Exposure to sulfur in soil explains pigmentation by pheomelanin in birds inhabiting Iceland

Melanins are the most common pigments in birds and mammals. The synthesis of the sulfurated form of melanin, termed pheomelanin, is promoted by high levels of thiols in melanocytes. Exposure to elemental sulfur in the environment can potentially influence intracellular thiol levels in terrestrial animals, but this has never been investigated in the wild. This is particularly relevant in areas with a high volcanic activity that facilitate sulfur exposure. We explored for an association between the pheomelanin content of feathers in 35 breeding birds from 13 phylogenetically diverse species and 29 locations in Iceland, and the sulfur content of nearby soil samples. After controlling for the effect of phylogenetic relatedness between species, we found that the pheomelanin content of feathers was significantly correlated with the sulfur content of soil, indicating that pheomelanin production increases with sulfur exposure. Our results suggest that environmental exposure to relatively high levels of elemental sulfur promotes the expression of pheomelanin-based pigmentation phenotypes in birds, likely by an influence on intracellular thiols during development. This is the first evidence that soil chemistry can affect the phenotype of animals. Highly volcanic areas such as Iceland are thus potential sources of global phenotypic diversity by promoting sulfur exposure in animals.