Structural colour: elusive iridescence strategies brought to light

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.     

The ‘chaotic’ flight feather moult of the Steppe Buzzard Buteo buteo vulpinus

Steppe Buzzards breed in Eurasia and spend the non-breeding season in Africa. Adults moult some flight feathers during the breeding season and some during the non-breeding season. Moult is arrested during migration. The extent of moult of flight feathers in adults is highly variable between individuals in southern Africa, with the renewal of two primaries, three secondaries and five rectrices as the most frequently encountered pattern. Time spent on the non-breeding grounds in South Africa is too short to allow for a sequential moult. Moult of flight feathers is restricted to the almost synchronous dropping of a number of feathers upon arrival, with few being replaced subsequently. Any of the flight feathers can be replaced in southern Africa, and the pattern of renewal in primaries and secondaries cannot be distinguished from random. Tail feathers are replaced in an alternating (transilient) pattern. Moult in the non-breeding areas may primarily be complementary to moult on the breeding grounds, but these two partial moults per year are insufficient to renew all flight feathers annually. Middle secondaries and central tail feathers are regularly carried over to a third moult, but this is rare for primaries.     

Feather isotope analysis discriminates age-classes of Western, Least, and Semipalmated sandpipers when plumage methods are unreliable

ABSTRACT Avian age‐class discrimination is typically based on the completeness of the first prebasic molt. In several calidrid sandpiper species, juvenal flight feathers grown on Arctic breeding grounds are retained through the first three migrations. Thereafter, flight feathers are grown annually at temperate migratory stopover sites during the fall or on the subtropical wintering grounds. Standard methods for distinguishing age classes of sandpipers rely on a combination of traits, including body plumage, coloration of protected inner median covert edges, and extent of flight feather wear. We tested the ability of stable hydrogen isotope ratios in flight feathers (δD_f) to distinguish young birds in their first winter through second fall from older adults in three calidrid sandpiper species, Western (Calidris mauri), Least (C. minutilla), and Semipalmated (C. pusilla) sandpipers. We compared the apparent reliability of the isotope approach to that of plumage‐based aging. The large expected differences in δD_f values of flight feathers grown at Arctic versus non‐Arctic latitudes enabled use of this technique to discriminate between age‐classes. We determined δD_f values of known Arctic‐grown feathers from juveniles that grew their flight feathers on the breeding grounds. Flight feather δD_f values of southward‐migrating adults showed bimodal distributions for all three species. Negative values overlapped with species‐specific juvenile values, identifying putative second fall birds with high‐latitude grown juvenal feathers retained from the previous year. The more positive values identified older adults who grew their feathers at mid‐ and low latitudes. Importantly, δD_f analysis successfully identified first‐winter and second‐fall birds not detected by plumage‐based aging. Flight feather wear alone was a poor basis for age classification because scores overlapped extensively between putative second fall birds and older adults. Flight feather hydrogen isotope analysis enables more definitive assignment of age classes when standard plumage methods are unreliable.     

Moult speed affects structural feather ornaments in the blue tit

Growing evidence suggests that structural feather colours honestly reflect individual quality or body condition but, contrary to pigment‐based colours, it is not clear what mechanism links condition to reflectance in structural feather colours. We experimentally accelerated the moult speed of a group of blue tits (Cyanistes caeruleus) by exposing them to a rapidly decreasing photoperiod and compared the spectral characteristics of their structural feather colours with those of control birds. Blue tits were sexually dimorphic on the UV/blue crown and on the white cheek feathers. Moult speed, however, dramatically reduced brightness and the saturation only on the UV/blue crown feathers, whereas structural white on the cheek feathers was basically unaffected by moult speed. Given that the time available for moulting is usually confined to the period between the end of the breeding season and migration or wintering, UV/blue colours, but not structural white, may convey long‐term information about an individual’s performance during the previous breeding season. The trade‐off between fast moulting and structural colour expression may represent a previously unrecognized selective advantage for early‐breeding birds.     

