Male Lawes's Parotia, a bird of paradise, use the highly directional reflection of the structurally colored, brilliant-silvery occipital feathers in their courtship display. As in other birds, the structural coloration is produced by ordered melanin pigmentation. The barbules of the Parotia's occipital feathers, with thickness ,3 mm, contain 6-7 layers of densely packed melanin rodlets (diameter ,0.25 mm, length ,2 mm). The effectively ,0.2 mm thick melanin layers separated by ,0.2 mm thick keratin layers create a multilayer interference reflector. Reflectance measurements yielded peak wavelengths in the near-infrared at ,1.3 mm, i.e., far outside the visible wavelength range. With the Jamin-Lebedeff interference microscopy method recently developed for pigmented media, we here determined the refractive index of the intact barbules. We thus derived the wavelength dependence of the refractive index of the barbules' melanin to be 1.7-1.8 in the visible wavelength range. Implementing the anatomical and refractive index data in an optical multilayer model, we calculated the barbules' reflectance, transmittance and absorptance spectra, thereby confirming measured spectra. Keywords: bird of paradise; interference reflector; iridescence; Jamin-Lebedeff microscopy; multilayer
INTRODUCTIONIn animal integuments melanins commonly produce dull red, brown and black colors. With nanoscale order, melanin in a matrix of vertebrate keratin or arthropod chitin can produce striking structural colors. [1][2][3][4] The magnificent displays of birds of paradise exemplify how the fine branches of the bird feathers, the barbules, are modified to achieve a diverse range of visual effects through structural coloration. In Lawes's Parotia (Parotia lawesii), the barbules of the males' breast feathers have a boomerang-shaped cross-section, which produces three directional-colored reflectors. 5 Here we investigate the male Parotia's occipital (or nape) feathers, which produce a shiny, silvery patch (Figure 1a and 1b). Compared to the breast feathers they are less colorful, but the barbules of the occipital feathers exhibit a mirror-like, directional reflection due to nanostructured melanin. 6 The uniquely colorful breast feathers allows the breast color to switch sharply between yellow, blue and black as the bird moves, during the ballerina dance, which is performed as part of the courtship display. [7][8][9] The shiny occipital feathers have a similar function. Recent behavioral observations on the closely related Wahnes's Parotia demonstrate that the occipital feather reflections are sharply directed to the observing females during part of the courtship performance, presumably to impress a potential mate, viewing from an elevated position on a tree branch. [6][7][8][9][10] To unravel the optical basis of the shiny occipital reflectors, we investigated the barbule anatomy. This revealed very regularly arranged melanosomes, i.e., small melanin rodlets, arranged in layers. To achieve an in-depth, quantitative understanding of the feathers'