Inter‐annual and body topographic consistency in the plumage bacterial load of Great Tits

ABSTRACT Plumage bacteria may play an important role in shaping the life histories of birds. However, to design suitable experiments to examine causal relationships between plumage bacteria and the fitness of host birds, natural variation in plumage bacterial communities needs to be better understood. We examined within‐individual consistency of plumage bacterial contamination in Great Tits (Parus major), comparing different body regions (ventral vs. dorsal) and comparing contamination between years. Numbers of free‐living and attached bacteria and the species richness of feather‐degrading bacterial assemblages were studied using flow cytometry and ribosomal intergenic spacer analysis (RISA). Numbers of both types of bacteria were higher on dorsal than on ventral feathers. Numbers of free‐living, but not attached, bacteria on the two body regions were highly positively correlated. There was also a strong within‐individual correlation between numbers of attached bacteria during the same breeding stages in different years. These results suggest that, despite variation in absolute levels of feather bacterial loads between years and different body regions, sampling individual birds can provide reliable estimates of relative levels of bacterial contamination, as long as sampling time and body region are carefully standardized.     

Sexually dimorphic melanin‐based colour polymorphism,feather melanin content,and wing feather structure in the barn owl (Tyto alba)

Feathers confer protection against biophysical agents and determine flying ability. The geometry and arrangement of the barbs, together with the keratin and pigments deposited in the feathers, determine the mechanical stability of the vane, and its stiffness and resistance to abrasive agents. In colour‐polymorphic species, individuals display alternative colour morphs, which can be associated with different foraging strategies. Each morph may therefore require specific flying abilities, and their feathers may be exposed to different abrasive agents. Feathers of differently coloured individuals may thus have a specific structure, and colour pigments may help resist abrasive agents and improve stiffness. We examined these predictions in the barn owl (Tyto alba), a species for which the ventral body side varies from white to dark reddish pheomelanic, and in the number and size of black spots located at the tip of the feathers. White and reddish birds show different foraging strategies, and the size of black feather spots is associated with several phenotypic attributes. We found that birds displaying a darker reddish coloration on the ventral body side deposit more melanin pigments in their remiges, which also have fewer barbs. This suggests that wear resistance increases with darkness, whereas feathers of lighter coloured birds may bend less easily. Accordingly, individuals displaying a lighter reddish coloration on the ventral body side, and those displaying larger black spots, displayed more black transverse bars on their remiges: as larger‐spotted individuals are heavier and longer‐winged birds also have more transverse bars, these bars may reduce feather bending when flying. We conclude that differently coloured individuals produce wing feathers of different strengths to adopt alternative behavioural and life history strategies. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 109 , 562–573.     

Frequency and consequences of feather holes in Barn Swallows Hirundo rustica

The relationship between feather quality, estimated through the prevalence and intensity of feather holes, and the breeding performance and survival of the Barn Swallow Hirundo rustica was studied over a 5-year period. In addition, we present some indirect data on the role of chewing lice in producing the feather holes. The balanced distribution of feather holes (high prevalence, low aggregation) corresponds with the pattern of distribution of lice on colonial birds, and in the Barn Swallow. Feather holes were significantly and positively associated with the arrival dates of the birds, as both males and females with an increased number of feather holes started laying later. Females that survived the winter had significantly fewer holes than non-surviving females, whereas there was no difference in feather hole number between surviving and non-surviving males. Given that there was no association between the number of feather holes and body condition indices, except for the tail length of male Barn Swallows, we suggest that the negative effect of feather holes on the fitness of the birds is mainly apparent during periods of intensive locomotor activity, such as migration. Alternatively, if feather holes are an indicator of quality, those birds with a high intensity of feather holes may have been of poor quality. These birds may have been less able to cope with the environmental conditions, resulting in the lower survival and later arrival of the birds to the breeding grounds. The negative relationship between the length of the outermost tail feathers of males and the incidence of feather holes suggests that the tail is a condition-dependent secondary sexual characteristic.     