Functional Microanatomy of the Feather-Bearing Integument: Implications for the Evolution of Birds and Avian Flight

A selective regime favoring a streamlining of body contoursand surfaces is proposed as having been instrumental in drivingthe morphological and functional transformations of an unfeatheredreptilian integument into a feather-bearing avian one. Thishypothesis is consistent with a new, structurally and functionallycoherent analysis of the microanatomy of the avian feather-bearingintegument as a complex, integrated organ system that includesan intricate, hydraulic skeleto-muscular apparatus of the feathers,a dermo-subcutaneous muscle system of the integument, and asubcutaneous hydraulic skeletal system formed by fat bodies.Key elements of the evidence supporting the new hypothesis are(1) the presence of depressor feather muscles that are not neededas antagonists for the erector feather muscles, but can counteractexternal forces, such as air currents; (2) the fact that thehighly intricate feather-bearing integument represents a machineryto move feathers or to stabilize them against external forces;(3) the crucial role of the coat of feathers in streamliningthe body contours and surfaces of birds; (4) the aerodynamicrole of feathers as pressure and turbulence sensors and as controllabletemporary turbulators; and (5) the critical role that a streamlinedbody plays in avian flight and is likely to have played in theevolutionary transformations from ecologically and locomotorilyversatile quadrupedal reptiles to volant bipedal birds withoutpassing through parachuting or gliding stages. These transformationsare likely to have occurred more than once. The ancestral birdswere probably small, arboreal, hopping, and using flap-bounding,or intermittent bounding, flight.     

Different scales of spatial segregation of two species of feather mites on the wings of a passerine bird

The "condition-specific competition hypothesis" proposes that coexistence of 2 species is possible when spatial or temporal variations in environmental conditions exist and each species responds differently to those conditions. The distribution of different species of feather mites on their hosts is known to be affected by intrinsic host factors such as structure of feathers and friction among feathers during flight, but there is also evidence that external factors such as humidity and temperature can affect mite distribution. Some feather mites have the capacity to move through the plumage rather rapidly, and within-host variation in intensity of sunlight could be one of the cues involved in these active displacements. We analyzed both the within- and between-feather spatial distribution of 2 mite species, Trouessartia bifurcata and Dolichodectes edwardsi , that coexist in flight feathers of the moustached warbler Acrocephalus melanopogon. A complex spatial segregation between the 2 species was observed at 3 spatial levels, i.e., "feather surfaces," "between feathers," and "within feathers." Despite certain overlapping distribution among feathers, T. bifurcata dominated proximal and medial regions on dorsal faces, while D. edwardsi preferred disto-ventral feather areas. An experiment to check the behavioral response of T. bifurcata to sunlight showed that mites responded to light exposure by approaching the feather bases and even leaving its dorsal face. Spatial heterogeneity across the 3 analyzed levels, together with response to light and other particular species adaptations, may have played a role in the coexistence and segregation of feather mites competing for space and food in passerine birds.     

On the Evolution of Feathers from an Aerodynamic and Constructional View Point

The evolution of birds and feathers are examined in terms ofthe aerodynamic constraints imposed by the arboreal and cursorialmodels of flight evolution. The cursorial origin of flight isassociated with the putative coelurosaurian ancestry of birds.As presently known, coelurosaurs have a center of mass locatedin the pelvic region and an elongated pubis that is ventrallyor anteriorly directed. Both of these characteristics make itdifficult to postulate an origin of flight that would involvea gliding phase because the abdomen cannot be flattened intoan aerodynamic shape. Moreover, the cursorial model must counteractgravity using the hindlimb and, thus, selection for the powerrequirement for lift-off would not focus on the forelimb. Therefore,if the hypothesis proposing a coelurosaurian ancestry of birdsis to remain viable, it must be via an as yet undiscovered taxonthat is compatible with the morphological and aerodynamic constraintsimposed by flight evolution. The arboreal model, currently centers around non-dinosauriantaxa and is more parsimonious in that early archosaurs haveshort pubes that do not preclude an aerodynamic body profile.Moreover, the arboreal proavis uses gravity to create the airflowover the body surfaces and is, thus, energy efficient. Considerationof the initial aerodynamic roles of feathers and feather designare consistent with a precursory gliding phase. Whether avianancestry lies among coelurosaur theropods or earlier archosaurs,we must remain mindful of the complex aerodynamic dictates ofgliding and powered flight and avoid formalistic approachesthat co-opt sister taxa, with their known body form, as functionalancestors.     

