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Hummingbirds are perhaps the most exquisite bird species because of their prominent iridescence, created by stacks of melanosomes in the feather barbules. The feather colours crucially depend on the nanoscopic dimensions of the melanosome, and the displayed iridescence can distinctly vary, dependent on the spatial organization of the barbs and barbules. We have taken the genus Coeligena as a model group, with species having feathers that strongly vary in their spatial reflection properties. We studied the feather morphology and the optical characteristics. We found that the coloration of Coeligena hummingbirds depends on both the Venetian-blind-like arrangement of the barbules and the V-shaped, angular arrangement of the barbules at opposite sides of the barbs. Both the nanoscopic and microscopic organization of the hummingbird feather components determine the bird's macroscopic appearance.

Among the many richly coloured birds, hummingbirds with their brilliant colouration are outstanding. We studied the plumage of male and female Blue-throated Starfrontlet,Coeligena helianthea, which exhibits a marked sexual dichromatism. The wide diversity of coloured feathers (blue, purple, golden, green, red) makes it an attractive species to investigate the structural basis of the colouration and to study the connection between the displayed colours and the perception by conspecifics. We analysed the optical properties of the feather barbules, applying spectrophotometry, scatterometry, and electron microscopy. Using the anatomical results, the spectral data can be interpreted by optical modelling. The reflectance spectra of the feathers of maleC. heliantheastrikingly overlap with the spectral sensitivities of bird photoreceptors, which suggests that the feather and photoreceptor spectra are tuned.

Morpho butterflies are universally admired for their iridescent blue coloration, which is due to nanostructured wing scales. We performed a comparative study on the coloration of 16 Morpho species, investigating the morphological, spectral and spatial scattering properties of the differently organized wing scales. In numerous previous studies, the bright blue Morpho coloration has been fully attributed to the multi-layered ridges of the cover scales' upper laminae, but we found that the lower laminae of the cover and ground scales play an important additional role, by acting as optical thin film reflectors. We conclude that Morpho coloration is a subtle combination of overlapping pigmented and/or unpigmented scales, multilayer systems, optical thin films and sometimes undulated scale surfaces. Based on the scales' architecture and their organization, five main groups can be distinguished within the genus Morpho, largely agreeing with the accepted phylogeny.

The male Anna's hummingbird features a brightly reddish-pink reflecting gorget, due to large stacks of melanosomes in the feather barbules, arranged in layers separated by keratin. Direct observations together with detailed scatterometry demonstrated that the barbules reflect incident light in an approximately specular manner. The structural colouration is iridescent, i.e. varies with a changing angle of light incidence. Spectrophotometrical measurements of the barbule reflectance and absorbance can be well interpreted with calculated spectra obtained with a transfer matrix method for optical multilayers, using anatomical data and measured refractive index spectra. The organization of the reflectors as a Venetian blind presumably functions to create a high spectral contrast of the male's plumage during courtship.

Butterflies belonging to the nymphalid subfamily, Morphinae, are famous for their brilliant blue wing coloration and iridescence. These striking optical phenomena are commonly explained as to originate from multilayer reflections by the ridges of the wing scales. Because the lower lamina of the scales of related nymphalid butterflies, the Nymphalinae, plays a dominant role in the wing coloration, by acting as a thin film reflector, we investigated single blue scales of three characteristic Morpho species: M. epistrophus, M. helenor and M. cypris. The experimental data obtained by spectrophotometry, scatterometry and scanning electron microscopy demonstrated that also in the Morpho genus the lower lamina of both the cover and ground scales acts as an optical thin film reflector, contributing importantly to the blue structural coloration of the wings. Melanin pigment has a contrast-enhancing function in a sub-class of ground scales. Electronic supplementary material The online version of this article (doi:10.1007/s00359-016-1084-1) contains supplementary material, which is available to authorized users.

Background: Ultrastructures in butterfly wing scales can take many shapes, resulting in the often striking coloration of many butterflies due to interference of light. The plethora of coloration mechanisms is dazzling, but often only single mechanisms are described for specific animals.Results: We have here investigated the male Rajah Brooke's birdwing, Trogonoptera brookiana, a large butterfly from Malaysia, which is marked by striking, colorful wing patterns. The dorsal side is decorated with large, iridescent green patterning, while the ventral side of the wings is primarily brown-black with small white, blue and green patches on the hindwings. Dense arrays of red hairs, creating a distinct collar as well as contrasting areas ventrally around the thorax, enhance the butterfly's beauty. The remarkable coloration is realized by a diverse number of intricate and complicated nanostructures in the hairs as well as the wing scales. The red collar hairs contain a broad-band absorbing pigment as well as UV-reflecting multilayers resembling the photonic structures of Morpho butterflies; the white wing patches consist of scales with prominent thin film reflectors; the blue patches have scales with ridge multilayers and these scales also have centrally concentrated melanin. The green wing areas consist of strongly curved scales, which possess a uniquely arranged photonic structure consisting of multilayers and melanin baffles that produces highly directional reflections.Conclusion: Rajah Brooke's birdwing employs a variety of structural and pigmentary coloration mechanisms to achieve its stunning optical appearance. The intriguing usage of order and disorder in related photonic structures in the butterfly wing scales may inspire novel optical materials as well as investigations into the development of these nanostructures in vivo.

SUMMARY The wings of the swordtail butterfly Graphium sarpedon nipponum contain the bile pigment sarpedobilin, which causes blue/green colored wing patches. Locally the bile pigment is combined with the strongly blue-absorbing carotenoid lutein, resulting in green wing patches and thus improving camouflage. In the dorsal forewings, the colored patches lack the usual wing scales, but instead have bristles. We have found that on the ventral side most of these patches have very transparent scales that enhance, by reflection, the wing coloration when illuminated from the dorsal side. These glass scales furthermore create a strongly polarized iridescence when illuminated by obliquely incident light from the ventral side, presumably for intraspecific signaling. A few ventral forewing patches have diffusely scattering, white scales that also enhance the blue/green wing coloration when observed from the dorsal side.

The angular and spectral reflectance of single scales of five different butterfly species was measured and related to the scale anatomy. The scales of the pierids Pieris rapae and Delias nigrina scatter white light randomly, in close agreement with Lambert's cosine law, which can be well understood from the randomly organized beads on the scale crossribs. The reflectance of the iridescent blue scales of Morpho aega is determined by multilayer structures in the scale ridges, causing diffraction in approximately a plane. The purple scales in the dorsal wing tips of the male Colotis regina act similarly as the Morpho scale in the blue, due to multilayers in the ridges, but the scattering in the red occurs as in the Pieris scale, because the scales contain beads with pigment that does not absorb in the red wavelength range. The green-yellow scales of Urania fulgens backscatter light in a narrow spatial angle, because of a multilayer structure in the scale body.