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Many studies have shown how pigments and internal nanostructures generate color in nature. External surface structures can also influence appearance, such as by causing multiple scattering of light (structural absorption) to produce a velvety, super black appearance. Here we show that feathers from five species of birds of paradise (Aves: Paradisaeidae) structurally absorb incident light to produce extremely low-reflectance, super black plumages. Directional reflectance of these feathers (0.05-0.31%) approaches that of man-made ultra-absorbent materials. SEM, nano-CT, and ray-tracing simulations show that super black feathers have titled arrays of highly modified barbules, which cause more multiple scattering, resulting in more structural absorption, than normal black feathers. Super black feathers have an extreme directional reflectance bias and appear darkest when viewed from the distal direction. We hypothesize that structurally absorbing, super black plumage evolved through sensory bias to enhance the perceived brilliance of adjacent color patches during courtship display.

Flight feathers are unique among a variety of keratinous appendages in that they are lightweight, stiff and strong. They are designed to withstand aerodynamic forces, but their morphology and structure have been oversimplified and thus understudied historically. Here we present an investigation of the shaft from seagull primary feathers, elucidate the hierarchical fibrous and porous structure along the shaft length, and correlate the tensile and nanomechanical properties to the fiber orientation. An analysis of the compressive behavior of the rachis based on a square-section model shows a good fit with experimental results, and demonstrates the synergy between the cortex and medulla. Flexural properties of the shaft along the shaft length, analyzed as a sandwich composite, reveal that although all flexural parameters decrease towards the distal shaft, the specific equivalent flexural modulus and strength increase by factors of 2 and 3, respectively. The failure mode in flexure for all specimens is buckling on the compressive surface, whereas the foamy medulla prevents destructive axial cracking and introduces important toughening mechanisms: crack deflection, fiber bridging, and microcracking., Statement of Significance: Using mechanics principles, we analyze the feather shaft as a composite beam and demonstrate that the flexural strength is extraordinary, considering its weight and tailored along the length. The cross section changes from circular in the proximal base to square/rectangular in the distal end. We also discovered that the composite design, a solid shell enclosing a foam core, produces synergistic strengthening and toughening to the feather at a minimum of weight., (Copyright © 2016 Acta Materialia Inc. Published by Elsevier Ltd. All rights reserved.)

Males of many species often use colourful and conspicuous ornaments to attract females. Among these, male manakins (family: Pipridae) provide classic examples of sexual selection favouring the evolution of bright and colourful plumage coloration. The highly iridescent feather colours of birds are most commonly produced by the periodic arrangement of melanin-containing organelles (melanosomes) within barbules. Melanin increases the saturation of iridescent colours seen from optimal viewing angles by absorbing back-scattered light; however, this may reduce the wide-angle brightness of these signals, contributing to a dark background appearance. We examined the nanostructure of four manakin species (Lepidothrix isidorei, L. iris, L. nattereri and L. coeruleocapilla) to identify how they produce their bright plumage colours. Feather barbs of all four species were characterized by dense and fibrous internal spongy matrices that likely increase scattering of light within the barb. The iridescent, yet pale or whitish colours of L. iris and L. nattereri feathers were produced not by periodically arranged melanosomes within barbules, but by periodic matrices of air and β-keratin within barbs. Lepidothrix iris crown feathers were able to produce a dazzling display of colours with small shifts in viewing geometry, likely because of a periodic nanostructure, a flattened barb morphology and disorder at a microstructural level. We hypothesize that iridescent plumage ornaments of male L. iris and L. nattereri are under selection to increase brightness or luminance across wide viewing angles, which may potentially increase their detectability by females during dynamic and fast-paced courtship displays in dim light environments., (© 2016. Published by The Company of Biologists Ltd.)

During the breeding season male, but not female, individuals of the purple sunbird possess colourful plumage of chiefly blue and black coloration with a splatter of orange and yellow on the chest. Representative feathers of these colours were collected from male birds during the breeding season and analyzed by reflectance and scanning electron microscopy. The rachis, which is the central support of a feather on which various barbs and barbules are arranged, is spongy and made up of keratin layers with rod-shaped melanosomes sparsely distributed within these layers. Barbs and barbules are the structural units of the feather and depending on how they are arranged provide a characteristic shape to the feather. X-Ray diffraction (XRD) analyses of the feathers revealed the presence of various metal elements that might contribute to the feathers' colorations. Blue feathers are iridescent and contain mainly iron, copper, zinc and cobalt (in that order of abundance); black feathers were also found to contain mainly iron and copper, but chromium instead of zinc and cobalt, while yellow feathers were found to contain predominantly cobalt and nickel. The metal content of the feathers in an as yet unknown way may be involved in the production of the distinct absorbance and reflectance patterns that the brilliant plumage of the purple sunbird is renowned for., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Eurasian Jay (Garrulus glandarius) feathers display periodic variations in the reflected colour from white through light blue, dark blue and black. We find the structures responsible for the colour are continuous in their size and spatially controlled by the degree of spinodal phase separation in the corresponding region of the feather barb. Blue structures have a well-defined broadband ultra-violet (UV) to blue wavelength distribution; the corresponding nanostructure has characteristic spinodal morphology with a lengthscale of order 150 nm. White regions have a larger 200 nm nanostructure, consistent with a spinodal process that has coarsened further, yielding broader wavelength white reflectance. Our analysis shows that nanostructure in single bird feather barbs can be varied continuously by controlling the time the keratin network is allowed to phase separate before mobility in the system is arrested. Dynamic scaling analysis of the single barb scattering data implies that the phase separation arrest mechanism is rapid and also distinct from the spinodal phase separation mechanism i.e. it is not gelation or intermolecular re-association. Any growing lengthscale using this spinodal phase separation approach must first traverse the UV and blue wavelength regions, growing the structure by coarsening, resulting in a broad distribution of domain sizes.

