern during early pupal development, we examined whether a scale's color and its microstructure could be uncoupled. The effect of these manipulations appears to be different in B. anynana and H. melpomene. In Bicyclus, "black" scales induced by wing damage at an ectopic location normally containing brown scales acquire both an intermediate structure and color between that of brown and normal black scales. In Heliconius, however, intermediate colors or scale structure were never observed, and scales with an altered color (due to damage) always have the same structure as normal scales with that color. The results are discussed on the basis of gene expression patterns, variability in rates of scale development and pigment, and scale sclerotization pathways.