The function and evolution of the tail streamer in hirundines

The morphology of a bird's tail may result from compromisesbetween aerodynamic efficiency, phylogenetic constraints andselection for non-aerodynamic characteristics, such as mateattraction. A good example of a trait shaped by trade-offsbetween aerodynamic efficiency and reproductive benefits mediatedthrough female preference is the tail streamer of the barn swallow. Here we use a standardized task to measure the impactof manipulated tail streamer lengths on maneuvering flightin the barn swallow and in the sand martin, a closely relatedspecies that lacks a streamer. Our results show that the tailstreamer of the barn swallow has a role in maneuvering flight. However, the outer tail feather is approximately 12 mm (9-20%)longer than the aerodynamic optimum for maneuvering flight.Furthermore, we show that the addition of artificial tail streamersto the sand martin, enhances maneuverability even at smallstreamer lengths, thereby implying that tail streamers mayhave evolved via natural selection for increased flight performance.Our results therefore suggest that initial tail streamer elongationin the barn swallow has a functional explanation in terms of increased aerodynamic performance. However, female choice hasbecome associated with this trait, promoting the developmentof a costly handicap.     

A comparative study of aerodynamic function and flexural stiffness of outer tail feathers in birds

To transmit aerodynamic forces to the body, tail feathers should be stiff to resist lift forces with minimum deformation. Because aerodynamic theory predicts that such feathers do not produce lift forces beyond the point of the maximum continuum width of the tail, species with deeply forked tails should not require stiff outer rectrices distal to that point. I tested this prediction by comparing the relative thickness of the outer rectrix rachis between species with deeply forked tails to those with triangular or shallowly forked tails. Eleven pairs of closely related species belonging to families Fregatidae, Phalacrocoracidae, Accipitridae, Sternidae, Caprimulgidae, Trochilidae, Coraciidae, Tyrannidae, Cotingidae, and Hirundinidae were compared. All but one of the phylogenetically independent comparisons showed that the species with triangular or shallowly forked tails have higher relative rachis thickness than their deeply forked relatives. In addition, nine out of eleven of the species with deeply forked tails showed a proportionately greater increase in relative rachis thickness from distal to proximal parts of the feather. In contrast, triangular and shallowly forked tails showed an approximately linear relation between relative rachis thickness and relative rachis length. These results considered together are consistent with the idea that the distal part of outer rectrix rachis in species with deeply forked tails has not been selected to resist lift forces and may be adaptively reduced to attenuate the costs of a hypertrophied ornament.     

Egg size decreases with increasing female tail fork depth in family Hirundinidae

Female ornamentation evolves as a nonfunctional copy of functional male ornamentation or through direct selection on female ornamentation. Because females invest many resources in both reproduction and ornamentation, an evolutionary tradeoff between female ornamentation and maternal investment can be predicted, but it has rarely been demonstrated. Using phylogenetic comparative analysis, we studied egg size in relation to female fork depth in the family Hirundinidae, which has a wide range of fork depth (from 0 to >50 mm). Because deeply forked tails are aerodynamically costly, we predicted that species in which females have deeply forked tails suffer reduced egg size due to inefficient foraging during the egg formation period. Egg length was significantly related to nest type but not to female or male tail fork depth, whereas we found a significant negative relationship between egg breadth and female, but not male, tail fork depth. As a consequence, a significant negative relationship was found between egg volume and female, but not male, tail fork depth. Because female and male fork depths were not significantly related to clutch size, clutch size would not compensate the relationship between egg size and fork depth. The current finding supports the hypothesis that female ornamentation trades off with maternal investment.     

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.     