On the identification of feather structures in stem-line representatives of birds: evidence from fossils and actuopalaeontology

Dinosaurs with fossilized filamentous integument structures are usually preserved in a highly flattened state. Several different feather types have been described on this basis, but the two-dimensional preservation of specimens during fossilization makes the identification of single feather structures difficult due to overlapping feather structures in vivo. Morphological comparison with the diversity of recent feather types is therefore absolutely vital to avoid misinterpretation. To simulate the preservation process, a cadaver of recent Carduelis spinus (European siskin) was flattened in a printing press. Afterwards, the structure of the plumage was compared with the morphology of a single body feather from the same specimen. In comparison with the single feather, the body plumage of the flattened bird looked rather filamentous. It was almost impossible to identify single structures, and in their place, various artefacts were produced. The investigation of plumage in a specimen of the Mesozoic bird Confuciusornis sanctus reveals similar structures. This indicates that flattening of specimens during fossilization amplifies the effect of overlapping among feathers and also causes a loss of morphological detail which can lead to misinterpretations. The results are discussed in connection with some dubious feather morphologies in recently described theropods and basal birds. Based on recent feather morphology, the structure of so-called proximal ribbon-like pennaceous feathers (PRPFs) found in many basal birds is reinterpreted. Furthermore, the morphology of a very similar-looking feather type found in the forelimb and tail of an early juvenile oviraptorosaur is discussed and diagnosed as the first feather generation growing out of the feather sheath. Thus, the whole plumage of this theropod might represent neoptile plumage.     

Losing the last feather: feather loss as an antipredator adaptation in birds

Birds often lose feathers during predation attempts, and thisability has evolved as a means of escape. Because predatorsare more likely to grab feathers on the rump and the back thanon the ventral side of an escaping bird, we predicted that theformer feathers would have evolved to be relatively looselyattached as an antipredator strategy in species that frequentlydie from predation. We estimated the force required to removefeathers from the rump, back, and breast by pulling featherswith a spring balance from a range of European bird speciesin an attempt to investigate ecological factors associated withease of feather loss during predation attempts. The force requiredto loosen a feather from the rump was less than that requiredto loosen a feather from back, which in turn was less than thatrequired to loosen a feather from the breast. The relative forceneeded to loosen rump feathers compared with feathers from theback and the breast was smaller for prey species preferred bythe most common predator of small passerine birds, the sparrowhawkAccipiter nisus. Likewise, the relative force was also smallerin species with a high frequency of complete tail loss amongfree-living birds, which we used as an index of the frequencyof failed predation attempts. The relative force required toremove feathers from the rump was smaller in species with ahigh frequency of fear screams, another measure of the relativeimportance of predation as a cause of death. Feather loss requiredparticularly little force among solitarily breeding bird speciesthat suffer the highest degree of predation. Antipredator defensein terms of force required to remove feathers from the rumpwas larger in species with a strong antiparasite defense interms of T-cell–mediated immune response. These findingsare consistent with the hypothesis that different defenses areantagonistic and that they are traded off against each other.     

石油污染对鸟类的影响及石油污染清洁修复技术的研究进展

石油污染物通过自然泄漏和各种人类活动进入生态环境,对鸟类的生存产生影响。本文综述了国内外有关石油污染物对鸟类所产生的影响、受污染鸟类及其栖息地的清洁修复技术的研究现状以及发展趋势。现有研究发现,石油的毒性和石油对羽毛结构与功能的破坏是石油污染影响鸟类的主要原因。石油污染不仅对鸟类有直接的致死作用,而且能产生许多慢性危害,包括引起鸟类溶血性贫血、使其免疫能力下降、降低羽毛的保温和防水能力等。对鸟类栖息地中的石油污染进行及时清理并开展对受污染鸟类的清洁修复工作,是减轻石油污染对鸟类影响的重要手段。我国作为石油消费大国,关于石油污染对鸟类影响的研究与国外相比滞后,亟需在石油污染对鸟类的短期和长期影响、受污染鸟类及其栖息地的清洁修复技术等方面开展更多的研究。     

Follicle characteristics and follicle developmental related Wnt6 polymorphism in Chinese indigenous Wanxi-white goose