Avian plumage colours are model traits in understanding the evolution of sexually selected ornamental traits. Paradoxically, iridescent structural colours, probably the most dazzling of these traits, remain the most poorly understood. Though some data suggest that expression of bright iridescent plumage colours produced by highly ordered arrays of melanosomes and keratin is condition-dependent, almost nothing is known of their ontogeny and thus of any developmental mechanisms that may be susceptible to perturbation. Here, we use light and electron microscopy to compare the ontogeny of iridescent male and non-iridescent female feathers in blue-black grassquits. Feather barbules of males contain a single layer of melanosomes bounded by a thin layer of keratin-producing blue iridescent colour, while those of females contain disorganized melanosomes and no outer layer. We found that nanostructural organization of male barbules occurs late in development, following death of the barbule cell, and is thus unlikely to be under direct cellular control, contrary to previous suggestions. Rather, organization appears to be caused by entropically driven self-assembly through depletion attraction forces that pin melanosomes to the edge of barbule cells and to one another. These forces are probably stronger in developing barbules of males than of females because their melanosomes are (i) larger, (ii) more densely packed, and (iii) more homogeneously distributed owing to the more consistent shape of barbules during keratinization. These data provide the first proposed developmental pathway for iridescent plumage colours, and suggest that any condition dependence of iridescent barbules is likely driven by factors other than direct metabolic cost.

Objective: Present study was performed to record the oviposition and egg laying pattern of two phthirapteran species infesting red whiskered Bulbul., Methods: The number of egg laid by each species on different parts of body was recorded by direct observation under stereozoom binocular microscope. Feathers bearing egg were subjected to SEM to observe the pattern of egg laying., Results: The maximum percentage of egg of the ischnoceran louse, Brueelia guldum were recorded on feather of back region, followed by neck and breast. The amblyceran louse, Menacanthus eurysternus prefers to lay eggs mainly on neck, head and nape feathers., Conclusion: The ischnoceran louse, Brueelia guldum exhibits widespread oviposition sites while amblyceran louse, Menacanthus eurysternus shows restricted oviposition sites on the host body.

The breast-plate plumage of male Lawes' parotia (Parotia lawesii) produces dramatic colour changes when this bird of paradise displays on its forest-floor lek. We show that this effect is achieved not solely by the iridescence--that is an angular-dependent spectral shift of the reflected light--which is inherent in structural coloration, but is based on a unique anatomical modification of the breast-feather barbule. The barbules have a segmental structure, and in common with many other iridescent feathers, they contain stacked melanin rodlets surrounded by a keratin film. The unique property of the parotia barbules is their boomerang-like cross section. This allows each barbule to work as three coloured mirrors: a yellow-orange reflector in the plane of the feather, and two symmetrically positioned bluish reflectors at respective angles of about 30°. Movement during the parotia's courtship displays thereby achieves much larger and more abrupt colour changes than is possible with ordinary iridescent plumage. To our knowledge, this is the first example of multiple thin film or multi-layer reflectors incorporated in a single structure (engineered or biological). It nicely illustrates how subtle modification of the basic feather structure can achieve novel visual effects. The fact that the parotia's breast feathers seem to be specifically adapted to give much stronger colour changes than normal structural coloration implies that colour change is important in their courtship display.

The results of comparative electron microscope study of the fine structure of the definitive contour feathers often Crows species (Perisoreus infaustus, Garrulus glandarius, Cyanopica cyanus, Pica Pica, Podoceus panderi, Nucifraga caryocatactes, Corvus monedula, C. frugilegus, C. cornix, C. corax) are presented. The results of the research allowed us to conclude that crows, along with the traditional elements of feather architectonics, have a number of species-specific microstructure characteristics that are taxonomically important.