Coevolution of caudal skeleton and tail feathers in birds

Birds are capable of a wide range of aerial locomotor behaviors in part because of the derived structure and function of the avian tail. The tail apparatus consists of a several mobile (free) caudal vertebrae, a terminal skeletal element (the pygostyle), and an articulated fan of tail feathers that may be spread or folded, as well as muscular and fibroadipose structures that facilitate tail movements. Morphological variation in both the tail fan and the caudal skeleton that supports it are well documented. The structure of the tail feathers and the pygostyle each evolve in response to functional demands of differing locomotor behaviors. Here, I test whether the integument and skeleton coevolve in this important locomotor module. I quantified feather and skeletal morphology in a diverse sample of waterbirds and shorebirds using a combination of linear and geometric morphometrics. Covariation between tail fan shape and skeletal morphology was then tested using phylogenetic comparative methods. Pygostyle shape is found to be a good predictor of tail fan shape (e.g., forked, graduated), supporting the hypothesis that the tail fan and the tail skeleton have coevolved. This statistical relationship is used to reconstruct feather morphology in an exemplar fossil waterbird, Limnofregata azygosternon. Based on pygostyle morphology, this taxon is likely to have exhibited a forked tail fan similar to that of its extant sister clade Fregata, despite differing in inferred ecology and other aspects of skeletal anatomy. These methods may be useful in reconstructing rectricial morphology in other extinct birds and thus assist in characterizing the evolution of flight control surfaces in birds. J. Morphol. 275:1431–1440, 2014. © 2014 Wiley Periodicals, Inc.     

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.     

The evolution of sexually dimorphic tail feathers is not associated with tail skeleton dimorphism

Sexual selection can influence the evolution of sexually dimorphic exaggerated display structures. Herein, we explore whether such costly ornamental integumentary structures evolve independently or if they are correlated with phenotypic change in the associated skeletal system. In birds, elongate tail feathers have frequently evolved in males and are beneficial as intraspecific display structures but impart a locomotor/energetic cost. Using the sexually dimorphic tail feathers of several passeriform species as a model system, we test the hypothesis that taxa with sexually dimorphic tail feathers also exhibit sexual dimorphism in the caudal skeleton that supports the muscles and integument of the tail apparatus. Caudal skeletal morphology is quantified using both geometric morphometrics and linear morphometrics across four sexually dimorphic passeriform species and four closely related monomorphic species. Sexual dimorphism is assessed using permutational MANOVA. Sexual dimorphism in caudal skeletal morphology is found only in those taxa that exhibit active functional differences in tail use between males and females. Thus, dimorphism in tail feather length is not necessarily correlated with the evolution of caudal skeletal dimorphism. Sexual selection is sufficient to generate phenotypic divergence in integumentary display structures between the sexes, but these change are not reflected in the underlying caudal skeleton. This suggests that caudal feathers and bones evolve semi‐independently from one another and evolve at different rates in response to different types of selective pressures.     

早白垩世鸟类尾综骨与尾羽之间的形态协同演化（英文）

从兽脚类恐龙中爬行类的骨质长尾,到以尾综骨为终端,并附着具有空气动力学功能的扇状尾羽的短巧尾部,是早期鸟类演化中最显著的适应性转变之一。但能直接反映这一转变的化石记录匮乏,而且对中生代鸟类尾部形态结构,以及尾综骨和尾羽早期演化的认知也相对不足。在此对早白垩世鸟类的尾综骨形态予以概述并将其与现生鸟类尾部结构类比。本研究强调了非鸟手盗龙类中尾椎的联合骨化与早期鸟类的尾综骨实属趋同演化。本研究表明,会鸟形类、孔子鸟形类、反鸟类和今鸟型类的尾综骨结构存在明显差异。今鸟型类尾综骨和尾羽(舵羽)与现代鸟类的相似,而相对更原始的鸟类的尾综骨,从形态来看,并不能支持舵羽球状膨大和必要的肌肉附着来操控具有空气动力学功能的扇状尾羽。由此可见,舵羽球状膨大、舵羽扇面与犁铧状的尾综骨是在今鸟型类演化早期相伴相生的。相对于从前的认知,本研究还发现孔子鸟类的尾综骨与反鸟类的有更多相似之处,与二者都具有的几乎相同的装饰性尾羽相符合。     

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.     