In birds, downy feather quantity mainly affected by the follicles. Wnt6, a secreted cysteine-rich protein, plays a key role in follicular development as an intercellular signaling molecule. The present study was to investigate the follicle development and Wnt6 polymorphism in Wanxi-white geese, a Chinese indigenous breed. In total, 300 fertilized eggs were hatched. At embryonic stage and on early birth goslings, the diameter and density of follicles from different sites were examined after sectioning and staining. The results showed that the diameter of primary feather follicles in thorax, venter, dorsum and flank had no difference at embryonic stage. In contrast, after birth, thorax and ventral feather follicles had greater diameter than those on dorsum and flank. Similarly, the primary feather follicle density was higher in thorax and venter than in dorsum and flank at embryonic stage. The secondary feather follicle diameter in flank was greater than that in other sites examined. The secondary follicle showed lush growth in E27 with thickest in ventral and thorax. Overall, follicle formed consistently in dorsal and flank, and follicle in thorax and ventral formed in another consistent way. The polymorphism study showed 2 single nucleotide polymorphisms of Wnt6 and 3 genotypes identified. Sequencing revealed two nucleotide transitions, T451C and A466G, which were synonymous mutations causing codons for aspartate and lysine to change from GAU to GAC and from AAA to AAG, respectively. This information about follicle development and Wnt6 polymorphisms would provide potential utilization in marker-assisted selection program for down feather selection.     

Comparative evidence for costs of secondary sexual characters: adaptive vane emargination of ornamented feathers in birds

We used a comparative approach, by comparing bird species with tail ornamentation with sister taxa without ornamentation, to deduce the aerodynamic function of extravagant feather ornaments and the costs of such ornaments in birds. First, the aerodynamic function of tail feather ornaments in birds can be deduced from asymmetry in the width of tail feather vanes, since flightless birds have symmetrical vanes while flying birds without feather exaggeration by sexual selection have asymmetrical vanes. Distal inner vanes at the tip of tail feathers were more narrow in ornamented as compared to nonornamented birds, and vane asymmetry at the tip of the feather was therefore reduced in ornamented species, suggesting marginal aerodynamic function of the distal part of extravagant feather ornaments. Second, the cost of feather ornaments due to parasite drag is proportional to the area of feathers extending beyond the maximum continuous width of the tail, and aerodynamic costs of long tails could therefore be diminished by a reduction in feather width. Consistent with this prediction, the outermost tip of feather ornaments was narrower than the homologous character in nonornamented sister taxa, while the base of the feather had similar width in the two groups of birds. These results suggest that the costs of extravagant ornamentation have been diminished by a reduction in feather width, leading to a reduction in drag. Costs of feather ornaments, as demonstrated by their fine morphology, thus appear to have been extensive during the evolution of these characters.     

Fitness and feather wear in the Collared Flycatcher Ficedula albicollis

Individual variation in the degree of feather wear is potentially a useful marker of individual quality or fitness, but next to nothing is known about causes and fitness consequences of feather wear in birds. We studied the effects of sex, age, year and experimental manipulation of brood size on primary feather wear in Collared Flycatchers Ficedula albicollis , and related variation in degree of feather wear to differences in fitness (viz. recruitment, survival). At the end of the breeding period, females and young birds had more worn flight feathers than males and adult birds, and the sexual difference in the degree of feather wear was particularly pronounced in one of the two study years. Experimental reduction of brood size reduced the degree of primary feather wear, whereas experimental enlargement of brood size did not lead to increased feather wear. In both sexes, there was a clear tendency for very old (>5 years old) birds to have more worn feathers than middle aged birds. The individual differences in the degree of feather wear were not correlated with individual differences in recruitment rate of young, but survival probability to the next breeding season increased with increasing degree of feather wear.     

