Convergent evolution of super black plumage near bright color in 15 bird families

Abstract

ABSTRACT We examined extremely low-reflectance, velvety black plumage patches in 32 bird species from 15 families and five orders and compared them with 22 closely related control species with normal black plumage. We used scanning electron microscopy to investigate microscopic feather anatomy, and applied spectrophotometry and hyperspectral imaging to measure plumage reflectance. Super black plumages are significantly darker and have more broadband low reflectance than normal black plumages, and they have evolved convergently in 15 avian families. Super black feather barbules quantitatively differ in microstructure from normal black feathers. Microstructural variation is significantly correlated with reflectance: tightly packed, strap-shaped barbules have lower reflectance. We assigned these super black feathers to five heuristic classes of microstructure, each of which has evolved multiple times independently. All classes have minimal exposed horizontal surface area and 3D micrometer-scale cavities greater in width and depth than wavelengths of light. In many species, barbule morphology varied between the super black exposed tip of a feather and its (i) concealed base or (ii) iridescently colored spot. We propose that super black plumages reduce reflectance, and flatten reflectance spectra, through multiple light scattering between the vertically oriented surfaces of microscale cavities, contributing to near-complete absorption of light by melanin. All super black plumage patches identified occur adjacent to brilliant colored patches. Super black plumage lacks all white specular reflections (reference points used to calibrate color perception), thus exaggerating the perceived brightness of nearby colors. We hypothesize that this sensory bias is an unavoidable by-product of color correction in variable light environments.