Brood size manipulations reveal relationships among physiological performance,body condition and plumage colour in Northern Flicker Colaptes auratus nestlings

Visual signals of quality in offspring, such as plumage colour, should honestly advertise need and/or body condition, but links between nutritional status, physiological performance and the expression of colours are complex and poorly understood. We assess how food stress during rearing affected two physiological measures (T‐cell‐mediated immune function and corticosterone level in feathers: CORT_f) and how these two variables were related to carotenoid and melanin coloration in Northern Flicker Colaptes auratus nestlings. We were also interested in how these two physiological measures were influenced by the sex of the nestling. We experimentally manipulated brood size to alter levels of food availability to nestlings during development. We measured carotenoid‐based colour (chroma and brightness) in wing feathers and the size of melanin spots on breast feathers. In agreement with our prediction, nestlings in the reduced brood treatment had better body condition and stronger immune responses than those in the control and brood enlargement treatments. This supports the hypothesis that immune responses are energetically costly. In contrast, CORT_f was not related to nestling body condition or sex and was unaffected by brood size manipulation. Nestlings of both sexes with stronger T‐cell‐mediated immune responses had larger melanin spots but only males with higher immune responses also had brighter flight feathers. Feather brightness decreased with increasing CORT_f levels. Our study is one of the few to examine the relationship between multiple physiological and plumage measures in nestlings and shows that plumage colour and immune function signalled body condition of nestlings, but that feather corticosterone levels did not.     

THE DISTRIBUTION OF STRUCTURAL COLOURS AND UNUSUAL PIGMENTS IN THE CLASS AVES.

Structural colours in birds are surveyed systematically. From this survey, the common structural configurations in feathers, producing bright colours of the iridescent and non-iridescent types respectively, are seen to exclude one another
Feather pigments of unusual shades or chemical constitutions have been included in the survey, since some of them produce colour effects similar to structural colours of the non-iridescent type     

Feather holes and flight performance in the barn swallow Hirundo rustica

Feather holes are small (0.5–1?mm in diameter) deformities that appear on the vanes of flight feathers. Such deformities were found in many bird species, including galliforms and passerines. Holey flight feathers may be more permeable to air, which could have a negative effect on their ability to generate aerodynamic forces. However, to date the effects of feather holes on flight performance in birds remained unclear. In this study we investigated the relationship between the number of feather holes occurring in the wing or tail feathers and short term flight performance traits – aerial manoeuvrability, maximum velocity and maximum acceleration – in barns swallows, which are long distance migrating aerial foragers. We measured short-term flight performance of barn swallows in a standardized manner in flight tunnels. We found that acceleration and velocity were significantly negatively associated with the number of holes in the wing flight feathers, but not with those in the tail feathers. In the case of acceleration the negative relationship was sex specific – while acceleration significantly decreased with the number of feather holes in females, there was no such significant association in males. Manoeuvrability was not significantly associated with the number of feather holes. These results are consistent with the hypothesis that feather holes are costly in terms of impaired flight. We discuss alternative scenarios that could explain the observed relationships. We also suggest directions for future studies that could investigate the exact mechanism behind the negative association between the number of feather holes and flight characteristics.     