Migratory behaviour affects the trade-off between feather growth rate and feather quality in a passerine bird

Migratory birds have less time for moulting than sedentary birds, which may force them to produce their feathers faster at the expense of reducing feather quality. However, the effects of migration on the trade-off between moult speed and plumage quality remain to be studied in natural populations. We analysed the relationship between growth rate and quality of individual feathers, taking advantage of natural variation between migratory and sedentary populations of blackcaps Sylvia atricapilla . As predicted by life-history theory, individual blackcaps showed variable individual quality, which was revealed by positive correlations between feather growth rate and feather mass within populations. However, migrants grew up their feathers faster, producing lighter feathers than sedentary blackcaps. These results support the idea that feather growth rate and feather quality are traded against each other in blackcaps. Such a trade-off is apparently caused by different selection associated to migratory and sedentary life styles, which opens new insights into the diversification of moult patterns in birds.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 98–105.     

Relationship between feather pecking and ground pecking in laying hens and the effect of group size

The aim of this experiment was to study the relationship between feather pecking and ground pecking in laying hens and the effect of group size on feather pecking behaviour. Hisex White hens were kept in floor pens in group sizes of 15, 30, 60 and 120 birds, each with four replicates. Behavioural observations were performed at four different ages and focused on the number of feather pecks and aggressive pecks, both given and received. The part of the body pecked and the location of the bird was recorded as well as the number of pecks made to the floor, feeder and drinker.The results showed that most feather pecking activity occurred in the largest group size (120 birds) and there was some evidence of an increasing frequency of aggressive pecks with increasing group size. The parts of the body which were targets for feather pecking varied depending on the location of the bird giving the peck and the bird receiving it. When looking at the behaviour of individuals, birds doing a lot of feather pecking also showed more ground pecking.     

Feather mites (Acari: Astigmata) and body condition of their avian hosts: a large correlative study

Feather mites are arthropods that live on or in the feathers of birds, and are among the commonest avian ectosymbionts. However, the nature of the ecological interaction between feather mites and birds remains unclear, some studies reporting negative effects of feather mites on their hosts and others reporting positive or no effects. Here we use a large dataset comprising 20 189 measurements taken from 83 species of birds collected during 22 yr in 151 localities from seven countries in Europe and North Africa to explore the correlation between feather mite abundance and body condition of their hosts. We predicted that, if wing‐dwelling feather mites are parasites, a negative correlation with host body condition should be found, while a mutualistic interaction should yield positive correlation. Although negative relationships between feather mite abundance and host body condition were found in a few species of birds, the sign of the correlation was positive in most bird species (69%). The overall effect size was only slightly positive (r =0.066). The effect of feather mite abundance explained <10% of variance in body condition in most species (87%). Results suggest that feather mites are not parasites of birds, but rather that they hold a commensalistic relationship where feather mites may benefit from feeding on uropygial gland secretions of their hosts and birds do not seem to obtain a great benefit from the presence of feather mites.     

Body condition and feather molt of a migratory shorebird during the non‐breeding season

Migratory shorebirds have some of the highest fat loads among birds, especially species which migrate long distances. The upland sandpiper Bartramia longicauda makes long‐distance migrations twice a year, but variation in body condition or timing of feather molt during the non‐breeding season has not been studied. Molt is an important part of the annual cycle of migratory birds because feather condition determines flight performance during migration, and long‐distance movements are energetically costly. However, variation in body condition during molt has been poorly studied. The objective of our field study was to examine the timing and patterns of feather molt of a long distance migratory shorebird during the non‐breeding season and test for relationships with body size, fat depots, mass, and sex. Field work was conducted at four ranches in the Northern Campos of Uruguay (Paysandú and Salto Departments). We captured and marked 62 sandpipers in a 2‐month period (Nov–Jan) during four non‐breeding seasons (2008–2012). Sex was determined by genetic analyses of blood samples taken at capture. Molt was measured in captured birds using rank scores based on published standards. Body mass and tarsus length measurements showed female‐biased sexual size dimorphism with males smaller than females. Size‐corrected body mass (body condition) showed a U‐shaped relationship with the day of the season, indicating that birds arrived at non‐breeding grounds in relatively good condition. Arriving in good body condition at non‐breeding grounds is probably important because of the energetic demands due to physiological adjustments after migration and the costs of feather molt.     

