ABSTRACT
Higher vertebrates synthesize two forms of melanin: eumelanin and pheomelanin. While the adaptive functions of eumelanin are diverse, those of pheomelanin, which is phototoxic and whose production consumes a key intracellular antioxidant (glutathione), are not clear apart from being involved in color patterns that confer concealment. The factors that have favored the evolution of pheomelanin thus remain a mystery, causing this pigment even to have been considered an "accident of nature." A recent hypothesis posits that pheomelanin has evolved because it represents an alternative mechanism to remove excess dietary cysteine, which can be toxic because of its oxidation. We tested for links between pheomelanin-based color and survival in both an intraspecific study of barn swallows Hirundo rustica and an interspecific study of 58 species of birds from North and Central America. As predicted on the basis that birds degrade excess dietary amino acids by transferring their amino group to uric acid synthesis, we found that under equal levels of uric acid in plasma, individuals or species with a higher intensity or greater proportion of plumage colored by pheomelanin (brown and chestnut coloration) had higher relative annual survival rates while controlling for the potentially confounding effects of age, sex, body size, and phylogenetic descent. Likewise, barn swallows with more intense pheomelanin-based coloration had higher prospects to survive the winter after controlling statistically for age, sex, body size, and level of uric acid. This supports the idea that pheomelanic traits evolve because of the removal of excess cysteine in nonstressful conditions, thus avoiding its toxic effects.