Understanding structural colours in nature requires the right set of optical experiments: this is illustrated by a new study on iridescent bird of paradise feathers, which suggests the potential behavioural importance of dynamic colour changes., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

Honest advertisement models posit that only individuals in good health can produce and/or maintain ornamental traits. Even though disease has profound effects on condition, few studies have experimentally tested its effects on trait expression and even fewer have identified a mechanistic basis for these effects. Recent evidence suggests that black and white, but not grey, plumage colors of black-capped chickadees (Poecile atricapillus) are sexually selected. We therefore hypothesized that birds afflicted with avian keratin disorder, a condition that affects the beak and other keratinized tissues, would show reduced expression of black and white, but not grey, color. UV-vis spectrometry of black-capped chickadees affected and unaffected by avian keratin disorder revealed spectral differences between them consistent with this hypothesis. To elucidate the mechanistic bases of these differences, we used scanning electron microscopy (SEM), electron-dispersive x-ray spectroscopy (EDX) and a feather cleaning experiment. SEM showed extreme feather soiling in affected birds, and EDX revealed that this was most likely from external sources. Experimentally cleaning the feathers increased color expression of ornamental feathers of affected, but not unaffected, birds. These data provide strong evidence that black and white color is an honest indicator in chickadees, and that variation in feather dirtiness, likely due to differences in preening behavior is a mechanism for this association.

Male Club-winged Manakins, Machaeropterus deliciosus (Aves: Pipridae), produce a sustained tonal sound with specialized wing feathers. The fundamental frequency of the sound produced in nature is approximately 1500 Hz and is hypothesized to result from excitation of resonance in the feathers' hypertrophied shafts. We used laser Doppler vibrometry to determine the resonant properties of male Club-winged Manakin's wing feathers, as well as those of two unspecialized manakin species. The modified wing feathers exhibit a response peak near 1500 Hz, and unusually high Q-values (a measure of resonant tuning) for biological objects (Q up to 27). The unmodified wing feathers of the Club-winged Manakin do not exhibit strong resonant properties when measured in isolation. However, when measured still attached to the modified feathers (nine feathers held adjacent by an intact ligament), they resonate together as a unit near 1500 Hz, and the wing produces a second harmonic of similar or greater amplitude than the fundamental. The feathers of the control species also exhibit resonant peaks around 1500 Hz, but these are significantly weaker, the wing does not resonate as a unit and no harmonics are produced. These results lend critical support to the resonant stridulation hypothesis of sound production in M. deliciosus.

Non-iridescent structural plumage color is typically produced by coherent scattering of light within a matrix of keratin and air (a ;spongy layer') in feather barbs. It remains unclear what role, if any, the basal melanin layer underlying this spongy layer plays in the production of coloration. Amelanism in birds with structural color is a ;natural experiment' in which melanin pigmentation is lost, allowing us to identify the effects of the loss of melanin on structural color production. Here we use full-spectrum spectrometry, transmission electron microscopy and Fourier analysis to compare the color and nanostructure of an amelanotic Steller's jay (Cyanocitta stelleri Gmelin) feather with a normal blue Steller's jay feather. As a control, we also examined a white domestic chicken (Gallus gallus Linnaeus) feather. The pale amelanotic jay feather had a broad reflectance curve with a blue/green peak, while the typical blue feather had a typical distinct bell-shaped blue curve with a UV/violet peak. The white chicken feather had a typical white reflectance curve with no discrete peaks. Electron microscopy revealed that both the amelanotic and blue feather barbs contained well-formed spongy layers that were of the correct size and arrangement to produce their measured peak reflectance values, whereas the chicken feather had no spongy layer. The washed-out color of the amelanotic jay feather was thus most probably caused by the loss of the basal melanin layer, suggesting that melanin functions to absorb incoherently scattered white light from the feather barb thereby increasing the purity of the color produced by the spongy layer.

Iridescence is produced by coherent scattering of light waves from alternating layers of materials of different refractive indices. In birds, iridescent colours are produced by feather barbules when light is scattered from alternating layers of keratin, melanin and air. The structure and organization of these layers, and hence the appearance of bird species with different types of plumage iridescence, varies extensively. One principal distinction between different types of iridescent colours is whether they are produced by a single pair of layers or by multiple pairs of layers. Multi-layer iridescence, such as that displayed by hummingbirds, has been relatively well characterized, but single-layer iridescence has only recently been modeled successfully. Here we use electron microscopy, spectrometry and thin-film optical modeling to investigate the glossy, ultraviolet-blue iridescent plumage colouration of adult male satin bowerbirds Ptilonorhynchus violaceus minor. The flattened barbules of adult males are composed of a superficial keratin layer overlying a melanin layer that is several granules thick. A thin-film model based on the thickness of the keratin layer and its two associated interfaces (air/keratin and keratin/melanin) generates predicted reflectance spectra that closely match measured spectra. In addition, hues predicted from this model are positively correlated with measured hues. As predicted from our thin-film model, measured hues shifted to shorter wavelengths at increasing angles of incidence and reflectance. Moreover, we found that individual variation in barbule nanostructure can predict measured variation in both hue and UV-chroma. Thus, we have characterized the microstructure of satin bowerbird barbules, uncovered the mechanisms responsible for producing ultraviolet iridescence in these barbules, and provided the first evidence of a nanostructural basis for individual variation in iridescent plumage colour.