Development of fluctuating asymmetry in tail feathers of the barn swallow Hirundo rustica

Fluctuating asymmetry represents usually small, random deviations from symmetry in bilateral morphological characters. The ontogeny of asymmetry in morphological characters may reveal information about developmental processes in a general sense. I studied the development of fluctuating asymmetry in feather characters of the barn swallow Hirundo rustica, that are developed repeatedly during the single annual moult, with the following results. First, the side developing a larger feather was found to be partially biased, as demonstrated by one side consistently developing a larger feather under natural and experimentally induced growth episode events. Second, asymmetric feathers were found to consist of asymmetric daily growth increments, and the size of the increments developing under different environmental conditions were positively correlated. Third, fluctuating asymmetries of feathers developing under different environmental conditions were positively correlated, although the level of asymmetry was larger under adverse environmental conditions. Fourth, individual asymmetries in tail length and growth bar length were unrelated to the duration of the developmental period, although late growth increments were smaller and more symmetric than early increments. These observations suggest that fluctuating asymmetry partially arises as a consequence of a random bias in the feather follicles and differences in environmental conditions during ontogeny of feathers.     

Fluttering wing feathers produce the flight sounds of male streamertail hummingbirds

Sounds produced continuously during flight potentially play important roles in avian communication, but the mechanisms underlying these sounds have received little attention. Adult male Red-billed Streamertail hummingbirds (Trochilus polytmus) bear elongated tail streamers and produce a distinctive 'whirring' flight sound, whereas subadult males and females do not. The production of this sound, which is a pulsed tone with a mean frequency of 858 Hz, has been attributed to these distinctive tail streamers. However, tail-less streamertails can still produce the flight sound. Three lines of evidence implicate the wings instead. First, it is pulsed in synchrony with the 29 Hz wingbeat frequency. Second, a high-speed video showed that primary feather eight (P8) bends during each downstroke, creating a gap between P8 and primary feather nine (P9). Manipulating either P8 or P9 reduced the production of the flight sound. Third, laboratory experiments indicated that both P8 and P9 can produce tones over a range of 700-900 Hz. The wings therefore produce the distinctive flight sound, enabled via subtle morphological changes to the structure of P8 and P9.     

Population differences in density and resource allocation of ornamental tail feathers in the barn swallow

Many organisms show well‐defined latitudinal clines in morphology, which appear to be caused by spatially varying natural selection, resulting in different optimal phenotypes in each location. Such spatial variability raises an interesting question, with different prospects for the action of sexual selection on characters that have a dual purpose, such as locomotion and sexual attraction. The outermost tail feathers of barn swallows (Hirundo rustica) represent one such character, and their evolution has been a classic model subject to intense debate. In the present study, we examined individuals from four European populations to analyze geographical variation in the length and mass of tail feathers in relation to body size and wing size. Tail feather length differed between sexes and populations, and such variation was a result of the effects of natural selection, acting through differences in body size and wing size, as well as the effects of sexual selection that favours longer tails. The extra enlargement of the tail promoted by sexual selection (i.e. beyond the natural selection optimum) could be achieved by increasing investment in ornaments, and by modifying feather structure to produce longer feathers of lower density. These two separate processes accounting for the production of longer and more costly tail feathers and less dense feathers, respectively, are consistent with the hypothesis that both Zahavian and Fisherian mechanisms may be involved in the evolution of the long tails of male barn swallows. We hypothesize that the strength of sexual selection increases with latitude because of the need for rapid mating as a result of the short duration of the breeding season at high latitudes. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 925–936.     

Mechanical and structural adaptations to migration in the flight feathers of a Palaearctic passerine

Current avian migration patterns in temperate regions have been developed during the glacial retreat and subsequent colonization of the ice‐free areas during the Holocene. This process resulted in a geographic gradient of greater seasonality as latitude increased that favoured migration‐related morphological and physiological (co)adaptations. Most evidence of avian morphological adaptations to migration comes from the analysis of variation in the length and shape of the wings, but the existence of intra‐feather structural adjustments has been greatly overlooked despite their potential to be under natural selection. To shed some light on this question, we used data from European robins Erithacus rubecula overwintering in Campo de Gibraltar (Southern Iberia), where sedentary robins coexist during winter with conspecifics showing a broad range of breeding origins and, hence, migration distances. We explicitly explored how wing length and shape, as well as several functional (bending stiffness), developmental (feather growth rate) and structural (size and complexity of feather components) characteristics of flight feathers, varied in relation to migration distance, which was estimated from the hydrogen stable isotope ratios of the summer‐produced tail feathers. Our results revealed that migration distance not only favoured longer and more concave wings, but also promoted primaries with a thicker dorsoventral rachis and shorter barb lengths, which, in turn, conferred more bending stiffness to these feathers. We suggest that these intra‐feather structural adjustments could be an additional, largely unnoticed, adaptation within the avian migratory syndrome that might have the potential to evolve relatively quickly to facilitate the occupation of seasonal environments.     