Uropygial gland size,feather holes and moult performance in the House Sparrow Passer domesticus

Feather holes represent damage to the plumage of birds and are correlated with delayed moult. Uropygial gland size is negatively correlated with feather holes. Consequently, it was predicted that birds with smaller uropygial glands would have more feather holes, and that this would affect moult performance. I examined this prediction in the House Sparrow Passer domesticus. Individuals with smaller uropygial glands had more feather holes, and those with more feather holes moulted later and faster. Therefore, uropygial gland size seemed to affect moult performance via its effect on feather holes. Uropygial gland size may have a positive effect on plumage quality, through a negative effect on feather holes, and therefore on moult timing and speed.     

Is the denser contour feather structure in pale grey than in pheomelanic brown tawny owls Strix aluco an adaptation to cold environments?

In colour polymorphic species morphs are considered to be adaptations to different environments, where they have evolved and are maintained because of their differential sensitivity to the environment. In cold environments the plumage insulation capacity is essential for survival and it has been proposed that plumage colour is associated with feather structure and thereby the insulation capacity of the plumage. We studied the structure of contour feathers in the colour polymorphic tawny owl Strix aluco. A previous study of tawny owls in the same population has found strong selection against the brown morph in cold and snowy winters whereas this selection pressure is absent in mild winters. We predicted that grey morphs have a denser and more insulative plumage, enabling them to survive better in cold climate compared to brown ones. The insulative plumulaceous part of the dorsal contour feathers was larger and the fine structure of the plumulaceous part of the feather was denser in grey tawny owls than in brown ones. In the ventral contour feathers the plumulaceous part of the feather was denser in females than in males and in older birds without any differences between morphs. Our study suggests that insulative microscopical feather structures differ between colour morphs and we propose that feather structure may be a trait associated with morph‐specific survival in cold environments.     

The scaling of the size and stiffness of primary flight feathers

Birds encompass a large range of body sizes, yet the importance of body size on feather morphology and mechanical properties has not been characterized. In this study, I examined the scaling relationships of primary flight feathers within a phylogenetically diverse sample of avian species varying in body size by nearly three orders of magnitude. I measured the scaling relationships between body mass and feather linear dimensions as well as feather flexural stiffness. The resnlts of an independent contrasts analysis to test the effects of phylogenetic history on the characters measured had no effect on the scaling relationships observed. There was slight, but not significant, positive allometry in the scaling of shaft diameter with respect to feather length across a range of body masses. The scaling of feather length and diameter against body mass was not significantly different from isometry. Flexural stiffness, however, exhibited strong negative allometry. Therefore, larger birds have relatively more flexible feathers than smaller birds. The more flexible primary feathers of large birds may reduce stresses on the wing skeleton during take-off and landing and also make these feathers less susceptible to mechanical failure. Conversely, the greater flexibility of these feathers may also reduce their capacity to generate aerodynamic lift.     

THE CONTAMINATION OF BIRDS' FEATHERS BY FUNGI

Fungi have regularly been recorded on feathers removed from birds. Cellulose-decomposing spores which are merely carried on the feathers were common on birds sampled in Britain, but were fewer than expected on the basis of random distribution on birds sampled in India. Hyaline-spored fungi were less common on Indian birds, and more abundant on British birds than expected. Keratinophilic fungi, which are related to the dermatophytes, were particularly associated with related groups of birds: thus Turdus spp. regularly yielded Arthroderma curreyi, and partridges were contaminated with Ctenomyces serratus. Chrysosporium spp. were widespread on birds in India and Britain. Where associations have been shown, feather fats from these birds stimulate, or do not depress the growth of the keratinophilic fungi. Otherwise, feather fats were inhibitory to the fungi. The temperature relationships of the fungi, showing no growth at 40°C, would explain the restriction of the keratinophilic fungi to the outer, contour feathers, and of dermatophytes mainly to the non-insulated parts of the birds' surface.     

