Cosmetic coloration in birds: occurrence, function, and evolution.

Colorful plumages are conspicuous social signals in birds, and the expression of these colors often reflects the quality of their bearers. Since mature feathers are dead structures, plumage color is often considered a static signal that does not change after molt. Feathers, however, can and do deteriorate between molts, and birds need to invest heavily in plumage maintenance. Here we argue that this need for preserving plumage condition and hence signaling content might have given rise to a novel type of sexual signal: cosmetic coloration. Cosmetic coloration occurs when the substances used for plumage maintenance change the color of the feathers, thereby becoming a signal themselves. Our review of cosmetic coloration in birds demonstrates that it is more widespread than currently realized, occurring in at least 13 bird families. Cosmetics have varied origins: they can be produced by the bird itself (uropygial and skin secretions, feather powder) or obtained from the environment (soil, iron oxide). Intraspecific patterns of cosmetic use (sex, age, and seasonal dimorphism) suggest that in many cases it may act as a sexual signal. However, more information is required on function, mechanisms, and costs to understand the evolution of cosmetic coloration and to confirm its signaling role.