Colour composition of nest lining feathers affects hatching success of barn swallows,Hirundo rustica (Passeriformes: Hirundinidae)

Many bird species use feathers as lining material, and its functionality has traditionally been linked to nest insulation. However, nest lining feathers may also influence nest detection by predators, differentially affect reproductive investment of mates in a post‐mating sexual selection process, and affect the bacterial community of the nest environment. Most of these functions of nest lining feathers could affect hatching success, but the effect might vary depending on feather coloration (i.e. pigmented versus white feathers). This would be the case if coloration is related to: (1) thermoregulatory properties; (2) attractiveness of feathers in the nest for mates; (3) eggshell bacterial density. All of these hypothetical scenarios predict that feathers of different colours would differentially affect the hatching success of birds, and that birds should preferentially choose the most beneficial feather colour for lining their nests. Results from two different experiments performed with a population of Danish barn swallow, Hirundo rustica, were in accordance with these predictions. First, H. rustica preferentially selected white experimentally offered feathers for lining their nests. Second, the experimental manipulation of the feather colour composition of nests of H. rustica had a significant effect on hatching success. Experimental nests with more white feathers added at the beginning of incubation had a lower probability of hatching failures, suggesting differential beneficial effects of lining nests with feathers of this colour. We discuss the relative importance of hypothetical functional scenarios that predicted the detected associations, including those related to sexual selection or to the community of microorganisms associated with feathers of different colours. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 67–74.     

Phylogenetic Context for the Origin of Feathers

A number of hypotheses have been suggested for the origin ofbirds and feathers. Although distributions of functional complexeshave frequently been used to test phylogenetic hypotheses, analysisof the origin of feathers remains hampered by the incompletefossil record of these unmineralized structures. It is alsocomplicated by approaches that confuse the origins of birds,feathers, and flight without first demonstrating that theserelate to the same historical event. Functional speculationregarding the origin of feathers usually focuses on three possiblealternatives: (1) flight; (2) thermal insulation; or (3) display.Recent fossil finds of Late Cretaceous feathered dinosaurs inChina have demonstrated that feathers appear to have originatedin taxa that retained a significant number of primitive nonavianfeatures. Current evidence strongly suggests that birds aretheropod dinosaurs, and that the most primitive known feathersare found on non-flying animals. This further suggests thatfeathers did not evolve as flight structures. Thermoregulatory,display, and biomechanical support functions remain possibleexplanations for the origin of feathers. As the earliest functionof feathers was probably not for aerial locomotion, it may bespeculated that the transitional animals represented by theChinese fossils possessed skin with the tensile properties ofreptiles and combined it with the apomorphic characteristicsof feathers.     

Increase of feather quality during moult: a possible implication of feather deformities in the evolution of partial moult in the great tit Parus major

Here we investigate the change in feather quality during partial post‐juvenile and complete post‐breeding moult in great tit Parus major by measuring the change in the number of fault bars and feather holes on wing and tail feathers. Feathers grown during ontogeny usually are of lower quality than feathers grown following subsequent moults at independence. This is reflected by higher number of fault bars and feather holes on juveniles compared to adults. Fault bars are significantly more common on tail and proximal wing feathers than on the distal remiges, indicating a mechanism of adaptive allocation of stress induced abnormalities during ontogeny into the aerodynamically less important flight feathers. On the contrary, feather holes produced probably by chewing lice have a more uniform distribution on wing and tail feathers, which may reflect the inability of birds to control their distribution, or the weak natural selection imposed by them. The adaptive value of the differential allocation of fault bar between groups of feathers seems to be supported by the significantly higher recapture probability of those juvenile great tits which have fewer fault bars at fledging on the aerodynamically most important primaries, but not on other groups of flight feathers. The selection imposed by feather holes seems to be smaller, since except for the positive association between hatching date, brood size and the number of feather holes at fledging, great tits' survival was not affected by the number of feather holes. During post‐juvenile moult, the intensity of fault bars drops significantly through the replacement of tail feathers and tertials, resulting in disproportional reduction of the total number of fault bars on flight feathers related to the number of feathers replaced. The reduction in the number of fault bars during post‐juvenile moult associated with their adaptive allocation to proximal wing feathers and rectrices may explain the evolution of partial post‐juvenile moult in the great tit, since the quality of flight feathers can be increased significantly at a relatively small cost. Our results may explain the widespread phenomenon of partial post‐juvenile moult of flight feathers among Palearctic passerines. During the next complete post‐breeding moult, the total number of fault bars on flight feathers has remained unchanged, indicating the effectiveness of partial post‐juvenile moult in reducing the number of adaptively allocated fault bars. The number of feather holes has also decreased on groups of feathers replaced during partial post‐juvenile moult, but the reduction is proportional with the number of feathers moulted. In line with this observation, the number of feather holes is further reduced during post‐breeding moult on primaries and secondaries, resulting in an increase in feather quality of adult great tits.     