Fine‐tuned distribution of feather mites (Astigmata) on the wing of birds: the case of blackcaps Sylvia atricapilla

The distribution of feather mites (Astigmata) along the wing of passerine birds could change dramatically within minutes because of the rapid movement of mites between feathers. However, no rigorous study has answered how fine‐tuned is the pattern of distribution of feather mites at a given time. Here we present a multiscale study of the distribution of feather mites on the wing of non‐moulting blackcaps Sylvia atricapilla in a short time period and at a single locality. We found that the number and distribution of mites differed among birds, but it was extremely similar between the wings of each bird. Moreover, mites consistently avoided the first secondary feather, despite that it is placed at the centre of the feathers most used by them. Thus, our results suggest that feather mites do precise, feather‐level decisions on where to live, contradicting the current view that mites perform “mass”, or “blind” movements across wing feathers. Moreover, our findings indicate that “rare” distributions are not spurious data or sampling errors, but each distribution of mites on the wing of each bird is the outcome of the particular conditions operating on each ambient‐bird‐feather mite system at a given time. This study indicates that we need to focus on the distribution of feather mites at the level of the individual bird and at the feather level to improve our understanding of the spatial ecology of mites on the wings of birds.     

Barb geometry of asymmetrical feathers reveals a transitional morphology in the evolution of avian flight

The geometry of feather barbs (barb length and barb angle) determines feather vane asymmetry and vane rigidity, which are both critical to a feather''s aerodynamic performance. Here, we describe the relationship between barb geometry and aerodynamic function across the evolutionary history of asymmetrical flight feathers, from Mesozoic taxa outside of modern avian diversity (Microraptor, Archaeopteryx, Sapeornis, Confuciusornis and the enantiornithine Eopengornis) to an extensive sample of modern birds. Contrary to previous assumptions, we find that barb angle is not related to vane-width asymmetry; instead barb angle varies with vane function, whereas barb length variation determines vane asymmetry. We demonstrate that barb geometry significantly differs among functionally distinct portions of flight feather vanes, and that cutting-edge leading vanes occupy a distinct region of morphospace characterized by small barb angles. This cutting-edge vane morphology is ubiquitous across a phylogenetically and functionally diverse sample of modern birds and Mesozoic stem birds, revealing a fundamental aerodynamic adaptation that has persisted from the Late Jurassic. However, in Mesozoic taxa stemward of Ornithurae and Enantiornithes, trailing vane barb geometry is distinctly different from that of modern birds. In both modern birds and enantiornithines, trailing vanes have larger barb angles than in comparatively stemward taxa like Archaeopteryx, which exhibit small trailing vane barb angles. This discovery reveals a previously unrecognized evolutionary transition in flight feather morphology, which has important implications for the flight capacity of early feathered theropods such as Archaeopteryx and Microraptor. Our findings suggest that the fully modern avian flight feather, and possibly a modern capacity for powered flight, evolved crownward of Confuciusornis, long after the origin of asymmetrical flight feathers, and much later than previously recognized.     

Variations in the morphology,distribution, and arrangement of feathers in domesticated birds

Domesticated birds exhibit a greater diversity in the morphology of their integument and its appendages than their wild ancestors. Many of these variations affect the appearance of a bird significantly and have been bred selectively by poultry and pigeon fanciers and aviculturists for the sake of visual appeal. Variations in feather distribution (e.g., feathering of legs and feet, featherless areas in normally feather-bearing skin) are widespread in chickens and pigeons. Variations in the number of feathers (e.g., increased number of tail feathers, lack of tail feathers) occur in certain pigeon and poultry breeds. Variations in feather length can affect certain body regions or the entire plumage. Variations in feather structure (e.g., silkiness, frilled feathering) can be found in exhibition poultry as well as in pet birds. Variations in feather arrangement (e.g., feather crests and vortices) occur in many domesticated bird species as a results of mutation and intense selective breeding. The causes of variations in the structure, distribution, length and arrangement of feathers is often unknown and opens a wide field for scientific research under various points of view (e.g., morphogenesis, pathogenesis, ethology, etc.). To that extent, variations in the morphology, distribution and arrangement of feathers in domesticated birds require also a concern for animal welfare because certain alleles responsible for integumentary variations in domesticated birds have pleiotropic effects, which often affect normal behaviour and viability.