A potential role for bat tail membranes in flight control

Wind tunnel tests conducted on a model based on the long-eared bat Plecotus auritus indicated that the positioning of the tail membrane (uropatagium) can significantly influence flight control. Adjusting tail position by increasing the angle of the legs ventrally relative to the body has a two-fold effect; increasing leg-induced wing camber (i.e., locally increased camber of the inner wing surface) and increasing the angle of attack of the tail membrane. We also used our model to examine the effects of flying with and without a tail membrane. For the bat model with a tail membrane increasing leg angle increased the lift, drag and pitching moment (nose-down) produced. However, removing the tail membrane significantly reduced the change in pitching moment with increasing leg angle, but it had no significant effect on the level of lift produced. The drag on the model also significantly increased with the removal of the tail membrane. The tail membrane, therefore, is potentially important for controlling the level of pitching moment produced by bats and an aid to flight control, specifically improving agility and manoeuvrability. Although the tail of bats is different from that of birds, in that it is only divided from the wings by the legs, it nonetheless, may, in addition to its prey capturing function, fulfil a similar role in aiding flight control.     

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.     

Resistance of flight feathers to mechanical fatigue covaries with moult strategy in two warbler species

Flight feather moult in small passerines is realized in several ways. Some species moult once after breeding or once on their wintering grounds; others even moult twice. The adaptive significance of this diversity is still largely unknown. We compared the resistance to mechanical fatigue of flight feathers from the chiffchaff Phylloscopus collybita, a migratory species moulting once on its breeding grounds, with feathers from the willow warbler Phylloscopus trochilus, a migratory species moulting in both its breeding and wintering grounds. We found that flight feathers of willow warblers, which have a shaft with a comparatively large diameter, become fatigued much faster than feathers of chiffchaffs under an artificial cyclic bending regime. We propose that willow warblers may strengthen their flight feathers by increasing the diameter of the shaft, which may lead to a more rapid accumulation of damage in willow warblers than in chiffchaffs.     

Spina cortica and Tapetum spinosus, two new microstructures of flight feathers: description, function and distribution in modern birds

The importance of feathers for the avian group has made them one of the most studied epidermal structures both from the morphological and evolutionary point of view. Surprisingly, our observations by Scanning Electron Microscopy detected the presence of two structures widely distributed within different avian groups and not yet described. In this paper we describe these two new structures (Spina cortica and Tapetum spinosus) and map their distribution within modern birds. The S. cortica is a thorn-like microstructure that grows on the barb cortex and the T. spinosus is the assemblage of these thorns. The distribution of these new structures among birds and their morphological diversity could be of great interest to taxonomists and evolutionary biologists interested in the origin of bird flight.     

The growth bars on tail feathers in the male Styan's Grasshopper Warbler may indicate quality

We propose that growth bar size may indicate the quality of individuals in Styan's Grasshopper Warblers Locustella pleskei . Positive correlations of an individual's standardized growth bar width between years showed that some individuals of both sexes were usually in better condition than others. The survival rate of males with wider growth bars was higher than that of other males. These survivors might have survived better because of a better body condition. We also examined the relationships between growth bar width and reproductive parameters (arrival date, pairing date and reproductive success). Males with wider growth bars arrived earlier, acquired a mate earlier and achieved higher total reproductive success by breeding twice. In females, growth bar width was correlated with reproductive parameters (arrival date, pairing date and total reproductive success), but not with survival. These results suggest that males with wider growth bars were of higher quality than those with narrower bars. The contrasting results for the two sexes may be explained by differences in their breeding tactics.