Morphological basis of glossy red plumage colours

Brightly coloured feathers, including the brilliant reds produced by carotenoids, are sometimes shiny in appearance. Gloss is a common property of materials and usually arises through specular reflection from smooth, flat surfaces. However, the production of gloss on red feathers has never been examined. In the present study, we compared the optical and structural properties of glossy and matte carotenoid‐based red feathers of multiple species to identify the proximate basis for their glossiness. Although specular reflectance did not differ between glossy and matte feathers, diffuse reflectance was lower in glossy than in matte feathers, leading to a higher contrast gloss. Compared to matte feathers, glossy red feathers had thicker barbs with a flatter and more homogeneous morphology, consistent with expectations, as well as thicker outer keratin cortices. Moreover, glossiness was predicted by a principal component regression using these same morphological traits. We demonstrate that the gloss of carotenoid‐based red feathers is produced at least in part by a smooth, flattened barb microstructure and an enhanced nanostructure, illustrating a novel colour‐producing interaction that neither pigment, nor microstructure could alone attain. How the ecology and evolution of species with glossy red feather differ from those with typical matte red feathers represent rich areas for future study.     

Colourful parrot feathers resist bacterial degradation

The brilliant red, orange and yellow colours of parrot feathers are the product of psittacofulvins, which are synthetic pigments known only from parrots. Recent evidence suggests that some pigments in bird feathers function not just as colour generators, but also preserve plumage integrity by increasing the resistance of feather keratin to bacterial degradation. We exposed a variety of colourful parrot feathers to feather-degrading Bacillus licheniformis and found that feathers with red psittacofulvins degraded at about the same rate as those with melanin and more slowly than white feathers, which lack pigments. Blue feathers, in which colour is based on the microstructural arrangement of keratin, air and melanin granules, and green feathers, which combine structural blue with yellow psittacofulvins, degraded at a rate similar to that of red and black feathers. These differences in resistance to bacterial degradation of differently coloured feathers suggest that colour patterns within the Psittaciformes may have evolved to resist bacterial degradation, in addition to their role in communication and camouflage.     

Is the structural and psittacofulvin‐based coloration of wild burrowing parrots Cyanoliseus patagonus condition dependent?

The bright colours of parrots are caused by psittacofulvin pigments, which appear unique to this Order, and by structural colours. We measured red (psittacofulvin), green (mixed) and blue (structural) colours of wild burrowing parrots Cyanoliseus patagonus of northeastern Patagonia, Argentina, and measured nestlings regularly to obtain data on breeding success and nestling growth. As adult feathers are moulted outside the breeding season, adult body condition could not be measured directly during feather growth, and climatic conditions were used as an indirect parameter. The colony of burrowing parrots is surrounded by Monte steppe habitat, the breeding success has been shown to depend strongly on the climatic patterns. The area experienced a drought with very poor breeding success as well as a year of above‐average precipitation during the study period, serving as a natural experiment. We thus analysed the variability of colouration within the population among and within breeding seasons. We observed strong inter‐annual differences in nestling and adult colouration. Nestlings grew blue feathers with lower achromatic brightness during better conditions, and when controlling for year effects, nestlings with higher mass and from more successful families also had blue feathers with lower achromatic brightness. Adult blue feathers showed the same trend, with lower achromatic brightness in the moult following breeding seasons of better conditions. In contrast, during better conditions, adults grew red feathers with higher achromatic brightness and the colour hue was also affected, and the hue of the red region of nestlings varied with the hatching order. The colour of all three regions of nestlings varied between nests, and the colour of the red region of adult males positively correlated with breeding success (clutch size, brood size). In summary, the present data suggest that environmental conditions contribute to variability in both structural and the psittacofulvin‐based colours of wild burrowing parrots.     

