Cassowary gloss and a novel form of structural color in birds

One of the two lineages of extant birds resulting from its deepest split, Palaeognathae, has been reported not to exhibit structural coloration in feathers, affecting inferences of ancestral coloration mechanisms in extant birds. Structural coloration in facial skin and eggshells has been shown in this lineage, but has not been reported in feathers. We present the first evidence for two distinct mechanisms of structural color in palaeognath feathers. One extinct volant clade, Lithornithidae, shows evidence of elongate melanin-containing organelles uniquely associated with glossy/iridescent color, a structural color mechanism found in fossil outgroups and neognath birds. We also demonstrate a structural basis for the exceptional gloss in extant cassowary feathers. We propose gloss as an intermediate phenotype between matte and iridescent plumage, conferred by a thick and smooth feather rachis. Rachis-based structural color has not been previously investigated. The new data illuminate the relationships between avian melanin-based coloration and feather structure.