Iridescent ultraviolet signal in the orange sulphur butterfly (Colias eurytheme): spatial, temporal and spectral properties

Many of nature's most striking animal colours are iridescent, exhibiting a high degree of spectral purity and strong angular dependence of intensity and hue. Although a growing number of studies have detailed the intricate mechanisms responsible for producing iridescent colours, few attempts have been made to describe their dynamic appearance in ecologically and behaviourally realistic contexts. We suggest that the optical properties unique to iridescent structural colours are important for understanding how they function as signals during behavioural interactions. Using males of the orange sulphur butterfly, Colias eurytheme , which exhibit an iridescent ultraviolet (UV) reflectance on their dorsal wing surfaces, we develop a holistic framework for inferring the appearance of this signal to conspecifics under field conditions that incorporate data on their spectral sensitivity. We show that, during flight, the UV signal is brightest within a wing beat cycle when viewed from directly above the male. Spectral properties of the signal under natural lighting indicate that male wing colour should be readily perceived and distinguished from that of females and from the dark green visual background of UV-absorbing vegetation. Finally, our analyses permit predictions regarding how signal senders and receivers should orientate themselves for maximal transmission and reception of this ultraviolet iridescent signal.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 349–364.     

Carotenoids need structural colours to shine

The bright colours of feathers are among the most striking displays in nature and are frequently used as sexual signals. Feathers can be coloured by pigments or by ordered tissue, and these mechanisms have traditionally been treated as distinct modes of display. Here we show that some yellow plumage colour is created both by reflection of light from white structural tissue and absorption of light by carotenoids. Thus, structural components of feathers contribute substantially to yellow 'carotenoid' displays, but the effect of variation in structural components on variation in colour displays is, to our knowledge, unstudied. The presence of structural colour in some carotenoid-based colour displays will have to be considered in studies of colour signalling.     

Bionik der Flugfeder

Bionik of flight feathers Large soaring landbirds depend on cross‐country flights as cheaply as possible. The migration of eastern European White Storks to southerly wintering areas with a length of more than 8000 km as well as the foraging flights of vultures over huge areas are physiologically only possible thanks to optimized soaring flight capabilities of the birds. The mechanisms to increase uplift and to reduce unwanted friction and vortex drags as presented here show that these adaptations are realized down to the microstructural level of the flight feathers. These are in particular: the airstream permeable structures along the feather shaft, the flow adaptive curvature of the feather profile and the shaping of the feather tip. These mechanisms serve a significant reduction of drag which was prerequisite to a successful development of the flight feather cascade. In conferred form they can be found also in technical missiles.     

Snipe in Eastern Africa

Summary— The Jack Snipe is a very distinctive bird and should not be confused with the other four species. Its flight also is slower and more butterfly-like.
The Pintail Snipe is not readily distinguishable in the field from the Common Snipe, but in the hand the extraordinarily thin outer tail feathers are unmistakable.
The Common Snipe is easily distinguishable by the outer tail feathers, which can be seen at close quarters when the bird is in flight. It also has a more rapid flight than either the Double or African Snipe, and looks rather smaller on the wing.
The African Snipe has a heavier and rather slower flight than the Common Snipe and usually looks larger on the wing. The much narrower white outer tail feathers are the most outstanding character.
The Double Snipe is the one that is usually confused with the African Snipe, as its general appearance and*outer tail feathers are much the same when seen in flight. In the hand the barred belly is a most distinctive character. The outer tail feathers are broader than those of the African Snipe. It should not be confused with the Common Snipe, and certainly not with the Pintail Snipe.
When a bag of mixed Double and African Snipe are laid out, the white unbarred belly of the latter is very apparent.